"Science, Faculty of"@en . "Zoology, Department of"@en . "DSpace"@en . "UBCV"@en . "McPhail, John Donald"@en . "2012-01-12T20:42:51Z"@en . "1959"@en . "Master of Science - MSc"@en . "University of British Columbia"@en . "Sympatric populations of S. malma and S. alpinus from Alaska were compared using a discriminant function analysis. The comparison indicated little or no hybridization. S. malma is, therefore, regarded as a distinct species. S. malma has fewer gill rakers, pyloric caeca and pores along the lateral line than S. alpinus. Five hundred specimens of S. malma from 42 North American localities were examined. Considerable geographic variability was observed. The variation showed no correlation with latitude. Evidence is presented that subspecific distinctions\r\nare invalid in S. malma. It is suggested that S. malma evolved in the North Pacific area sometime during the Pliestocene."@en . "https://circle.library.ubc.ca/rest/handle/2429/40050?expand=metadata"@en . "A SYSTEMATIC STUDY OF THE DOLLY VARDEN, SALVELINUS MALMA (WALBAUM) by JOHN DONALD McPHAIL B.A., U n i v e r s i t y o f B r i t i s h C o l u m b i a , 1957 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE o i n t h e D e p a r t m e n t o f ZOOLOGY We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o t h e s t a n d a r d r e q u i r e d f r o m c a n d i d a t e s f o r t h e d e g r e e o f MASTER OF SCIENCE Members o f t h e Dep a r t m e n t o f Z o o l o g y THE UNIVERSITY OF BRITISH COLUMBIA May, 1959 ABSTRACT S y m p a t r i c p o p u l a t i o n s o f S. malma and S. a l p i n u s f r o m A l a s k a were compared u s i n g a d i s c r i m i n a n t f u n c t i o n a n a l y s i s . The c o m p a r i s o n i n d i c a t e d l i t t l e o r no h y b r i d i z a t i o n . S. malma i s , t h e r e f o r e , r e g a r d e d as a d i s t i n c t s p e c i e s . S. malma has f e w e r g i l l r a k e r s , p y l o r i c c a e c a and p o r e s a l o n g t h e l a t e r a l l i n e t h a n S_. a l p i n u s . F i v e h u n d r e d s p e c i m e n s o f S. malma f r o m 42 N o r t h A m e r i c a n l o c a l i t i e s were examined. C o n s i d e r a b l e g e o g r a p h i c v a r i a b i l i t y was o b s e r v e d . The v a r i a t i o n showed no c o r r e l a t i o n w i t h l a t i t u d e . E v i d e n c e i s p r e s e n t e d t h a t s u b s p e c i f i c d i s t i n c -t i o n s a r e i n v a l i d i n S. malma. I t i s s u g g e s t e d t h a t S. malma e v o l v e d i n t h e N o r t h P a c i f i c a r e a sometime d u r i n g t h e P l i e s t o c e n e . In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make i t freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of The University of British Columbia, Vancouver 8, Canada. i ACKNOWLEDGMENTS Dr. C. C. L i n d s e y s u g g e s t e d t h e p r o b l e m and d i r e c -t e d t h r o u g h o u t t h e i n v e s t i g a t i o n . H i s g u i d a n c e and a d v i c e a r e g r a t e f u l l y a c k n o w l e d g e d . D. R. M i l l e r a s s i s t e d w i t h most o f t h e f i e l d c o l l e c t i o n s . Mr. T. Ueno p r o v i d e d t r a n s l a t i o n s o f J a p a n e s e l i t -e r a t u r e . M i s s M. J u r k e l a p r o v i d e d t r a n s l a t i o n s o f R u s s i a n l i t e r a t u r e . The k i n d a s s i s t a n c e o f t h e s t a f f a t t h e U.B.C. c o m p u t i n g c e n t e r i s a l s o a c k n o w l e d g e d . F i n a n c i a l s u p p o r t f o r t h e s t u d y was p r o v i d e d by t h e A r c t i c I n s t i t u t e o f N o r t h A m e r i c a and t h e I n s t i t u t e o f F i s h e r i e s , U.B.C. i i TABLE OF CONTENTS Page I . I n t r o d u c t i o n 1 I I . M a t e r i a l s and Methods 4 S p e c i m e n s examined 4 S t a i n i n g p r o c e d u r e 4 Measurements 5 C o u n t s 5 C o l o u r p a t t e r n . . . . . # T r e a t m e n t o f D a t a \u00C2\u00A3 I I I . R e s u l t s 9 Taxonomic R a l a t i o n s h i p o f S. malma and S. a l p i n u s 9 K a r l u k L a k e 9 F r a s e r L a k e 27 B r o o k s L a k e 3& K e n a i R i v e r 42 Summary o f t a x o n o m i c r e l a t i o n s h i p o f S. malma and S. a l p i n u s 43 G e o g r a p h i c v a r i a b i l i t y 43 D e s c r i p t i o n o f S. malma 52 Key t o t h e a d u l t c h a r o f w e s t e r n N o r t h A m e r i c a . . 54 D i s t r i b u t i o n o f S. malma 56 IV. D i s c u s s i o n 59 V. Summary 65 B i b l i o g r a p h y 67 A p p e n d i x I 72 A p p e n d i x I I 73 i i i L I S T OF TABLES TABLE PAGE I C o m p a r i s o n o f M e r i s t i c d a t a f o r S. malma and S. a l p i n u s i n K a r l u k L a k e ( d a t a f r o m D e l a c y and M o r t o n 1942). 10 I I C o r r e l a t i o n o f g i l l r a k e r s , p y l o r i c c a e c a and p o r e s a l o n g t h e l a t e r a l l i n e w i t h f o r k l e n g t h f o r S. malma and S. a l p i n u s i n K a r l u k L a k e . 22 I I I A c o m p a r i s o n o f t h e m a j o r t o x o n o m i c c h a r a c -t e r s o f S. malma and S. a l p i n u s . 54 iv LIST OF FIGURES FIGURE PAGE 1. Hap showing world distribution of Salvelinus. 2 2. Measurements used in the present study. 6 3. Distribution of g i l l rakers in the complete Karluk Lake sample. 12 4. Distribution of g i l l rakers and pyloric caeca in Karluk Lake Salvelinus. 14 5. Distribution of pores along the lateral line and spots in area B for Karluk Lake Salvelinus. 15 6. Distribution of tooth counts in Karluk Lake Salvelinus. 16 7. Correlations of g i l l rakers on fork length in Karluk Lake Salvelinus. 1$ 8. Correlations of pyloric caeca and pores along the lateral line on fork length in Karluk Lake Salvelinus. 19 9. Correlation of pyloric caeca on fork length for S. alpinus in Fraser Lake. 21 10. Discriminant scores for Karluk Lake and Fraser Lake. 25 11. Iterated discriminant scores for Karluk Lake 26 12. Distribution of g i l l rakers in the Fraser Lake sample of Salvelinus. 29 13. Distribution of g i l l rakers and pyloric caeca in Fraser Lake Salvelinus. 30 FIGURE PAGE 14. D i s t r i b u t i o n o f p o r e s a l o n g t h e l a t e r a l l i n e , s p o t s i n a r e a B and s i z e o f s p o t s i n F r a s e r L a k e S a l v e l i n u s . 31 15. D i s t r i b u t i o n o f t e e t h i n F r a s e r L a k e S a l v e l i n u s . 32 16. D i s t r i b u t i o n o f b r a n c h i o s t e g a l r a y s i n F r a s e r L a k e S a l v e l i n u s . 33 17. a) R e l a t i v e g r o w t h o f he a d i n F r a s e r L a k e S a l v e l i n u s . b) R e l a t i v e g r o w t h o f i n t e r o r b i t a l w i d t h i n F r a s e r L a k e S a l v e l i n u s . 35 I d . a) R e l a t i v e g r o w t h o f v e r t i c a l eye d i a m e t e r i n F r a s e r L a k e S a l v e l i n u s . b) R e l a t i v e g r o w t h o f c a u d a l p e d u n c l e i n F r a s e r L a k e S a l v e l i n u s . 36 19. R e l a t i v e g r o w t h o f m a x i l l a r y i n F r a s e r L a k e S a l v e l i n u s . 37 20. D i s t r i b u t i o n o f t a x o n o m i c c h a r a c t e r s e x amined on B r o o k s L a k e s a m p l e . 39 21. D i s c r i m i n a n t s c o r e s f o r combined K a r l u k , F r a s e r L a k e s , B r o o k s L a k e and K e n a i R i v e r . 41 22. G e o g r a p h i c v a r i a t i o n i n g i l l r a k e r s on t h e l o w e r l i m b o f S. malma. 44 23. G e o g r a p h i c v a r i a t i o n i n p y l o r i c c a e c a o f S. malma. 45 24. G e o g r a p h i c v a r i a t i o n i n t h e number o f p o r e s a l o n g t h e l a t e r a l l i n e o f S. malma. 46 FIGURE PAGE 25. G e o g r a p h i c v a r i a t i o n o f r e l a t i v e h e a d l e n g t h o f S. malma. 47 26. a) G e o g r a p h i c v a r i a t i o n o f t e e t h on t o n g u e o f S. malma. b) G e o g r a p h i c v a r i a t i o n i n v e r t e b r a e number i n S. malma. 4# 27. a) G e o g r a p h i c v a r i a t i o n i n mean g i l l r a k e r c o u n t . b) G e o g r a p h i c v a r i a t i o n i n mean p y l o r i c c a e c a c o u n t . 50 2o*. a) C o r r e l a t i o n o f mean g i l l r a k e r number and mean p y l o r i c c a e c a number, b) C o r r e l a t i o n o f g i l l r a k e r c o u n t and p y l o r i c c a e c a i n S e l t a t C r e e k . 51 29. Map i n d i c a t i n g N o r t h A m e r i c a n r a n g e o f S. malma. 561 1 I . INTRODUCTION The c h a r ( S a l v e l i n u s ) a r e a h o l a r c t i c genus o f S a l m o n i d f i s h e s . C h a r c l o s e l y r e s e m b l e t r o u t ( S a l m o ) , b u t a r e d i s t i n g u i s h e d by t h e p r e s e n c e o f l i g h t s p o t s a g a i n s t a d a r k b a c k g r o u n d and t h e a b s e n c e o f t e e t h on t h e s h a f t o f t h e vomer. M o r p h o l o g i c a l l y c h a r a r e v e r y p l a s t i c . S t u n t e d p o p u l a t i o n s and c o l o u r v a r i a n t s a r e common. As a r e s u l t numerous s p e c i e s o f S a l v e l i n u s have b e e n d e s c r i b e d , many o f w h i c h a r e p r o b a b l y synonymous. F i g u r e 1 p r e s e n t s t h e w o r l d d i s t r i b u t i o n o f S a l v e l i n u s . J o r d a n , Evermann and C l a r k (1930) l i s t t h i r t e e n s p e c i e s o f S a l v e l i n u s f r o m N o r t h A m e r i c a . T h e s e c h a r f a l l i n t o t h r e e d i s t i n c t g r o u p s ; S_. namaycush, S. f o n t i n a l i s , a n d t h e S. a l p i n u s g r o u p . The f i r s t two e a c h c o n s i s t o f a s i n g l e s p e c i e s endemic t o N o r t h A m e r i c a . The t h i r d g r o u p i s c i r c u m -p o l a r i n d i s t r i b u t i o n , and c o n t a i n s an u n d e t e r m i n e d number o f s p e c i e s . T h i s g r o u p i s r e f e r r e d t o as t h e A r c t i c c h a r s , o r t h e n S . a l p i n u s c omplex.