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Moult and pelage patterns in the Pacific harbour seal Phoca vitulina Stutz, Sherwood S. 1966

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MOULT AND PELAGE PATTERNS IN THE PACIFIC HARBOUR SEAL PHOCA VITULINA by SHERWOOD S. STUTZ B.Sc, Humboldt State College, 1963 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department of Zoology We accept t h i s thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA A p r i l , 1966 In presenting 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 for 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 for reference and study. I further agree that per-mission for extensive copying of t h i s t h e s i s for s c h o l a r l y purposes may be granted by the Head of my Department or by h i s representatives,, 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 for 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 A p r i l 29, 1966 ABSTRACT Phoca v i t u l i n a uses a v a r i e t y o f s h a l l o w c o a s t a l and f r e s h w a t e r h a b i t a t s , i s n o n - m i g r a t o r y and r a r e l y found more t h a n f i f t e e n m i l e s from s h o r e . The s e a l p o s s e s s e s a complete mammalian h a i r system ( l e s s t h e e r e c t o r p i l i muscle) c o n s i s t i n g of guard h a i r s , u n d e r h a i r s , sebaceous and sweat g l a n d s . The h a i r p r o v i d e s p r i m a r i l y m e c h a n i c a l p r o t e c t i o n f o r t h e s k i n r a t h e r t h a n i n s u l a t i o n a g a i n s t heat l o s s . A f o e t a l moult, n o r m a l l y i n t r a u t e r i n e (except i n P. v. l a r g h a , w h i c h may not be a harbour s e a l ) d i f f e r s i n p a t t e r n sequence from t h e a d u l t moult. The a d u l t moult o c c u r s d u r i n g August - October f o r most specimens. There i s some moult a c t i v i t y among a d u l t s d u r i n g almost e v e r y month of t h e y e a r . D i s t r i b u t i o n of two c o l o u r p a t t e r n s , b l a c k and common, i n d i c a t e l o c a t i o n s of d i s c r e t e p o p u l a t i o n s whose e x i s t e n c e has management i m p l i c a t i o n s . Phoca v i t u l i n a . Mother and pup, August, 1962. (Photograph c o u r t e s y o f P r o f e s -s o r R.J. H a r r i s o n , London H o s p i t a l M e d i c a l C o l l e g e . ) i i i TABLE OF CONTENTS Page INTRODUCTION 1 The Study 1 The Seal, General Information 2 METHODS AND MATERIALS 6 Skin Sampling, Sources and Limitations . . . . 6 Preparation of Specimens and Data 7 MOULT PROGRESSION 9 Morphology of the Seal Skin 9 The Pre-Natal and Neo-Natal Pelage 18 Moult of the Adult Pelage 25 PELAGE PATTERNS AND COLOURS 36 Description 36 Colour Pattern - Size Relationships i n Commercial Skins 38 The Geographical D i s t r i b u t i o n of Colour and Pattern 39 Foreign Growths and Stains of Hair 41 DISCUSSION 4 5 RECOMMENDATIONS 56 SUMMARY 5 9 ACKNOWLEDGEMENTS LITERATURE CITED APPENDIX V LIST OF TABLES Table Page I. Moult progression by months, percentages 27 II. Pelage types of harbour seal, Phoca v i t u l i n a , skins c o l l e c t e d on the P a c i f i c Coast 37 III. Comparison of samples 38 IV. D i s t r i b u t i o n of pelage types by area 40 Appendix A. Horizontal sections through f o l -l i c l e groups, with hair measure-ments, males 65 B. Horizontal sections through f o l -l i c l e groups, with hair measure-ments, females . . 