\" B e r g (1943) r e c o g n i z e s e l e v e n s p e c i e s o f A r c t i c c h a r s f r o m R u s s i a . J o r d a n , Evermann and C l a r k (1930) l i s t f o u r s p e c i e s o f A r c t i c c h a r f r o m G r e e n l a n d and t h e C a n a d i a n a r c t i c . V l a d y k o v (1954) and W a l t e r s (1955) s u g g e s t t h e use o f t h e s i n g l e name S a l v e l i n u s a l p i n u s . f o r t h e w h o l e g r o u p u n t i l more m a t e r i a l i s a v a i l a b l e f o r s t u d y . 1. J o r d a n , Evermann and C l a r k (1930) c o n s i d e r t h e l a k e c h a r ( S a l v e l i n u s namaycush) a d i s t i n c t genus C r i s t i v o m e r . F o r a d i s c u s s i o n on t h e v a l i d i t y o f C r i s t i v o m e r s e e M o r t o n and M i l l e r (1954) and V l a d y k o v (195477 2 F i g . 1. Map showing world d i s t r i b u t i o n of S a l v e l i n u s . 3 A char closely related to the S. alpinus complex is found along the Pacific coasts of North America and Asia. This is the Dolly Varden, Salvelinus malma (Walbaum). Several authors (Dymond 1936, 1947, Carl and Clemens 1953, and Lindsey 1956) have considered the Dolly Varden a part of the S. alpinus complex and refer to i t by the subspecific name Salvelinus alpinus malma. Other authors (Delacy and Morton 1942, Morton and Miller 1954) consider the Dolly Varden a distinct species, Salvelinus malma. The present study is an attempt to cl a r i f y the systematic status of the Dolly Varden, and to determine the variabil i t y within the species. New material was collected during the summer of 195S, including specimens from areas where both S. malma and S. alpinus had been reported. The 195$ collections, mostly made in Alaska, were sponsered by the Arctic Institute of North America and the Institute of Fisheries, U.B.C. 4 II. MATERIALS AND METHODS Specimens E x a m i n e d The m a t e r i a l u s e d c o n s i s t e d o f c o l l e c t i o n s o f w i l d f i s h f r o m B r i t i s h C o l u m b i a , A l b e r t a , t h e Yukon T e r r i t o r y , A l a s k a and Montana. S p e c i m e n s f r o m 42 l o c a l i t i e s , t o t a l i n g 500 S. malma and 100 S. a l p i n u s , were examined. I n d i v i d u a l s r a n g e d f r o m 20 t o 500 mm. i n l e n g t h . B o t h anadromous and n o n -m i g r a t o r y p o p u l a t i o n s o f S. malma were s a m p l e d . The c o l l e c -t i o n s a r e d e p o s i t e d 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 Museum, U n i v e r s i t y o f B r i t i s h C o l u m b i a and t h e F i s h e r i e s Museum, U n i v e r s i t y o f W a s h i n g t o n . A p p e n d i x I l i s t s t h e l o c a l i t y and c o l l e c t i o n numbers f o r e a c h c o l l e c t i o n o f S. malma examined. The number i n f r o n t o f e a c h c o l l e c t i o n c o r r e s p o n d s t o t h e number on t h e map ( F i g u r e 29). S t a i n i n g P r o c e d u r e S e r i e s o f f i s h were s t a i n e d and c l e a r e d u s i n g a t e c h n i q u e s i m i l a r t o t h a t d e s c r i b e d b y H o l l i s t e r (1934). No whole f i s h o v e r 130 mm. l o n g was s t a i n e d and c l e a r e d , b u t t h e l o w e r jaw and b r a n c h i a l a r c h e s o f some l a r g e r f i s h were removed and s t a i n e d . 5 M o r p h o m e t r i c Measurements Body p a r t s were m e a s u r e d u s i n g a d i a l c a l i p e r a c c u r a t e t o 0.1 ram. L e n g t h was m e a s u r e d on a m e a s u r i n g b o a r d . The f o l l o w i n g measurements, i l l u s t r a t e d i n F i g u r e 2, were made: 1. F o r k l e n g t h - d i s t a n c e f r o m t h e t i p o f t h e s n o u t t o t h e end o f t h e m i d d l e c a u d a l r a y s . 2. Head l e n g t h - t i p o f s n o u t t o t h e most p o s t e r i o r edge o f t h e o p e r c u l a r membrane. 3. M a x i l l a r y l e n g t h - t i p o f s n o u t t o p o s t e r i o r t i p o f m a x i l l a r y . 4. Eye d i a m e t e r - d o r s o - v e n t r a l d i s t a n c e between t h e o r b i t a l r i m s . 5. C a u d a l p e d u n c l e d e p t h - l e a s t d e p t h o f c a u d a l p e d u n c l e . 6. I n t e r o r b i t a l w i d t h - minimum d i s t a n c e between t h e t o p m a r g i n s o f t h e o r b i t s . M e r i s t i c C o u n t s C o u n t s o f f i n r a y s , v e r t e b r a e , b r a n c h i o s t e g a l r a y s and t e e t h were made u n d e r a b i n o c u l a r m i c r o s c o p e , on c l e a r e d s p e c i m e n s . G i l l r a k e r s , p y l o r i c c a e c a and p o r e s a l o n g t h e l a t e r a l l i n e were c o u n t e d on f i s h o f a l l s i z e s . R 6 - i A 1 \u00E2\u0080\u0094 \u00E2\u0080\u0094 F i g . 2. Measurements used i n the present study. 1. Fork length 2. Head length 3. M a x i l l a r y length 4. Eye diameter 5. Caudal peduncle depth 6. I n t e r o r b i t a l width A. Area i n which pores were counted B. Area i n which spots were counted 7 1. F i n r a y s - I n t h e d o r s a l and a n a l f i n s a l l r a y s were c o u n t e d . C o u n t s i n c l u d e d t h e s h o r t , r u d i m e n t a r y r a y s a t t h e a n t e r i o r o f t h e d o r s a l and a n a l f i n s . The p o s t e r i o r s p l i t r a y o f t h e d o r s a l and a n a l f i n was c o u n t e d a s one r a y . 2. V e r t e b r a e - A l l v e r t e b r a e were c o u n t e d , i n c l u d i n g t h e t h r e e c o m p r i s i n g t h e h y p u r a l . 3. B r a n c h i o s t e g a l r a y s - B r a n c h i o s t e g a l r a y s were c o u n t e d o n l y on s t a i n e d f i s h . A l l b r a n c h i o s t e g a l r a y s were c o u n t e d . 4. G i l l r a k e r s - A l l g i l l r a k e r c o u n t s were made on t h e l e f t , a n t e r i o r g i l l a r c h . R u d i m e n t a r y r a k e r s were i n c l u d e d . The a r c h was d i s s e c t e d t o f a c i l i t a t e c o u n t i n g . G i l l r a k e r s on t h e u p p e r and l o w e r l i m b s o f t h e a r c h were c o u n t e d s e p a r a t e l y . The r a k e r s t r a d d l i n g t h e a n g l e o f t h e a r c h was i n c l u d e d i n t h e c o u n t o f t h e l o w e r l i m b . A l l g i l l r a k e r c o u n t s were made u s i n g a b i n o c u l a r m i c r o s c o p e . 5. P y l o r i c c a e c a - P y l o r i c c a e c a were c o u n t e d i n d i v i d -u a l l y a s t h e y were s e p a r a t e d f r o m t h e m a i n mass o f c a e c a . E a c h t i p was c o u n t e d a s a n i n d i v i d u a l caecum. 6. Number o f p o r e s a l o n g t h e l a t e r a l l i n e - A l l p o r e c o u n t s were made u s i n g a b i n o c u l a r m i c r o s c o p e . O n l y t h e p o r e s p o s t e r i o r t o t h e p o i n t where t h e l a t e r a l l i n e t u r n s d o r s a l l y t o f o l l o w t h e c o n t o u r o f t h e o p e r c u l u m were c o u n t e d ( F i g u r e 2 ) . P o r e s i n s m a l l , a c c e s s o r y b r a n c h e s o f t h e l a t e r a l l i n e were n o t c o u n t e d . 7. T o o t h c o u n t s - T o o t h c o u n t s were made on t h e t o n g u e and b a s i b r a n c h i a l bone. A l v e o l e s f r o m w h i c h t e e t h had b e e n l o s t were c o u n t e d a s t e e t h . 8 C o l o u r P a t t e r n 1. Number o f s p o t s - The number o f s p o t s i n t h e a r e a u n d e r t h e d o r s a l f i n and above t h e l a t e r a l l i n e were c o u n t e d ( F i g u r e 2, A r e a B ) . Any s p o t s p a r t i a l l y i n t h i s a r e a were i n c l u d e d . S p o t s were o n l y c o u n t e d on f i s h on w h i c h t h e number o f s p o t s c o u l d be a c c u r a t e l y d e t e r m i n e d . 2. S i z e o f s p o t s - The h o r i z o n t a l d i a m e t e r o f t h e l a r g e s t s p o t t o u c h i n g t h e l a t e r a l l i n e was u s e d as a measure o f s p o t s i z e . S p o t s i z e v a r i e s w i t h l e n g t h , and was t h e r e f o r e e x p r e s s e d as a p r o p o r t i o n o f t h e i n t e r o r b i t a l w i d t h . T r e a t m e n t o f D a t a A l l body measurements f o r F r a s e r L a k e c h a r were p l o t t e d a g a i n s t f o r k l e n g t h o r h e a d l e n g t h on l o g a r i t h m i c a x i s ( M a r t i n 1949). L i n e s o f b e s t f i t were f i t t e d b y e y e . S i g n i f i c a n c e o f c o r r e l a t i o n s was c r u d e l y t e s t e d b y C h i - s q u a r e a n a l y s i s o f t h e number o f i n d i v i d u a l s w h i c h com-b i n e d more t h a n o r l e s s t h a n t h e mean measurement o r c o u n t o f e a c h o f t h e v a r i a b l e s . A d i s c r i m i n a n t f u n c t i o n a n a l y s i s ( F i s h e r 1937, Rao 194$) was u s e d on d a t a f r o m f o u r A l a s k a n c o l l e c t i o n s o f c h a r . The c o m p u t a t i o n s were done a t t h e U.B.C. C o m p u t i n g C e n t e r on an Alwac H I E computor. The method i s d i s c u s s e d l a t e r . G e o g r a p h i c v a r i a b i l i t y o f d a t a was p r e s e n t e d g r a p h i c a l l y u s i n g Hubbs and Hubbs' (1953) m o d i f i c a t i o n o f t h e D i c e - L e r a a s t e c h n i q u e . 