66 C. Dates of moult stages, males 67 D. Dates of moult stages, females 70 E. Dates of moult stages, sex undetermined 74 v i LIST OF FIGURES F i g u r e Page 1. Habour s e a l (Phoca v i t u l i n a ) sub-s p e c i e s i n t h e P a c i i i c ocean. Phoca i n s u l a r i s i s s y m p a t r i c w i t h P. v. l a r g h a I n t h e K u r i l e I s l a n d s , Fetween Japan and Kamchatka 3 2. O v e r h a i r and u n d e r h a i r f o l l i c l e s , 20X. Note t h e s i n g l e p i l a r y c a n a l i n w hich a l l h a i r s j o i n b e f o r e emerging from t h e s k i n s u r f a c e 11 3. F o l l i c l e groups ( h a i r b u n d l e s ) and sebaceous g l a n d s , 80X. Note t h e f o l l i c l e group, i n t h e c e n t e r of t h e p i c t u r e , w hich shows an o v e r h a i r , f o u r u n d e r h a i r s , and a horseshoe shaped group o f sebaceous g l a n d s 12 4. C l u b h a i r f o l l i c l e , 52X. Note c l u b h a i r , t e l o g e n b u l b and p a p i l l a remnant, c r o s s s e c t i o n s o f c o n v o l u -t e d sweat g l a n d 14 5. Anagen, e a r l y , 20X. Note t h e o v e r -h a i r , e x t e n d i n g from t h e anagen b u l b i n t h e lower r i g h t , w hich i s growing a l o n g s i d e and p a s t a c l u b h a i r 17 6. Phoca v i t u l i n a r i c h a r d i female pup c a p t u r e d at P o r t Moody, B.C., on 10 August, 1965. Weight 17 l b s . U m b i l i -cus 3| i n c h e s l o n g , p i n k r e d . F o e t a l (lanugo) h a i r coat prominent except on t o p of head, f a c e , and f o r e f l i p -p e r s . Photographed at s e r v i c e y a r d , S t a n l e y P a r k , Vancouver, B.C., 13 August, 1965 20 7. Tanned s k i n of Phoca v i t u l i n a l a r g h a pup from B e r i n g Sea. Depth o f t h e w h i t e wooly c o a t 30 mm. Note t h e s h o r t , s p o t t e d a d u l t t y p e coat v i s i b l e i n t h e t h r e e s m a l l p l u c k e d areas . . . . . 21 F i g u r e v i i Page 8. Tanned s k i n s l i c e s viewed i n c r o s s s e c t i o n . M a n g i f i c a t i o n about 3X. a. Phoca v i t u l i n a r i c h a r d i f o e t u s . b. Phoca v i t u l i n a l a r g h a w h i t e -coat pup 22 9. The f o e t a l moult p a t t e r n i n P. v. r i c h a r d i . . . 7 . — 24 10. H i s t o g r a m of t h e annual moult i n Phoca v i t u l i n a r i c h a r d i 28 11. T y p i c a l moult p r o g r e s s i o n i n t h e harbour s e a l of B r i t i s h Columbia. a. E a r l y moult b. A l a t e r s t a g e of moult. .'. 31 12. Three year o l d female common s e a l , Phoca v i t u l i n a v i t u l i n a photographed November, 1960. Compare the spot p a t t e r n v i s i b l e a n t e r i o r t o t h e l e f t f r o n t f l i p p e r w i t h F i g u r e 13. 34 13. F i v e year o l d female P. v. v i t u l i n a photographed August, T96TJ. T h i s i s t h e same s e a l i l l u s t r a t e d i n F i g u r e 12. Note t h e unchanged spot p a t -t e r n a n t e r i o r t o t h e l e f t ' f r o n t f l i p p e r 35 14. S k e t c h o f a l g a e e n c r u s t e d p e l t of female harbour s e a l c o l l e c t e d 19 September, 1965, at Westham I s l a n d , mouth of t h e F r a s e r R i v e r , B r i t i s h C olumbia. The l i n e d a r e a was t h i c k l y e n c r u s t e d w i t h P l e u r o c a p s a sp. T h i s specimen was i n a c t i v e moult and o b v i o u s