9 I I I . RESULTS Taxonomic R e l a t i o n s h i p o f S a l v e l i n u s malma (Walbaum) and S a l v e l i n u s a l p i n u s ( L i n n a e u s ) . To d e t e r m i n e w h e t h e r S. malma i s a d i s t i n c t s p e c i e s , o r a s u b s p e c i e s o f S. a l p i n u s , i t i s n e c e s s a r y t o examine s a m p l e s f r o m a r e a s where t h e two f o r m s o c c u r s y m p a t r i c a l l y . F o u r s u c h a r e a s were s a m p l e d . K a r l u k L a k e K a r l u k L a k e ( l o c a t i o n 9, F i g u r e 29) i s t h e l a r g e s t l a k e on K o d i a k I s l a n d . I t d r a i n s t o t h e s e a by t h e K a r l u k R i v e r . D e l a c y and M o r t o n (1942) r e p o r t e d two s p e c i e s o f S a l v e l i n u s f r o m K a r l u k L a k e . They a s s i g n e d one s p e c i e s t o t h e S. a l p i n u s complex and r e g a r d e d t h e o t h e r s p e c i e s as S. malma. B e c a u s e t h e two f o r m s l i v e s y m p a t r i c a l l y D e l a c y and M o r t o n c o n t e n d e d t h a t S. malma must be a v a l i d s p e c i e s . They p r e -s e n t e d a m o r p h o m e t r i c and m e r i s t i c c o m p a r i s i o n o f t h e two s p e c i e s . E x a m i n a t i o n o f t h e i r d a t a i n d i c a t e s t h a t t h e r e a r e no m a j o r m o r p h o m e t r i c d i f f e r e n c e s t h a t may be u s e d t o s e p a r a t e t h e two n o m i n a l s p e c i e s . T a b l e I p r e s e n t s t h e modes and r a n g e s o f D e l a c y and M o r t o n ' s m e r i s t i c d a t a . T a b l e I i n d i c a t e s c o n s i d e r a b l e o v e r l a p i n a l l o f t h e c h a r a c t e r s p r e s e n t e d . D e l a c y and M o r t o n n o t e d m a j o r b e h a v i o u r a l d i f f e r -e n c e s between t h e two K a r l u k L a k e f o r m s o f S a l v e l i n u s . I n 10 TABLE I Meristic Data from Delacy and Morton (1942) S. malma G i l l rakers on upper limb mode G i l l rakers on lower limb mode Number of vertebrae mode Pyloric Caeca mode Dorsal f i n rays Anal f i n rays 5-11 8 3-12 10 60-64 62 21-39 30 mean 10.5 mean 9 S. alpinus 7-12 10 12-15 13 63-67 65 30-64 45 mean 10.0 mean 9 11 Karluk Lake S. malma i s anadromous, while S. alpinus i s non-migratory. This difference in behaviour might account for any morphometric differences between the two nominal species in Karluk Lake. Moreover, Delacy and Morton suggest that in the Karluk Lake system S. malma spawns in streams, while S. alpinus spawns in the lake. The overlap in a l l of the characters examined suggests that hybridization might occur in Karluk Lake. Delacy and Morton did not consider hybrids in their discussion. In June 1953, 65 specimens of Salvelinus were collected by the author from Karluk Lake. A l l collections were made by gillnets and seines. A taxonomic study of this sample was undertaken using the following characters; 1 - total number of g i l l rakers on f i r s t arch. 2 - number of g i l l rakers on upper limb of f i r s t arch. 3 - number of g i l l rakers on lower limb of f i r s t arch. 4 - number of pyloric caeca. 5 - number of pores along the lateral line. 6 - number of spots in area B (Figure 2 ) . 7 - number of teeth on tongue. 3 - number of basibranchial teeth. Field observations suggested two populations of char in Karluk Lake. One population was characterized by numerous, small, yellowish-red spots, and the other population by fewer, large, bluish-red spots. A few adults and most juveniles could not be separated on this character. Detailed examination of the characters mentioned above confirmed that there are two populations of char in Karluk Lake. The total number of g i l l rakers on the f i r s t arch and the number of g i l l rakers on the 12 20 o LU =5 O UJ a: 10 (a) 15 20 25 NO. OF GILL RAKERS ON FIRST ARCH 20 i o UJ => 10 o UJ cr u. ( b ) 8 10 12 14 NO OF GILL RAKERS ON LOWER LIMB F i g . 3. D i s t r i b u t i o n of g i l l rakers i n the complete Karluk Lake sample. 13 l o w e r l i m b o f t h e f i r s t a r c h show b i m o d a l d i s t r i b u t i o n s when t h e e n t i r e K a r l u k sample i s p l o t t e d ( F i g u r e 3). The numbers o f m a l e s and f e m a l e s a r e e q u a l l y d i s t r i b u t e d i n b o t h p e a k s . D e l a c y and M o r t o n (1942) f o u n d t h a t t h e number o f g i l l r a k e r s on t h e l o w e r l i m b o f t h e f i r s t a r c h r a n g e d f r o m \u00C2\u00A3-12 i n S. malma and f r o m 12-15 i n S. a l p i n u s . U s i n g t h i s c r i t e r i o n a l l s p e c i m e n s e x c e p t t h o s e w i t h 12 g i l l r a k e r s on t h e l o w e r l i m b c a n be d e s i g n a t e d as e i t h e r S. malma o r S. a l p i n u s . E x a m i n a t i o n o f S. malma f r o m o t h e r a r e a s where o n l y S. malma o c c u r s c o n f i r m e d t h a t t h e r a n g e o f g i l l r a k e r s on t h e l o w e r l i m b i s f r o m 8-12 t h r o u g h o u t t h e N o r t h A m e r i c a n r a n g e o f t h i s s p e c i e s . A c c o r d i n g l y , t h e e n t i r e K a r l u k L a k e sample was 2 s e g r e g a t e d i n t o \"S. malma\" and \"S. a l p i n u s \" on t h e b a s i s o f t h e number o f g i l l r a k e r s on t h e l o w e r l i m b o f t h e f i r s t a r c h . F i g u r e s 4,5 and 6 p r e s e n t f r e q u e n c y d i s t r i b u t i o n s o f a l l t h e i n v e s t i g a t e d c h a r a c t e r s , s e g r e g a t e d i n t o \"S. malma\" ( d a r k b a r s ) and \"S. a l p i n u s ( l i g h t b a r s ) . I n d i v i d u a l s w i t h a n i n t e r m e d i a t e number o f g i l l r a k e r s on t h e l o w e r l i m b o f t h e f i r s t a r c h a r e r e p r e s e n t e d b y s t i p p l e d b a r s , and a r e i n c l u d e d i n b o t h h i s t o g r a m s f o r e a c h c h a r a c t e r . F i g u r e s 4,5 and 6 i n d i c a t e t h a t no s i n g l e c h a r a c t e r c a n c o m p l e t e l y s e p a r a t e \"S. malma\" and \"S. a l p i n u s \" i n K a r l u k L a k e . A c c o r d i n g l y , t h e d i s c r i m i n a t i n g powers o f s e v e r a l 2. The v a l i d i t y o f S. malma has n o t y e t b e e n d i s c u s s e d . \"S. malma\" and \"S. a l p i n u s \" a r e u s e d h e r e f o r c o n v e n i e n c e i n r e f e r r i n g t o t h e two f o r m s o f K a r l u k c h a r . F o r a d i s -c u s s i o n o f t h e v a l i d i t y o f S. malma s e e s e c t i o n I V . F i g . 5\u00C2\u00AB D i s t r i b u t i o n s of pores along the l a t e r a l l i n e and spots i n area B f o r Karluk Lake S a l v e l i n u s . o UJ cr 5 -I I 16 \u00E2\u0080\u0094 20 3 1 \u00E2\u0080\u0094 3 5 46 \u00E2\u0080\u0094 50 NO. OF TEETH ON B A SI BRA NCHIA L BONE 61 \u00E2\u0080\u0094 65 81 \u00E2\u0080\u0094 85 156 \u00E2\u0080\u0094 160 >- 5 o z UJ o UJ cr u_ - ~ l f~l r ~ r ~ i /VTZL. 12 15 18 2! 28 NO. OF TEETH ON TONGUE F i g . 6* D i s t r i b u t i o n s of tooth counts i n Karluk Lake SalveXinus. 17 c h a r a c t e r s were combined i n t o a s i n g l e d i s c r i m i n a n t s c o r e . I n o r d e r t o do t h i s i t i s n e c e s s a r y t o e l i m i n a t e t h o s e t a x o n o m i c c h a r a c t e r s s u b j e c t t o e r r o r o r b i a s . O n l y c h a r a c t e r t h a t c o u l d be c o u n t e d a c c u r a t e l y on u n s t a i n e d f i s h c o u l d be u s e d . T h i s e l i m i n a t e d b r a n c h i o s t e g a l r a y c o u n t s and t o o t h c o u n t s . The f a c t t h a t t h e t e e t h a r e d e c i d u o u s i n l a r g e s p e c i m e n s o f S. a l p i n u s a l s o s u g g e s t s t h a t t o o t h c o u n t s s h o u l d n o t be u s e d . The number o f s p o t s c o u l d n o t be u s e d b e c a u s e t h e s p o t s f a d e i n p r e s e r v e d f i s h and o f t e n c a n n o t be c o u n t e d a c c u r a t e l y . Some t a x o n o m i c c h a r a c t e r s may v a r y w i t h t h e s i z e o f f i s h . The K a r l u k L a k e s a m p l e s o f \"S. malma\" and \"S. a l p i n u s \" a r e n o t c o m p a r a b l e i n l e n g t h ; \"S. malma s p e c i m e n s a r e m o s t l y j u v e n i l e s , w h i l e \"S. a l p i n u s a r e a l l a d u l t s . T h e r e f o r e , a l l c h a r a c t e r s c o r r e l a t e d w i t h l e n g t h had t o be d i s c a r d e d . S p o t s i z e i n c r e a s e s w i t h l e n g t h and was t h e r e f o r e d i s c a r d e d . F i g u r e s 7 and 8\" show g i l l r a k e r c o u n t s on u p p e r and l o w e r l i m b s o f t h e f i r s t a r c h , number o f p y l o r i c c a e c a , and number o f p o r e s a l o n g t h e l a t e r a l l i n e , e a c h p l o t t e d a g a i n s t f o r k l e n g t h . The c o r r e l a t i o n o f e a c h c h a r a c t e r on f o r k l e n g t h has b e en t e s t e d by t h e C h i - s q u a r e t e c h n i q u e . T a b l e I I p r e s e n t s t h e C h i - s q u a r e v a l u e s and p r o b a b i l i t i e s f o r e a c h c h a r a c t e r . F i g u r e 8b s u g g e s t s t h a t p y l o r i c c a e c a a r e c o r r e l a t e d w i t h f o r k l e n g t h i n K a r l u k L a k e \"S. a l p i n u s \" . I f t h i s i s s o 18 S. MALMA 100 200 FORK LENGTH MM. m 2 tx UJ Q-Q. ZD tr UJ < 8 (b) S. ALPINUS 200 300 FORK LENGTH MM. S. MALMA 100 200 FORK LENGTH MM. !d) m 5 16 cr UJ 3 14 tr < cr 10 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 S. ALPINUS \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2\u00E2\u0080\u00A2 200 300 FORK LENGTH MM. 7\u00C2\u00AB C o r r e l a t i o n s of g i l l rakers on f o r k length i n Karluk Lake S a l v e l i n u s . 19 S. MALMA S. ALPINUS 100 200 FORK LENGTH M M , 50 < cc o 0. 40 30 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 \u00E2\u0080\u00A2 200 300 FORK LENGTH MM. S. MALMA \u00C2\u00AB 150 140 < cr UJ -o z Ui ZD a UJ cc 10 -12 16 TOTAL NO. OF 20 GILL RAKERS 24 28 ON FIRST ARCH 30 >-<_> o UJ cc 20 10 9 12 15 NO. OF GILL RAKERS LOWER LIMB 12. D i s t r i b u t i o n of g i l l rakers i n the Fraser Lake sample of S a l v e l i n u s . o 12 16 20 24 28 9 12 15 2 6 10 TOTAL NO. GILL RAKERS ON FIRST ARCH NO. GILL RAKERS LOWER LIMB NO. GILL RAKERS UPPER LIMB 5 -> o z UJ o UJ ce 25 30 35 40 45 50 55 NO. PYLORIC CAECA F i g . 13. D i s t r i b u t i o n of g i l l rakers and p y l o r i c caeca i n Fraser Lake Salvelinus 31 o z UJ o M I N I d i d X L 115 125 135 145 PORES ALONG LATERAL LINE 155 >- 5 o z U I o U I cr u. - \u00E2\u0080\u0094 L J u H J mmm m m n rfh fl fh n n ITm n m n 20 30 40 NO. SPOTS IN AREA B. 50 10 > o z ui 2 20 cr 10 I.I \u00E2\u0080\u0094 2.0-2.1\u00E2\u0080\u00943.0-3.1 \u00E2\u0080\u0094 4.0-4.1\u00E2\u0080\u00945.0- 5.1\u00E2\u0080\u00946.0-6.1\u00E2\u0080\u00947.0-7.1 \u00E2\u0080\u00948.0 SPOT SIZES EXPRESSED AS A RATIO OF INTER0RBITAL SIZE F i g . 14\u00E2\u0080\u00A2 D i s t r i b u t i o n of pores along the l a t e r a l l i n e , spots i n area B and s i z e of spots i n Fraser Lake S a l v e l i n u s . 32 15 10 -o 5 -LLQ i \u00E2\u0080\u0094 I 12 15 18 21 24 NO. OF TEETH ON TONGUE 15 r 10 ->- 5 o z UJ o UJ cc J J = L n 1-5 16-20 31-36 46-50 61-65 76-80 . 91-95 106-110 141-145 NO. OF TEETH ON BA SI BRA NCHI AL BONE F i g . 15\u00E2\u0080\u00A2 D i s t r i b u t i o n of teeth i n Fraser Lake Salvelinus\u00C2\u00BB 34 discrete groups. The single individual intermediate in number of g i l l rakers on the lower limb of the f i r s t arch is s t i l l intermediate, but closer to the \"S. malma\" group than to the \"S. alpinus\" group. In spot number and spot size this individual l i e s within the range of both \"S. malma and \"S. alpinus\", but in both of these characters i t i s near the mean of the \"S. malma\" group and at the extreme range of the \"S. alpinus\" group. These facts suggest that the individual is \"S. malma\", but do not eliminate the possibility that the intermediate i s a hybrid. Figures 17,18 and 19 present the relative growth of measured body parts for Fraser Lake char. Head length i s plotted relative to fork length. Interorbital width, vertical eye diameter, caudal peduncle depth and maxillary length are plotted relative to head length. Figure 17a shows \"S. malma\" below 220 mm. have larger heads than \"S. alpinus\" of the same size, but that the head of \"S. alpinus\" grows faster, relative to fork length, than the head of \"S. malma\". In f i s h over 220 mm. the head of \"S. alpinus\" i s larger. Figure 17b shows the interorbital width of \"S. malma\" grows faster, relative to head length, than the interorbital width of \"S. alpinus\". Figure 18a shows that the vertical eye diameter starts out larger, relative to head length, in \"S. alpinus but grows faster in \"S. malma\". At a head length of 55 mm. the vertical eye diameter of \"S. malma\" becomes larger, relative to head length, than the vertical eye 30 20 10 cr UJ t-< UJ >-UJ < o I-cr UJ > (a) J J_ 20 40 60 80 100 HEAD LENGTH MM. F i g . 18 ( a ) . R e l a t i v e growth of v e r t i c a l eye diameter i n Fraser Lake S a l v e l i n u s . 30 20 5 2 X I-0. UJ o 3 Q UJ a. < Q < 10 (b) ON J I I L_l 20 60 40 HEAD LENGTH MM. F i g . 18 ( b ) . Rela t i v e growth of caudal peduncle i n Fraser Lake S a l v e l i n u s . 80 100 37 80 20 40 60 80 HEAD LENGTH MM. F i g . 19* R e l a t i v e growth of ma x i l l a r y i n Fraser Lake S a l v e l i n u s . 38 diameter of \"S. alpinus\". Figure 18b shows the caudal peduncle depth starts out larger, relative to head length, in \"S. alpinus\" but grows faster in \"S. malma\". At a head length of 35 mm. the caudal peduncle of \"S. malma\" becomes larger, relative to head length than the caudal peduncle of \"S. alpinus\". Figure 19 shows the maxillary starts out longer, relative to head length, in \"S. malma\", but grows faster in \"S. alpinus\". At a head length of 70 mm. the maxillary of \"S. alpinus\" becomes longer than the maxillary of \"S. malma\". Figures 17, 1$ and 19 show some differences in the relative growth of body parts of \"S. malma\" and \"S. alpinus\" in Fraser Lake, but none of these differences are sufficient to separate the two species. Brooks Lake and Tributaries Brooks Lake i s located on the Alaska Peninsula (locality 2, Figure 29) and i s tributary to Bristol Bay. In the summer of 195$ 21 char were collected from Brooks Lake and its tributaries by T. Merrill of the U.S. Fish and Wildlife Service. Twenty of the char are from tributaries and one i s from Brooks Lake proper. Figure 20 shows the frequency dis-tributions of g i l l rakers, pyloric caeca, and pores along the lateral line in the Brooks Lake sample. There is no indication of bimodality in any of these characters. Judging from Karluk and Fraser Lake samples, the dumber of g i l l rakers on the lower limb of the f i r s t arch i s 39 10 r o z UJ i 5 cr 10 18 21 24 TOTAL NO. GILL RAKERS FIRST ARCH > o \u00E2\u0080\u00A2 z UJ o UJ cr 9 12 15 NO. GILL RAKERS LOWER LIMB 10 o z UJ UJ cr u. 5 10 NO. GILL RAKERS UPPER LIMB 10 o 5 O ui cr n n n n n n 20 25 30 PYLORIC CAECA 35 40 10 r->-o z UJ g 5 UI cr u. IL rj rm llmlmlnh 110 120 130 PORES ALONG LATERAL LI-NE 140 150 F i g . 20. D i s t r i b u t i o n of taxonomic characters exandned on Brooks Lake sample. 40 the best morphological difference between \"S. malma\" and \"S. alpinus\". The mean number of g i l l rakers on the lower limb in the Brooks Lake sample i s 12.3, with a range of 11 - 14. The sample is therefore intermediate between \"S. malma\" and \"S. alpinus\" in this character. In order to include other characters in a consider-ation of the Brooks Lake sample a discriminant score was cal-culated for each f i s h . Because of the small size of the Brooks Lake sample, coefficients obtained from combining the Karluk and Fraser Lake samples were used to calculate the 3 Brooks Lake scores. Figure 21a presents the discriminant scores for the combined Karluk and Fraser Lake samples. Figure 21b presents the distribution of discriminant scores for the Brooks Lake sample. Figures 21a and 21b suggest that a l l of the char from the Brooks Lake basin with 12 g i l l rakers on the lower limb of the f i r s t arch are \"S. malma\". This suggestion i s further strengthened by the fact that a l l of the char with 12 g i l l rakers were collected in streams. Delacy and Morton (1942) suggest that in Karluk Lake \"S. alpinus\" i s mainly lacustrine and that \"S. malma\" inhabits both the lake and tributary streams. The one specimen from Brooks Lake proper is undoubt-ably \"S. alpinus\". Thirteen of the specimens from Brooks Lake can be 3. It i s realized that the combined Karluk and Fraser Lake coefficients are probably not the same as the Brooks Lake coefficients. They are used here as approximations. (a) 8 (b) COMBINED KARLUK AND FRASER FI n d tti HlQWLn. 10 12 13 14 15 BROOKS LAKE 8 (C) vl ' i t v n n i l i n n 9 10 II 12 13 KENAI R. 15 .EL X L 10 II 12 DISCRIMINANT SCORES 13 14 15 F i g . 21. Discriminant scores f o r combined Karluk, Fraser Lakes, Brooks Lake and Kenai River. 42 c o n s i d e r e d a s \"S. malma\" and 8\" as \"S. a l p i n u s \" . The mean number o f g i l l r a k e r s on t h e l o w e r l i m b o f t h e f i r s t a r c h o f t h e \"S. malma\" sample i s 11.7. T h i s i s t h e h i g h e s t mean f o r t h i s c h a r a c t e r f o u n d i n any o f t h e p o p u l a t i o n s o f \"S. malma\" examined. The mean number o f g i l l r a k e r s on t h e l o w e r l i m b i n t h e \"S. a l p i n u s \" sample i s 13.2. T h i s i s t h e l o w e s t mean f o r t h i s c h a r a c t e r f o u n d i n any o f t h e p o p u l a t i o n s o f \"S. a l p i n u s \" s t u d i e d . F i g u r e 21b i n d i c a t e s t h a t t h e d i s c r i m -i n a n t s c o r e s f o r t h e B r o o k s L a k e sample a r e more c l o s e l y g r o u p e d t h a n t h e s c o r e s f o r t h e combined K a r l u k and F r a s e r L a k e s a m p l e s . T h i s e v i d e n c e s u g g e s t s i n t r o g r e s s i o n b e t w e e n \"S. malma\" and \"S. a l p i n u s \" i n t h e B r o o k s L a k e a r e a . However, t h e s m a l l s i z e o f t h e s a m p l e s (13 \"S. malma.