l y s h e d d i n g i t s " o l d " h a i r 42 15. The B r i t i s h Columbia and s o u t h -e a s t e r n A l a s k a c o a s t s . S k e t c h map of key l o c a t i o n s 49 16. B l a c k s k i n . Male c o l l e c t e d near S k i d e g a t e , Queen C h a r l o t t e I s l a n d s , l a t e A p r i l , 1965 50 v i i i F i g u r e Page 17. Common s k i n . Male c o l l e c t e d from Puget Sound, e a r l y May, 1965 51 18. Muddy s k i n . Female from t h e Queen C h a r l o t t e I s l a n d s . C o l l e c t e d l a t e J u l y , 1964 52 ERRATA SHEET MOULT AND PELAGE PATTERNS IN THE PACIFIC HARBOUR SEAL PHOCA VITULINA Page 18. Line 8 reads: yllowish, should read; yellowish. Page 19. Top of page reads: tanned skin samples from a P. v. largha whitecoat foetus from B r i t i s h Colum-bia, should read: tanned skin samples from a P. v. r i c h a r d i whitecoat foetus from B r i t i s h Colum-bi a and a P. v. largha whitecoat pup from Bering Sea Page 60. Line 14 reads; Bennot, should read: Bohnot. Page 63. Add to top of page: Scheffer, V.B. 1962. Pelage and surface topogra phy of the northern fur s e a l . North American Fauna No. 64. U.S. Fish and W i l d l . Svc., Wash. D.C. 206 p. 1 INTRODUCTION THE STUDY Recent changes i n t h e economic v a l u e of t h e harbour s e a l , Phoca v i t u l i n a r i c h a r d i , of t h e P a c i f i c c o a s t of N o r t h America have i n c r e a s e d i n t e r e s t i n e f f e c t i v e economic manage-ment of t h e s p e c i e s . Development of a c o m p a r a t i v e l y h i g h economic v a l u e at t h e end of 1962 f o l l o w e d a p e r i o d of many ye a r s d u r i n g which t h e harbour s e a l was r e g a r d e d as a p u r e l y n u i s a n c e a n i m a l . A bounty was p a i d by t h e Canada Department of F i s h e r i e s t o encourage d e s t r u c t i o n of s e a l s . F o l l o w i n g a r i s e i n t h e p r i c e p a i d f o r raw s k i n s t o a h i g h average of $50.00 ( S t u t z , 1965) t h e bounty was removed e f f e c t i v e J u l y 31, 1964. T h i s s t u d y was begun, w i t h t h e support of t h e F i s h -e r i e s R e s e a r c h Board of Canada, i n 1964 t o o b t a i n some of t h e b i o l o g i c a l i n f o r m a t i o n needed f o r management of t h e r e s o u r c e . My f i e l d of i n v e s t i g a t i o n i n c l u d e d a g e n e r a l s t u d y o f t h e s e a l w i t h p a r t i c u l a r a t t e n t i o n t o t h e p e l t of t h e s p e c i e s and i t s s e a s o n a l and r e g i o n a l v a r i a t i o n s . F i e l d work was conducted i n A p r i l - J u l y , 1964, i n t h e lower F r a s e r V a l l e y , on t h e Skeena and E c s t a l l R i v e r s , and at R i v e r s I n l e t and Owikeno Lake. The c o o p e r a t i o n of a f u r buyer at L a n g l e y , B.C., Mr. H. Hansen, made a v e r y l a r g e number of p e l t s c o n t i n u o u s l y a v a i l -a b l e t o me at h i s p l a n t . H i s t o l o g i c a l work was conducted w i t h 2 t h e f a c i l i t i e s of t h e Department of Zoology, U n i v e r s i t y of B r i t i s h C olumbia. THE SEAL, GENERAL INFORMATION D i s t r i b u t i o n o f Phoca v i t u l i n a i n t h e P a c i f i c Ocean ( F i g u r e 1) extends from Japan and v i c i n i t y (Schwarz, 1942; N i s h i w a k i and N a g a s a k i , 1960; Imaizumi, 1961) northward a l o n g the A s i a t i c c o a s t