\" and 8 \"S. a l p i n u s \" ) make any i n t e r p r e t a t i o n a m a t t e r o f s p e c u l a t i o n . K e n a i R i v e r Two s p e c i m e n s were c o l l e c t e d i n t h e K e n a i R i v e r ( l o c a l i t y 22, F i g u r e 29). One c h a r had 12 g i l l r a k e r s on t h e l o w e r l i m b o f t h e f i r s t a r c h and numerous s m a l l s p o t s . The o t h e r c h a r had 13 g i l l r a k e r s o n t h e l o w e r l i m b o f t h e f i r s t a r c h and f e w e r , l a r g e r s p o t s . G i l l r a k e r s on t h e u p p e r l i m b , g i l l r a k e r s on t h e l o w e r l i m b , p y l o r i c c a e c a and p o r e s a l o n g t h e l a t e r a l l i n e were combined i n t o a d i s c r i m i n a n t s c o r e . The c o e f f i c i e n t s o b t a i n e d f r o m t h e combined K a r l u k and F r a s e r L a k e s a m p l e s were a g a i n u s e d t o c a l c u l a t e t h e s e s c o r e s . F i g u r e 21c p r e s e n t s t h e K e n a i R i v e r d i s c r i m i n a n t s c o r e s . The s p e c i m e n w i t h 12 g i l l r a k e r s on t h e l o w e r l i m b f a l l s i n t o t h e \"S. malma\" 43 g r o u p , w h i l e t h e s p e c i m e n w i t h 13 g i l l r a k e r s i s p r o b a b l y \"S. a l p i n u s \" . Summary o f Taxonomic R e l a t i o n s h i p o f S. malma and S. a l p i n u s . \"S. malma\" and \"S. a l p i n u s \" were f o u n d s y m p a t r i c a l l y i n K a r l u k and F r a s e r L a k e s . I n l a r g e s a m p l e s f r o m t h e s e l a k e s i t was p o s s i b l e t o s e p a r a t e t h e n o m i n a l s p e c i e s i n t o d i s c r e t e g r o u p s w i t h o n l y two q u e s t i o n a b l e h y b r i d s . E v i d e n c e was p r e -s e n t e d w h i c h i n d i c a t e s \"S. malma\" and \"S. a l p i n u s \" o c c u r sym-p a t r i c a l l y i n B r o o k s L a k e a n d t h e K e n a i R i v e r . B e c a u s e S. malma o c c u r s s y m p a t r i c a l l y w i t h S. a l p i n u s i t must be c o n s i d e r e d a d i s t i n c t s p e c i e s . G e o g r a p h i c V a r i a b i l i t y The g e o g r a p h i c v a r i a b i l i t y o f some t a x o n o m i c c h a r -a c t e r s o f S. malma were s t u d i e d . L o c a l i t i e s s t u d i e d l i e b etween 49\u00C2\u00B0N and 60\u00C2\u00B0N. C h a r a c t e r s e x amined were; g i l l r a k e r s on l o w e r l i m b o f f i r s t a r c h , p y l o r i c c a e c a , p o r e s a l o n g t h e l a t e r a l l i n e , number o f v e r t e b r a e , number o f t e e t h on t o n g u e , and hea d l e n g t h r e l a t i v e t o f o r k l e n g t h . F i g u r e s 22-26 p r e s e n t t h e g e o g r a p h -i c a l v a r i a t i o n i n t h e above c h a r a c t e r s . The s a m p l e s a r e a r r a n g e d i n a s e r i e s s t a r t i n g i n n o r t h w e s t e r n N o r t h A m e r i c a and e x t e n d i n g a r o u n d t h e r i m o f t h e N o r t h P a c i f i c t o J a p a n . D a t a f r o m t h e l i t e r a t u r e was i n c l u d e d where a v a i l a b l e . T h e r e i s c o n s i d e r a b l e g e o g r a p h i c v a r i a t i o n i n a l l o f t h e c h a r a c t e r s examined. T h i s v a r i a t i o n c o u l d be a r e s u l t o f e i t h e r e n v i r o n m e n t a l o r g e n e t i c d i f f e r e n c e s b etween 1 r \u00E2\u0080\u00A2 10 II NO. OF GILL RAKERS FRASER L. KARLUK L. MIDARM L. TURNAGAIN ARM SUMMIT L. SELTAT CR. TAKU R. BOWSER L. COTTONWOOD R. OCEAN FALLS SUMALLO R. SAGE CR. GLACIER NAT. PARK MEAN 2(ST. ERROR) ST. DEVIATION RANGE ( 0 ) INDICATES ANADROMOUS POPULATIONS F i g . 22. Geographic v a r i a t i o n i n g i l l rakers on the lower limb of S. malma. 45 a , , ! , , 10 20 30 40 50 NO. OF PYLORIC CAECA FRASER L. KARLUK L. MIDARM L. TURNAGAIN ARM. SUMMIT L. SELTAT CR . TAKU R. BOWSER L. COTTONWOOD R OCEAN FALLS SUMALL0 R . SAGE CR. GLACIER NAT. PA F i g . 23\u00C2\u00BB Geographic v a r i a t i o n i n p y l o r i c caeca of S. malma. 46 i mim\ i IHJLH i i mimt i i I J M I - i 1 \ 1\u00E2\u0080\u0094 110 120 130 140 NO. OF PORES ALONG THE LATERAL LINE FRASER L. 0 KARLUK L. MIDARM L a TURNAGAIN ARM SUMMIT L. SELTAT CR. a TAKU R. BOWSER L. COTTONWOOD R. a OCEAN FALLS SUMALLO R. SAGE CR. GLACIER NAT. PARK F i g . 24* Geographic v a r i a t i o n i n the number of pores along the l a t e r a l l i n e of S. malma. 47 2 , 3 4 5 RATIO OF HEAD LENGTH TO FORK LENGTH PETER THE GREAT BAY NELMA R. TARTARY ST. SHANTAR IS. TAUIS R. OLA R . KAMCHATICA COMANDOR IS. UNALASKA FRASER L . KARLUK L. MIDARM L. TURNAGAIN ARM SUMMIT L. SELTAT CR. TAKU R. BOWSER L . COTTONWOOD R OCEAN FALLS SUMALLO R . SAGE CR. GLACIER NAT. PARK 1 6 F i g . 25* Geographic v a r i a t i o n of r e l a t i v e head length of S. malma. 43 v 9 10 NO. OF TEETH 12 13 ON TONGUE 14 1 15 FRASER L. KARLUK L. MIDARM L. TURNAGAIN ARM SUMMIT L, PARSNIP R. BELLA COOLA R SHEEP CREEK COQUIHALLA R, SUMALLO R. (b) HOKKAIDO SOUTH OF AMUR GULF NORTH OF AMUR GULF TURNAGAIN ARM SELTAT CR. PARSNIP R. BELLA COOLA R. SHEEP CR. COQUIHALLA R. , , , , , 50 55 60 65 70 NO. OF VERTEBRAE F i g . 26. (a) Geographic v a r i a t i o n of teeth on tongue of S. malma. (b) Geographic v a r i a t i o n of vertebrae number i n S. malma. 49 populations, or an interaction of both. Populations located in similar habitats only a few miles apart (Fraser and Karluk Lakes) show large differences in some characters ( g i l l rakers and pyloric caeca), while populations several hundred miles apart (Seltat Creek and Sumallo River) are quite similar in these characters. Anadromous populations show no consistant differences from non-migratory populations. No obvious relationship is appar-ent between latitude or loca l i t y and variation of any char-acter. If the variation were due entirely to environment, clinal variation might be expected. However, most samples are from the mountainous coastal regions where the environment i s largely influenced by local factors such as proximity to the sea, altitude and glaciers. Such local factors could easily obliterate any c l i n a l changes in environment. Figure 27a presents the mean number of g i l l rakers on the lower limb of the f i r s t arch for 12 samples arranged in geographic sequence from south to north. There i s no apparent trend to the data. Figure 27b presents the mean pyloric caeca counts for the same 12 samples. The pattern of variation is very similar to that of g i l l rakers. Figure 28a indicates there i s a correlation between the mean g i l l raker counts and the mean pyloric caeca counts. If the correlation between the means of g i l l rakers and pyloric caeca was due to genetic linkage of these two characters i t would be reasonable to expect that the characters would be correlated within a single 50 F i g . 27. (a) (b) Geographic Geographic v a r i a t i o n v a r i a t i o n i n mean g i l l raker count, i n mean p y l o r i c caeca count. 51 (a ) in cc U J < cc _1 _ l 2 < 10 r : .87 P= .001 40 20 30 MEAN NO. PYLORIC CACEA 28 ( a ) . C o r r e l a t i o n of- mean g i l l raker number and mean p y l o r i c caeca count. \u00E2\u0080\u00A2 12 cc UJ o l l 10 - j 9 (b) 2 X = 0.4 P = 0.9 20 30 PYLORIC CAECA 28 ( b ) . C o r r e l a t i o n of g i l l raker count and p y l o r i c caeca i n S e l t a t Creek. 52 population. Figure 27b (Seltat Creek sample), shows that there is no correlation between g i l l rakers on the lower limb and pyloric caeca within a population. This suggests that the variation in g i l l rakers and pyloric caeca i s probably due to some environmental factor that effects both characters in a similar fashion. For most characters studied there is no indication whether the variation i s the result of environmental or genetic differences, and i t is probably the result of an interaction of both. Taxonomic interpretation of these results is dealt with in the Discussion. Description of Salvelinus malma A description of S. malma is included for several reasons. None of the previous descriptions (Jordan and Evermann 1898, Berg 1943, Aoyagi 1955), include a l l of the characters necessary to distinguish S. malma from S. alpinus. Most pre-vious decriptions have not incorporated published data from the entire range of S. malma. The number of g i l l rakers on the lower limb of the f i r s t arch is not included in most des-criptions. The presence of vermiculations on the dorsal sur-face of some S. malma has not been noted in previous descrip-tions. Salvelinus malma (Walbaum) Salmo malma Walbaum, Artedi P i s e , 1792, Kamchatka. Salmo curilus Pallas, Zoogr. Rosso-Asiat., III, 1811, Curile Islands. Salmo callaris Pallasj, loc. c i t . , Bering Sea. Salmo laevigatus^WallagX loc. c i t . , Curile Islands. 53 Salmo nummifer Cuvier and Valenciennes, Hist. Nat. Poiss., XXI, 1848, Kamchatka. Salmo erythrochychos Cuvier and Valenciennes, loc. c i t . , Kamchatka. Salmo tudes Cope, Proc. Amer. Phil. Soc. Phila., X l l l , !L8\"73, Captains Harbour, Unalaska. Salmo pluvius Hilgendorf, Mitth. Deutsch. Geo. Ostasiens, XI, 187b, Japan. Salmo spectabilis Girard, Proc. Acad. Mat. Sci. Phila., V l l l , 1856, Fort Dalles, Oregon. Salmo ba i r d i i Suckley, loc. c i t . , Flathead River, Montana. Salmo parkei Suckley, arm. Lyc. Nat. Hist. M.Y., V l l , 18&L, Kootenay River. Salmo campbelli Suckley, Loc. Cit., Fort Dalles, Oregon. Salmo Tbrdii Gi'inther, l a t . , VI, 1866, Skagit River. Head 3.2 to 5.8 into fork length. D. 13-16 ( a l l rays); A. 11-15 ( a l l rays). Body elongate, somewhat rounded; vertical eye diameter 4-5.5 into head; mouth terminal, large; maxillary usually reaching to a point behind posterior margin of the eye. Teeth on head of vomer only: 7-15 teeth on tongue; 0-44 teeth on the basibranchial bone, the number of teeth on the basibran-chial tends to increase with length. G i l l rakers on f i r s t arch 11-26 (13-23 maximum recorded range in North America); g i l l rakers on lower limb of f i r s t arch 8-12; g i l l rakers oh upper limb of f i r s t arch 3-9. Pyloric caeca 13-47. Branchiostegal rays 21-30, usually more on l e f t side than on right side. Total number of vertebrae 58-69. Lateral line slightly decurved anteriorly, then straight, often with a dip or arch posteriorly; pores along the lateral line 105-142. Colour variable; blue, olive green or brown on dorsal surface; occasionally bright red on sides; pelvic and anal fins often with creamy leading edges. Spotting: yellow, orange or red spots on dorsal surface and sides; spots small and numerous, the largest touching lateral line 3-7 times into interorbital width. Dorsal surface 54 4 occasionally with verraiculations. Key to the Adult Char of Western North America Previous keys (Jordan and Evermann 1898, Wilamovsky 195^) that include a l l of the chars of western North America use the presence or absence of vermiculations to distinguish S. fontinalis from S. malma. This character is not exclusive to S. fontinalis and the present key does not rely on i t to separate S. fontinalis from S. malma. A Caudal f i n deeply forked; spots not highly coloured, with irregular outlines; pyloric caeca 90-170. Lake Trout - Salvelinus namaycush (Walbaum) AA Caudal f i n truncate; spots yellow, orange or red, mostly round; less than 65 pyloric caeca. B Distinct dark marbling on back and dorsal f i n ; blue halos around some of the spots on the sides of body; usually no teeth on basibranchial bone. Eastern Brook Trout - Salvelinus fontinalis (Mitchill) BB Usually no distinct dark marbling on the back, no distinct dark marbling on dorsal f i n ; no blue halos around some of the spots on the sides of body; usually teeth on basibranchial bone. Dolly Varden - Salvelinus malma (Walbaum) or Arctic Char - Salvelinus alpinus. (Linnaeus). No satisfactory key has yet been made to distinguish a l l S. malma from a l l S. alpinus. Table III gives a comparison of the principal characters used to distinguish them. 4. Specimens of S. malma with vermicuations on the dorsal surface have Seen collected in the Sumallo River (south western B.C.), and in several North Coast streams. The vermiculations are distinct, but not as well developed as in S. fontinalis. 55 TABLE III A Comparison of Sj_ malma and Sj_ alpinus G i l l rakers on lower limb Pyloric caeca Pores along the lateral line Teeth on tongue Teeth on basibron-chial Branchiostegal rays Size of spots along lateral line S. malma 8-12 usually 10 or 11 13-47 105-142 7-15 0-44 usually in one or two rows 21-30 small 3-7 times into the interorbital, usually 5 S. alpinus 12-16 usually 14 or 15 25-64 120-155 10-28 16-160 usually in three or more rows 21-26 large 2-7 times into the interorbital, usually 3-5 56 Distribution of Salvelinus malma Salvelinus malma ranges from northern California around the rim of the North Pacific and Bering Sea to Japan and Korea. Over most of this range S. malma has both anadro-mous and non - migratory populations. At the southern extremes of i t s range S. malma i s usually found as a non - migratory form. Taranetz (1936) reports anadromous populations of S. malma as far south as Peter the Great Bay. Non - migratory 5 forms are found in southern Honshu and in the Talu River (Taranetz 1936). In North America, the southern extent of the range is represented by isolated, non-migratory, populations in northern Nevada (Miller and Morton 1952) and northern California (Wales 1957). In Asia S. malma is not reported north of the Anadyr River (Berg 1948). The exact northern limits of the range in North America are unknown. Salvelinus malma i s re-ported from the Arctic slope of Alaska and the Yukon Territory. 6 Murdock (1885a, 1885b) reported S. malma from the Colville 6 River mouth, Pergniak and Point Barrow. Scofield reports S. malma from Point Hope and Herschel Island. Salvelinus alpinus is common in the Bering Sea area and along the artic slope of Alaska and the Yukon Territory. Possibly some of the records of S. malma from areas where S. alpinus also occurs are 5. Many Japanese authors consider the char found on Honshu to be a distinct species Salvelinus pluvius (Hilgendorf). 6. References not seen, reports from Evermann and Goldberg (1906) and Walters (1955). 57 due t o c o n f u s i n g t h e two s p e c i e s . S. malma i s n o t common i n c e n t r a l A l a s k a and c e n t r a l Yukon T e r r i t o r y . I t a p p e a r s t o be c o n f i n e d t o h e a d w a t e r h a b i t a t s i n t h e s e a r e a s . Everraann and G o l d b e r g (1906) r e p o r t S. malma f r o m Mynock Gr e e k ( t r i b u t a r y o f t h e Yu k o n R i v e r ) n e a r Rampart. The o n l y S. malma o b t a i n e d f r o m c e n t r a l A l a s k a d u r i n g t h e 1958 c o l l e c t i o n s were f r o m D r y C r e e k , a s m a l l t r i b -u t a r y o f t h e Tanana R i v e r . No S. malma.was t a k e n i n t h e Tanana o r Yukon R i v e r s p r o p e r . Wynne - Edwards (1947) r e p o r t s S. malma f r o m t h e c l e a r s t r e a m s o f t h e M a c k e n z i e and R i c h a r d s o n M o u n t a i n s . P r e b l e (1908) r e p o r t s S. malma f r o m t h e h e a d w a t e r s o f t h e P e e l R i v e r . I n B r i t i s h C o l u m b i a S. malma i s w i d e l y d i s t r i b u t e d i n a l l r i v e r s y s t e m s e x c e p t t h e Okanagan ( C a r l and Clemens 1953). On t h e e a s t s l o p e o f t h e R o c k y M o u n t a i n s S. malma i s abun d a n t i n t h e h e a d w a t e r s o f t h e S o u t h S a s k a t c h e w a n , N o r t h S a s k a t c h e w a n , A t h a b a s k a and P e a c e R i v e r s b u t does n o t e x t e n d f a r o n t o t h e p l a i n s . S. malma i s w i d e l y d i s t r i b u t e d t h r o u g h -o u t most o f t h e C o l u m b i a s y s t e m , b u t has o n l y b e e n r e p o r t e d once (Hubbs and M i l l e r 194$) f r o m t h e Snake R i v e r , a m a j o r t r i b u t a r y o f t h e C o l u m b i a . F i g u r e 29 p r e s e n t s t h e N o r t h A m e r i c a n d i s t r i b u t i o n o f S. malma. 58 F i g . 29\u00E2\u0080\u00A2 Map i n d i c a t i n g North American range of S. malma. 59 IV. DISCUSSION The p r i m a r y c r i t e r i o n o f a b i o l o g i c a l s p e c i e s i s w h e t h e r h y b r i d i z a t i o n o c c u r s where t h e s p e c i e s o c c u r s s y m p a t r i -c a l l y w i t h a c l o s e l y r e l a t e d s p e c i e s (Mayr 1957, C a i n 1954). O c c a s i o n a l h y b r i d i z a t i o n c a n o c c u r b u t as l o n g as t h e h y b r i d s a r e r a r e t h e s p e c i e s may be c o n s i d e r e d v a l i d . G r a n t (1957) p o i n t s o u t t h a t an o c c a s i o n a l h y b r i d , i s a v e r y d i f f e r e n t s i t u a t i o n f r o m a mass f l o w o f g e n e s between s p e c i e s . The v a l i d i t y o f d o u b t f u l s p e c i e s p a i r s c a n o n l y be e s t a b l i s h e d i f t h e y o c c u r s y m p a t r i c a l l y w i t h o u t mass h y b r i d i z -a t i o n . The p r e s e n t s t u d y was u n d e r t a k e n t o d e t e r m i n e the. v a l i d i t y o f S. malma. . T h r e e l a k e s were s a m p l e d where S. malma and S. a l p i n u s o c c u r s y m p a t r i c a l l y . The e v i d e n c e i n d i c a t e s t h a t w i t h i n t h e s e l a k e s h y b r i d i z a t i o n between S. malma and S. a l p i n u s r a r e l y , i f e v e r , o c c u r s . T h e r e f o r e S. malma must be c o n s i d e r e d a v a l i d s p e c i e s . J o r d a n , Evermann and C l a r k (1930) s u g g e s t t h a t i n N o r t h A m e r i c a S. malma i n t e r g r a d e s s o u t h w a r d s t o f o r m t h e s u b -s p e c i e s S a l v e l i n u s malma s p e c t a b i l i s . S. m. malma i s d i s t i n g -u i s h e d f r o m S. m. s p e c t a b i l i s b y t h e l e n g t h o f t h e head ( l e s s t h a n \u00C2\u00A3 i n t o s t a n d a r d l e n g t h i n S. m. malma, more t h a n \ i n S. m. s p e c t a b i l i s ) . F i g u r e 25 i n d i c a t e s t h a t t h e r e i s some g e o g r a p h i c v a r i a b i l i t y i n t h e l e n g t h o f t h e h e a d r e l a t i v e t o body l e n g t h , b u t t h a t t h e r e i s no e v i d e n c e t o s u p p o r t s u b -s p e c i f i c d i s t i n c t i o n b a s e d on t h i s c h a r a c t e r . 60 T a r a n e t z (1936) and B e r g (194$) l i s t one s u b s p e c i e s , 7 S a l v e l i n u s malma k r a s h e n i n n i k o v l T a r a n e t z , f r o m R u s s i a . S. m. k r a s h e n i n n i k o v i i s d i s t i n g u i s h e d f r o m S. m. malma by t h e number o f v e r t e b r a e (59-64 i n S. m. k r a s h e n i n n i k o v i a nd 64-69 i n S. m. malma). F i g u r e 26b shows t h e g e o g r a p h i c v a r i a b i l i t y i n v e r t -e b r a e f o r S. malma. The f o r m o f S. malma s o u t h o f t h e G u l f o f Amur c a n n o t be r e a d i l y d i s t i n g u i s h e d f r o m o t h e r S. malma on t h e b a s i s o f v e r t e b r a l c o u n t s . U n l e s s S. m. k r a s h e n i n n i k o v i c a n be d i s t i n g u i s h e d f r o m S. m. malma on some o t h e r c h a r a c t e r S_. m. k r a s h e n i n n i k o v i c a n n o t be c o n s i d e r e d a v a l i d s u b s p e c i e s . J o r d a n , Evermann and C l a r k (1930) l i s t S a l v e l i n u s p l u v i u s ( H i l g e n d o r f ) as a synonym f o r S. malma. M o r i (1936) c o n s i d e r e d S. p l u v i u s a s u b s p e c i e s o f S. malma. R e c e n t J a p a n e s e a u t h o r s ( A o y a g i 1957, M a t s u b a r a 1955, Okado 1955) c o n s i d e r S. p l u v i u s t o be a d i s t i n c t s p e c i e s . A o y a g i (1957) d i s t i n g u i s h e s S. malma f r o m S. p l u v i u s on t h e b a s i s o f g i l l r a k e r s and p a r r marks (16-20 g i l l r a k e r s and no p a r r marks i n a d u l t S_. malma, 11-20 g i l l r a k e r s and p a r r marks i n a d u l t S. p l u v i u s ) . The g i l l raker c o u n t f o r S. p l u v i u s i s b a s e d on 7 s p e c i m e n s . E v e n w i t h t h i s s m a l l number t h e g i l l raker c o u n t c o n s i d e r a b l y o v e r l a p s t h a t g i v e n f o r S. malma. I n N o r t h A m e r i c a s t u n t e d p o p u l a t i o n s o f S. malma commonly have p a r r marks as a d u l t s . A o y a g i (1957) r e p o r t s S. malma f r o m H o k k a i d o and S_. p l u v i u s f r o m Honshu; p r e s u m a b l y t h e i r r a n g e s n e v e r o v e r l a p . The e v i d e n c e s u g g e s t s 7. T a r a n e t z (1936) l i s t s two i n f r a s p e c i e s f r o m R u s s i a , S a l v e l i n u s malma i n f r a s p e c i e s c u r i l u s ( P a l l a s ) and S a l v e l i n u s malma i n f r a s p e c i e s k u z n e t z ' o v i T a r a n e t z . T h e s e f o r m s a r e e c o t y p e s . 