t o t h e B e r i n g Sea (Ognev, 1935), t h e A l e u -t i a n I s l a n d s ( M u r i e , 1959) and from B e r i n g S t r a i t s o u t h a l o n g the N o r t h American c o a s t t o B a j a C a l i f o r n i a ( S c h e f f e r , 1958). Phoca v i t u l i n a r i c h a r d i , t h e harbour s e a l of t h e American shore of t h e P a c i f i c Ocean resembles P. v. v i t u l i n a of Europe and P. v. c o n c o l o r of t h e w e s t e r n A t l a n t i c i n i t s avoidance of i c e (Dunbar, 1949) and use of t i d a l a r e a s as pup-p i n g s i t e s ( S c h e f f e r and S l i p p , 1944; M a n s f i e l d , 1963; K i n g , 1964). Phoca v i t u l i n a l a r g h a o f n o r t h e a s t e r n A s i a (Ognev, 1935; Schwarz, 1942) i s m o r p h o l o g i c a l l y so s i m i l a r t o P. v. r i c h a r d i ( S c h e f f e r , 1958; K i n g , 1964) t h a t Ognev (1935) d e c l i n e d t o s e p a r a t e t h e American form, t a x o n o m i c a l l y , from l a r g h a . P. v. l a r g h a i s , however, r e p r o d u c t i v e l y i s o l a t e d from o t h e r s u b s p e c i e s of Phoca v i t u l i n a ( S t u t z , 1966) and i s s y m p a t r i c w i t h a form, Phoca i n s u l a r i s , d e s c r i b e d by B e l k i n (1964) which may be a s u b s p e c i e s of P. v i t u l i n a ( S t u t z , 1966). Harbour s e a l s t e n d t o remain near shore and are non F i g u r e 1. Harbour s e a l (Phoca v i t u l i n a ) s u b s p e c i e s i n t h e P a c i f i c Ocean. Phoca i n s u l a r i s i s s y m p a t r i c w i t h P. v. l a r g h a i n t h e K u r i l e I s l a n d s , between Japan and Kamchatka. 4 m i g r a t o r y (Ognev, 1935; S c h e f f e r and S l i p p , 1944; S p a l d i n g , 1964) but e x h i b i t p r e f e r e n c e s f o r areas of d i f f e r i n g c h a r a c -t e r i s t i c s w i t h i n t h e i r home ranges d u r i n g d i f f e r e n t seasons ( S p a l d i n g , 1964). F i s h e r (1952) r e p o r t e d a l o c a l upstream movement i n t h e Skeena R i v e r d u r i n g t h e e u l a c h o n , T h a l e i c h t h y s  p a c i f i c u s , and Chinook salmon, Oncorhynchus t s h a w y t s c h a , spawn-i n g r u n s . T h i s appeared t o be a movement o f i n d i v i d u a l s f o l -l o w i n g a f o o d s o u r c e and not a mass m i g r a t i o n . In June, 1964 I obse r v e d t h e use of sandbars on t h e Skeeria and E c s t a l l R i v e r s by females w i t h newborn pups. F i s h e r (1952) had obs e r v e d newborn s e a l s on t h e same b a r s e i g h t e e n y e a r s e a r l i e r . Phoca v i t u l i n a uses a v a r i e t y o f a q u a t i c h a b i t a t s . On t h e C a l i f o r n i a c o a s t I have noted harbour s e a l s i n s m a l l groups i n t i d e p o o l s and among t h e k e l p beds of r o c k y s h o r e -l i n e s f e a t u r i n g heavy s u r f . A few m i l e s away members of t h e same s p e c i e s and p o s s i b l y of t h e same p o p u l a t i o n were l o c a t e d i n s h e l t e r e d bays and i n s h a l l o w water o f f sandy beaches. Murie (1959) r e p o r t e d s i m i l a r use o f k e l p beds and sandy areas by harbour s e a l s i n w e s t e r n A l a s k a . The harbour s e a l f r e q u e n t l y e n t e r s f r e s h water streams and l a k e s ( K i n g , 1964; M a n s f i e l d , 1963; F i s h e r , 1952; Dunbar, 1949; Ognev, 1935) and one s u b s p