61 p l u v i u s i s a g e o g r a p h i c f o r m o f S. malma and p r o b a b l y s h o u l d n o t be c o n s i d e r e d a d i s t i n c t s p e c i e s , w h e t h e r S. p l u v i u s s h o u l d be c o n s i d e r e d a s e p a r a t e s u b s p e c i e s o f S. malma c a n n o t be d e -t e r m i n e d on t h e a v a i l a b l e e v i d e n c e . B a i l e y , Winn and S m i t h (1954) s u g g e s t t h a t , n a s p e c i e s i s p r o p e r l y d i v i s a b l e i n t o s u b s p e c i e s when d a t a have b e e n s u i t a b l y p u b l i s h e d t o d e m o n s t r -a t . a ' t h a t i t c o n s i s t s o f two o r more a l l o p a t r i c p o p u l a t i o n s , e a c h d i s p l a y i n g a h i g h d e g r e e o f u n i f o r m i t y o v e r i t s r a n g e and d i f f e r i n g w i t h a h i g h c o n s t a n c y ( a f i g u r e o f a t l e a s t 93 p e r c e n t o f i n d i v i d u a l s i s s u g g e s t e d ) f r o m o t h e r f o r m s , e a c h o f w h i c h i n t e r g r a d e s o v e r a r e l a t i v e l y n a r r o w g e o g r a p h i c a r e a w i t h a t l e a s t one o t h e r f o r m . \" None o f t h e named s u b s p e c i e s o f S. malma a p p e a r t o f u l f i l l t h e s e c r i t e r i a and t h e r e f o r e , s h o u l d n o t be r e c o g n i z e d a s v a l i d s u b s p e c i e s . O r i g i n o f S. malma The m o r p h o l o g i c a l s i m i l a r i t y between S. malma and S. a l p i n u s s u g g e s t s t h a t t h e y a r e d e r i v e d f r o m a common a n c e s t o r . Some s p e c u l a t i o n c a n be made on t h e t i m e a t w h i c h S. malma and S. a l p i n u s d i v e r g e d . Ekman (1953) p r e s e n t s e v i d e n c e i n d i c a t i n g a f a i r l y c o m p r e h e n s i v e f a u n i s t i c c o n n e c t i o n between t h e c o l d -t e m p e r a t e f a u n a s o f t he A t l a n t i c and P a c i f i c d u r i n g t h e l a t e P l i o c e n e o r e a r l y P l e i s t o c e n e . I f S. malma had b e e n d i f f e r -e n t i a t e d a t t h i s t i m e i t i s l i k e l y t h a t i t w o u l d have r e a c h e d t h e N o r t h A t l a n t i c . R e l i c t p o p u l a t i o n s o f S. malma i n n o r t h e r n C a l i f o r n i a , J a p a n and K o r e a i n d i c a t e t h a t S. malma was w i d e l y d i s t r i b u t e d i n t h e N o r t h P a c i f i c p r i o r t o t h e l a s t P l e i s t o c e n e 62 8 g l a e i a t i o n . R e l i c t p o p u l a t i o n s o f S. a l p i n u s i n New H a m p s h i r e , 8 Maine and E u r o p e i n d i c a t e t h a t S. a l p i n u s was w i d e l y d i s t r i -b u t e d i n t h e N o r t h A t l a n t i c p r i o r t o t h e l a s t P l e i s t o c e n e g l a e i a t i o n . S. malma and S. a l p i n u s must t h e r e f o r e have d i v e r g e d sometime i n t h e P l e i s t o c e n e p r i o r t o t h e l a s t g l a e i a t i o n . S. malma p r o b a b l y e v o l v e d i n t h e N o r t h P a c i f i c a r e a . The N o r t h P a c i f i c was i s o l a t e d f r o m t h e A r c t i c Ocean s e v e r a l t i m e s d u r i n g t h e P l e i s t o c e n e by t h e B e r i n g l a n d b r i d g e . I t i s g e n e r a l l y a c c e p t e d t h a t t h e l a n d b r i d g e was f l o o d e d d u r i n g t h e i n t e r g l a c i a l p e r i o d s , a l t h o u g h Ekman (1953) i n d i c a t e s t h a t v e r y l i t t l e i s known a b o u t p o s s i b l e i n t e r g l a c i a l c o m m u n i c a t i o n s b e -tween t h e N o r t h P a c i f i c a nd N o r t h A t l a n t i c . S. malma must have b e e n e v o l v e d p r i o r t o one o f t h e s e i n t e r g l a c i a l f l o o d i n g s , b u t d i d n o t e x t e n d i t s e l f i n t o t h e A t l a n t i c . Neave (195$) s u g g e s t s t h a t \" t h r o u g h o u t t h e a v a i l a b l e i n t e r g l a c i a l i n t e r v a l s t h e p h y -s i c a l c o n d i t i o n s o f t h e P o l a r S ea d i d n o t become much more f a v o u r a b l e t h a n t h e y a r e a t p r e s e n t . \" The p r e s e n t r a n g e o f S. malma does n o t e x t e n d f a r , i f a t a l l , i n t o t h e A r c t i c Ocean. I t may be s u p p o s e d t h a t t h e r e i s some p h y s i c a l f a c t o r l i m i t i n g t h e n o r t h w a r d d i s t r i b u t i o n o f S. malma and t h a t t h i s p r e v e n t e d t h e s p r e a d o f S. malma i n t o t h e A r c t i c Ocean d u r i n g t h e i n t e r -g l a c i a l p e r i o d s . 8\". T h e s e p o p u l a t i o n s a r e somewhat d i v e r g a n t and o f t e n c o n -s i d e r e d s e p a r a t e s p e c i e s , S a l v e l i n u s a u r e o l u s Bean i n New Hampshire and S a l v e l i n u s o p u a s s a ( G i r a r d ) i n M a i n e . Backus (1957) d i s c u s s e s t h e r e l a t i o n s h i p o f S. a l p i n u s , S. a u r e o l u s and S. o p u a s s a . S a l v e l i n u s m a r s t o n i (Garman) a d i v e r g a n t f o r m i n Quebec i s a l s o p r o b a b l y a r e l i c t p o p u l a t i o n o f S. a l p i n u s . 63 M o r p h o l o g i c a l l y S_. malma and S. a l p i n u s a r e v e r y s i m i l a r . B o t h s p e c i e s spawn i n t h e f a l l o r e a r l y w i n t e r . E v i d e n c e f r o m r e l a t e d s p e c i e s ( S t e n t o n 1952, Buss and W r i g h t 1956) s u g g e s t t h a t t h e i n t e r b r e e d i n g o f S. malma and S_. a l p i n u s w o u l d p r o d u c e f e r t i l e o f f s p r i n g . T h e r e f o r e , some mechanism f o r p r e v e n t i n g i n t e r b r e e d i n g must have e v o l v e d w h i l e S. malma and S. a l p i n u s were g e o g r a p h i c a l l y i s o l a t e d . I t i s s u g g e s t e d t h a t t h e i n c i p i e n t s p e c i e s d e v e l o p e d d i f f e r e n c e s i n s p a w n i n g b e h a v -i o u r w h i c h p r e v e n t e d i n t e r b r e e d i n g a f t e r t h e g e o g r a p h i c a l . b a r r i e r s s e p a r a t i n g them were b r o k e n down. T h i s s u g g e s t i o n i s s u p p o r t e d by t h e s p a w n i n g b e h a v i o u r o f t h e two s p e c i e s i n K a r l u k L a k e . I n t h e K a r l u k s y s t e m S. malma spawns i n t r i b u -t a r y s t r e a m s , and S. a l p i n u s spawns i n l a k e s . T h i s b e h a v i o u r a l d i f f e r e n c e , i f a b s o l u t e , p r o v i d e s a mechanism w h i c h p r e v e n t s i n t e r b r e e d i n g b etween S. malma and S. a l p i n u s . D i s p e r s i o n o f S. malma I n N o r t h A m e r i c a S. malma p r o b a b l y s u r v i v e d t h e l a s t P l e i s t o c e n e g l a c i a t i o n i n . t w o a r e a s . I s o l a t e d h e a d w a t e r p o p -u l a t i o n s f o u n d i n t h e Yukon V a l l e y s u g g e s t t h a t S. malma c o u l d have s u r v i v e d i n t h i s u n g l a c i a t e d a r e a o f A l a s k a . The p r e s e n t r a n g e o f S. malma e x t e n d s w e l l s o u t h o f t h e s o u t h e r n l i m i t s o f g l a c i a t i o n and s u g g e s t t h a t S. malma a l s o s u r v i v e d s o u t h o f t h e i c e s h e e t on t h e we s t s i d e o f t h e c o n t i n e n t a l d i v i d e . The p r e s e n t N o r t h A m e r i c a n d i s t r i b u t i o n o f S. malma h a s p r o b a b l y b e e n a t t a i n e d by b o t h f r e s h w a t e r and m a r i n e d i s p e r s i o n f r o m t h e s e c e n t e r s . S a l v e l i n u s malma i s o f t e n f o u n d i n h e a d w a t e r 64 s t r e a m s w h i c h a r e h i g h l y s u s c e p t i b l e t o p i r a c y f r o m o t h e r w a t e r -s h e d s . P o s s i b l y S. malma r e a c h e d t h e e a s t s l o p e o f t h e R o c k y M o u n t a i n s b y t h i s means. The anadromous h a b i t a c c o u n t s f o r t h e w i d e c o a s t a l d i s t r i b u t i o n o f S. malma. V. SUMMARY (1) S. malma and S. a l p i n u s o c c u r s y m p a t r i c a l l y w i t h l i t t l e o r no e v i d e n c e o f h y b r i d i z a t i o n . T h e r e f o r e , S. malma s h o u l d be r e g a r d e d as a d i s t i n c t s p e c i e s and n o t a s u b -s p e c i e s o f S. a l p i n u s . (2) S. malma has f e w e r g i l l r a k e r s , p y l o r i c c a e c a and p o r e s a l o n g t h e l a t e r a l l i n e t h a n S. a l p i n u s . G i l l r a k e r s , p y l o r i c c a e c a and p o r e s a l o n g t h e l a t e r a l l i n e a r e n o t c o r -r e l a t e d w i t h l e n g t h i n S. malma. (3) R e l a t i v e g r o w t h s t u d i e s on S. malma and S, a l p i n u s r e v e a l d i f f e r e n c e s i n t h e g r o w t h o f v a r i o u s body p a r t s r e l a t i v e t o f o r k l e n g t h and head l e n g t h . None o f t h e s e d i f -f e r e n c e s a r e l a r g e enough t o c h a r a c t e r i z e e i t h e r s p e c i e s . (4) T h e r e i s c o n s i d e r a b l e g e o g r a p h i c v a r i a t i o n i n S. malma. The v a r i a t i o n shows no c o r r e l a t i o n w i t h l a t i t u d e . (5) G e o g r a p h i c v a r i a t i o n i n S. malma i s p r o b a b l y due t o b o t h e n v i r o n m e n t a l and g e n e t i c d i f f e r e n c e s b etween p o p -u l a t i o n s . (6) T h e r e i s no e v i d e n c e t o s u p p o r t s u b s p e c i f i c s t a t u s f o r any p o p u l a t i o n o f S. malma. S. m. s p e c t a b i l i s and S. m. k r a s h e n i n n i k o v i a r e n o t v a l i d s u b s p e c i e s . 