e c i e s , Phoca v i t u l i n a  m e l l o n a e , i s e n t i r e l y r e s t r i c t e d t o f r e s h water ( S c h e f f e r , 1958). I n B r i t i s h Columbia harbour s e a l s u t i l i z e f r e s h w a t e r l a k e s t h a t are r e a d i l y a c c e s s i b l e from t h e s e a . F i s h e r i e s 5 O f f i c e r s , Mr. V. G i r a u d and Mr. M. Bussey, d e s c r i b e d a s m a l l p o p u l a t i o n s e a s o n a l l y p r e s e n t i n L a k e l s e Lake over 90 m i l e s upstream from t h e Skeena R i v e r mouth. In t h e F r a s e r d r a i n a g e two harbour s e a l s were shot on H a r r i s o n Lake i n t h e f a l l of 1965 by Mr. R. F a u l k n e r of Deroche, B.C., and s e a l s have been seen i n L i l l o o e t Lake by Mr. T. G j u r n e s s of t h e Salmon Commis-s i o n (DeBock, p e r s o n a l communication). I e s t i m a t e d a p o p u l a -t i o n of 30 s e a l s on Owikeno Lake i n J u l y , 1964. P e r s o n n e l of t h e F i s h C u l t u r e B r a n c h , Canada Department of F i s h e r i e s , s t a t e d t h a t t h e y had seen s e a l s i n Owikeno Lake d u r i n g e v e r y month of t h e y e a r . I noted at l e a s t one a d u l t - pup p a i r a p p a r e n t l y r e s i d e n t about 30 m i l e s from s a l t w a t e r , i n d i c a t i n g a p r o b a b i l -i t y t h a t t h e pup had been b o r n at Owikeno Lake. 6 METHODS AND MATERIALS SKIN SAMPLING, SOURCES AND LIMITATIONS The s o u r c e of most s k i n samples was a c ommercial f u r b uyer's p l a n t where a v e r y l a r g e volume of s k i n s from most areas of t h e c o a s t between w e s t e r n A l a s k a and Washington S t a t e were made a v a i l a b l e . S k i n samples were removed from h i d e s f o r which th e approximate g e o g r a p h i c a l o r i g i n s and d a t e s of c o l -l e c t i o n by t h e hunter were known. Two samples were t a k e n from each s k i n . Sample "A" was cut from th e v e n t r a l m i d l i n e between t h e p e c t o r a l f l i p p e r s . Sample "B" was t a k e n from th e d o r s a l m i d l i n e at t h e a n t e r i o r end of t h e p e l t , a l o c a t i o n which v a r i e d c o n s i d e r a b l y between p e l t s but was u s u a l l y t h e p o s t e r i o r p o r t i o n of t h e head. Most h i d e s sampled were a l s o photographed t o r e c o r d t h e p e l a g e t y p e . E x c e p t i n g o b v i o u s pup h i d e s no method of a g i n g s k i n s by e x a m i n a t i o n has been de v e l o p e d . The r e d u c t i o n of h a i r d e n s i t y r e p o r t e d by S c h e f f e r (1964) i s a f u n c t i o n of s i z e and i s not w e l l c o r r e l a t e d w i t h t h e age o f t h e specimen. S k i n s have t h e r e f o r e been c l a s s i f i e d as pups or as a d u l t s . S e x i n g of many specimens has been q u e s t i o n a b l e . On some h i d e s t h e p e n i a l o p ening remains v i s i b l e , i n s u r i n g i d e n t i -f i c a t i o n as a male. In o t h e r s , i n c l u d i n g n u r s i n g f e m a l e s , t h e n i p p l e s a r e l a r g e and prominent. More u s u a l l y t h e r e i s n e i t h e r a p e n i a l o p ening nor prominent n i p p l e s and s e x i n g has been 7 based on t h e s i z e of t h e n i p p l e s and t h e presence or absence of neck s c a r s ( c o n s i d e r e d a c h a r a c t e r i s t i c of a mature ma l