66 (7) S. p l u v i u s i s a g e o g r a p h i c f o r m o f S. malma. Whether S. p l u v i u s s h o u l d be r e g a r d e d as a s e p a r a t e s u b s p e c i e s o f S. malma c a n n o t be d e c i d e d on t h e a v a i l a b l e e v i d e n c e . (8) S. malma and S. a l p i n u s d i v e r g e d sometime i n P l e i s t o c e n e . S. malma e v o l v e d i n t h e N o r t h P a c i f i c a r e a . (9) S,. malma s u r v i v e d t h e l a s t P l e i s t o c e n e g l a c i -a t i o n i n N o r t h A m e r i c a i n two a r e a s , t h e Yukon V a l l e y and s o u t h o f t h e i c e s h e e t . The p r e s e n t N o r t h A m e r i c a n d i s t r i b u t i o n o f S. malma has b e e n a t t a i n e d by b o t h f r e s h w a t e r and m a r i n e d i s -p e r s i o n f r o m t h e s e r e f u g i a . 67 BIBLIOGRAPHY A o y a g i , H. 1957. G e n e r a l N o t e s on t h e F r e s h w a t e r F i s h e s o f t h e J a p a n e s e A r c h i p e l a g o . D a i s y u - K a n , Tokyo, 272 pp., i n J a p a n e s e . B a c k u s , R. H. 1957. The F i s h e s o f L a b r a d o r . B u l l . Am. Mus. N a t . , H i s t . , V o l . 113(4), PP. 273-338. B e r g , L . S. 194$. Ryby presmykh v o d S S S R i s o p r e d e l n y k h s t r a n . E d i t i o n 4. Z o o l . I n s t . Akad. Nauk SSSR, No. 27, pp. 1-466, I n R u s s i a n . B u s s , K., and W r i g h t , J . E . 1956. R e s u l t s o f s p e c i e s h y b r i d i z -a t i o n w i t h i n t h e f a m i l y S a l m o n i d a e . P r o g . F i s h -c u l t . , 18(4), pp. 149-157. C a i n , A. J . 1954. A n i m a l S p e c i e s and T h e i r E v o l u t i o n . H u t c h i n s o n ' s U n i v e r s i t y L i b r a r y , L o n d o n , 190 pp. C a r l , C. G., and Clemens, W. A. 1953. The f r e s h - w a t e r f i s h e s o f B r i t i s h C o l u m b i a . 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S y s t e m a t i c P o s i t i o n o f t h e L a k e T r o u t , S a l v e l i n u s namaycush. C o p e i a , No. 2} pp. 116-124. Neave, F. 1958. The O r i g i n and S p e c i a t i o n o f O n c o r h y n c h u s . T r a n s . Roy. Coc. Canada, V o l . L I T , S e r . I l l , pp. 2 5 -39. Okada, Y. 1955. F i s h e s o f J a p a n . M a r a z e n Co., Takyo, 434 P a g e s . P r e b l e , E. A. 1908. F i s h e s o f t h e A t h a b a s k a - M a c k e n z i e r e g i o n . N o r t h Amer. F a u n a , No. 2 7 , pp. 502-515. Rao, C. R. 1952. A d v a n c e d s t a t i s t i c a l methods i n b i o m e t r i c r e s e a r c h . J o h n W i l e y and Sons, Y. Y., pp. 286-322. S t e n t o n , J . E . 1952. A d d i t i o n a l i n f o r m a t i o n on e a s t e r n b r o o k t r o u t X l a k e t r o u t h y b r i d s . Can. F i s h . C u l t . , 13, pp. 15-21. T a r a n e t z , A. F r e s h w a t e r f i s h e s o f t h e b a s i n o f t h e n o r t h -w e s t e r n p a r t o f t h e J a p a n S e a . T r a n . I n s t . Z o o l . A c a d . S c i . U. S.S.R., I V , pp. 483-540. V l a d y k o v , V. 1954. Taxonomic C h a r a c t e r s o f t h e E a s t e r n N o r t h A m e r i c a C h a r s ( S a l v e l i n u s and C r i s t i v o m e r ) . J . F i s h . Res. Bd. Canada, 11(6), pp. 904-932. W a l t e r s , V. 1955. F i s h e s o f w e s t e r n a r c t i c A m e r i c a and E a s t e r n A r c t i c S i b e r i a . B u l l . Am. Mus. N a t . H i s t . , v o l , 106 (5), PP. 255-368. W e i s e l . G. F. 1957. F i s h G u i d e f o r I n t e r m o u n t a i n Montana, Montana S t a t e U n i v . P r e s s , M i s s o u l a , 88 pp. W a l e s , J . H. 1957. T r o u t o f C a l i f o r n i a . D e p t . o f F i s h and Game, S a c r a m e n t o , 55 pp. Wil i m o w s k y , N. J . 1958. P r o v i s i o n a l Keys t o t h e F i s h e s o f A l a s k a . F i s h e r i e s R e s e a r c h L a b o r a t o r y , U. S. F i s h and W i l d l i f e S e r v i c e , J u n e a u , 113 pp. 7 0 APPENDIX I A l i s t o f S. Malma examined d u r i n g t h e p r e s e n t s t u d y . '[Numbers p r e c e d i n g l o c a l i t i e s a r e t h o s e shown i n F i g . 2 9 and r e f e r r e d t o i n t h e t e x t . The l e t t e r s BC p r e c e d i n g c o l l e c t i o n numbers r e f e r t o s p e c i m e n s 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 , U.B.C. 1. D r y C r e e k . 144 42' W., 63 41'N. 2. B r o o k s L a k e . 156\u00C2\u00B0 00' W., 58\u00C2\u00B0 2 2 ' N. BC 53-414. 3 . T u r n a g a i n Arm. 149\u00C2\u00B0 13' W., 60\u00C2\u00B0 56' N. BC 53-200. 4. Summit L a k e . 149\u00C2\u00B0 31' N., 60\u00C2\u00B0 56' N. BC 53 - 2 0 3 . 5. Chenan L a k e . 144\u00C2\u00B0 26' W., 61\u00C2\u00B0 3 2 ' N. BC 53 - 2 2 7 . 6. S e l t a t C r e e k . 136\u00C2\u00B0 27' W., 59\u00C2\u00B0 36' N. BC 57-236. 7. T a k u R i v e r . 133\u00C2\u00B0 51' W., 53\u00C2\u00B0 32' N. BC 53-337. 3. S m a l l pond n e a r Midarm L a k e . 152\u00C2\u00B0 25' W., 53\u00C2\u00B0 11' N. BC 53-210. 9. K a r l u k L a k e . 154\u00C2\u00B0 05' W., 57\u00C2\u00B0 2 3 ' N. BC 53-219. 10. F r a s e r L a k e . 154\u00C2\u00B0 09' W., 57\u00C2\u00B0 15' N. BC 53-216. 71 11. Bowser L a k e . 130\u00C2\u00B0 00' W., 56\u00C2\u00B0 30' N. BC 56-537. 12. P a r s n i p R i v e r . 123\u00C2\u00B0 00*. W., 55\u00C2\u00B0 00' N. BC 56-457. * 13. Ocean F a l l s . 127\u00C2\u00B0 40' W., 52\u00C2\u00B0 20' N. BC 58-305. 14. B e l l a C o o l a R i v e r . 126\u00C2\u00B0 40' W., 52\u00C2\u00B0 15' N. BC 56-480. 15. Cottonwood R i v e r . 122o 20' W., 53\u00C2\u00B0 10' N. BC 56-448, 462, 463, 515, 561, 5$2. 16. C h e h a l i s L a k e . 122\u00C2\u00B0 00' W., 49\u00C2\u00B0 30' N. BC 59-131. 17. S u r a a l l o R i v e r . 121\u00C2\u00B0 20' W., 49\u00C2\u00B0 20' N. BC 55-316. 18\". Sheep C r e e k . 117\u00C2\u00B0 30' W., 49\u00C2\u00B0 10' N. BC 55-269. 19. Sage C r e e k . 114\u00C2\u00B0 30' W., 49\u00C2\u00B0 10' N. BC 57-327. 20. C o q u i h a l l a R i v e r . 121\u00C2\u00B0 30' W., 49\u00C2\u00B0 20' N. BC 59-29. 21. C u t h e a d L a k e . 115\u00C2\u00B0 40' W., 51\u00C2\u00B0 20' N. BC 59-169. 22. K e n a i R i v e r 151\u00C2\u00B0 24' W., 600 28' N. BC 5$-222. 23. G l a c i e r N a t i o n a l P a r k . 113\u00C2\u00B0 30' W., 48\u00C2\u00B0 30' N. 2341, 3002, 334$, ( S p e c i m e n s l o a n e d f r o m t h e U n i v e r s i t y o f Washington). Upper T r a i l L a k e . 149\u00C2\u00B0 24T W., 60\u00C2\u00B0 31' N. BC 58-202. K e l s a l l L a k e . 136\u00C2\u00B0 30' W., 59\u00C2\u00B0 50' N. BC 57-238. S t u h i n n i C r e e k . 133\u00C2\u00B0 32' W., 58\u00C2\u00B0 37' N. BC 58-376. P i t t L a k e . 122\u00C2\u00B0 30' W., 49\u00C2\u00B0 20' N. BC 57-274, 277. Swan L a k e . 131\u00C2\u00B0 24' W., 59\u00C2\u00B0 53' N. BC 57-226. Homathko R i v e r . 1 2 5 \u00C2\u00B0 00' W., 59\u00C2\u00B0 10' N. BC 56-485. P e a c e R i v e r . 120\u00C2\u00B0 30' W., 56\u00C2\u00B0 10' N. BC 58-171. L i t t l e K i t o i L a k e . 152\u00C2\u00B0 23' W., 58\u00C2\u00B0 12' N. BC 58-211. Takhanne R i v e r . 136\u00C2\u00B0 56' W., 60\u00C2\u00B0 0 6 ' N. BC 57-240. D e z a d e a s h L a k e . 137\u00C2\u00B0 00' W., 60\u00C2\u00B0 27' N. BC 57-234. S k i l a k L a k e . 150\u00C2\u00B0 30' W.,60\u00C2\u00B0 40' N. BC 58-223. G o l d C r e e k . Q 117 W., 49 N. BC 57-398. R i v e r s I n l e t 121\u00C2\u00B0 30' \u00C2\u00A5., 51\u00C2\u00B0 30' N. BC 57-52. K l u k s h u L a k e . 137 00' V/ . , 60 17>' N. BC 57-242. 73 APPENDIX I I P r o c e d u r e u s e d i n c a l c u l a t i n g d i s c r i m i n a n t s c o r e s . F o r any i n d i v i d u a l t h e d i s c r i m i n a n t s c o r e n D \" i s ; A/ V A, V A . V \ X 4 = D T> , X^, X^ and X^ _ a r e c h a r a c t e r s 1,2,3, and 4 f o r any i n d i v i d u a l , and A,, A 2 , A^, and A^ a r e c o e f f i c i e n t s o b t a i n e d b y s o l v i n g t h e f o l l o w i n g e q u a t i o n s ; *, % + S,2 + A* X\u00E2\u0080\u009E - X /2 K S2, + A, + A3 SZ3 + A4 = -A, + A 2 + A3 S\u00C2\u00BB * A 4 V = K S4/ A Z A3 + A+ - X ^ z Sy/, S , e t c . i s t h e c o v a r i a n c e o f t h e p a r t i c u l a r c o m b i n a t i o n o f c h a r a c t e r s : X//f X,, ..etc. i s t h e mean o f c h a r a c t e r 1 i n p o p u l a t i o n 1, c h a r a c t e r 1 i n p o p u l a t i o n 2 e t c . "@en . "Thesis/Dissertation"@en . "10.14288/1.0106102"@en . "eng"@en . "Zoology"@en . "Vancouver : University of British Columbia Library"@en . "University of British Columbia"@en . "For non-commercial purposes only, such as research, private study and education. Additional conditions apply, see Terms of Use https://open.library.ubc.ca/terms_of_use."@en . "Graduate"@en . "A systematic study of the Dolly Varden, Salvelinus malma (Walbaum)"@en . "Text"@en . "http://hdl.handle.net/2429/40050"@en .