e ) . These methods are not c o n s i d e r e d r e l i a b l e . Some s k i n samples were a bypr o d u c t of a r e p r o d u c t i v e c y c l e / l i f e h i s t o r y s t u d y c o nducted by Mr. M i c h a e l B i g g . These were i n i t i a l l y accompanied by t h e r e p r o d u c t i v e t r a c t s and lower jaws of t h e specimens. A c c u r a t e s e x i n g f rom t h e t r a c t s and a g i n g t h r o u g h s t u d y of c a n i n e t o o t h s e c t i o n s by t h e method o u t l i n e d by M a n s f i e l d and F i s h e r (1960) were a c c o m p l i s h e d by Mr. B i g g . PREPARATION OF SPECIMENS AND DATA Samples f o r h i s t o l o g i c a l s t u d y were f i x e d i n f o r m a l i n u n t i l r eady f o r p r e p a r a t i o n . T i s s u e samples were r i n s e d , d e h y d r a t e d , embedded i n parawax, s e c t i o n e d at 8 m i c r o n s , mounted i n s e r i a l s e c t i o n s , and s t a i n e d w i t h h a e m a t o x y l i n and e o s i n . To r e v e a l t h e e x t e n t of h a i r b u l b development and r e g r e s s i o n , samples from 261 s e a l s were s e c t i o n e d v e r t i c a l l y ( p a r a l l e l t o t h e h a i r f o l l i c l e s ) . A d d i t i o n a l samples were s e c t i o n e d h o r i z o n t a l l y f o r s t u d y of f o l l i c l e groups and measure-ment of i n d i v i d u a l h a i r d i a m e t e r s . Measurements were made w i t h an o c u l a r micrometer which was c a l i b r a t e d w i t h a s t a g e micrometer b e f o r e t h e r e a d i n g s were r e c o r d e d i n f i n a l form. 8 Comparison of w h i t e c o a t p e l a g e of Phoca v i t u l i n a r i c h a r d i w i t h t h e w h i t e c o a t of P. v. l a r g h a was made t h r o u g h e x a m i n a t i o n of c r o s s s e c t i o n a l s l i c e s of t a nned s k i n c u t w i t h a r a z o r b l a d e . Photography and p e r s o n a l s c r u t i n y of s e a l p e l t s and c a r c a s s e s p r o v i d e d t h e b a s i s f o r d e s c r i p t i o n s of p a t t e r n s of moult p r o g r e s s i o n f o r t h e f o e t a l and p o s t - n a t a l p e l a g e s of P. v. r i c h a r d i . 9 MOULT PROGRESSION MORPHOLOGY OF THE SEAL SKIN The h i s t o l o g y and h i s t o c h e m i s t r y of t h e s k i n of Phoca v i t u l i n a v i t u l i n a i n England were i n v e s t i g a t e d by Mon-t a g n a and H a r r i s o n (1957). Over t h e e n t i r e body t h e e p i d e r m i s measured 1/2 t o 1 mm i n t h i c k n e s s . S okolov (1959, 1960a) r e p o r t e d t h a t t h e e p i d e r m i s of a "young male" P. v. l a r g h a from t h e K u r i l e I s l a n d s averaged 106 m i c r a i n t h i c k n e s s . The t h i c k e s t r e g i o n , 150 m i c r a , was on t h e c h e s t between t h e f o r e -f l i p p e r s . Montagna and H a r r i s o n (1957) noted t h a t no s t r a t u m granulosum was p r e s e n t and t h a t t h e s t r a t u m corneum was incom-p l e t e l y k e r a t i n i z e d . They b e l i e v e d t h a t t h i s i n c o m p l e t e k e r a -t i n i z a t i o n was an a q u a t i c a d a p t a t i o n s i n c e a " s c a l y " k e r a t i n s u r f a c e , t y p i c a l of p u r e l y t e r r e s t r i a l mammals, might be sub-j e c t t o w a t e r l o g g i n g and e x c e s s i v e f l a k i n g . S i m i l a r l y Palmer and W a d d e l l (1964) noted t h a t t h e e p i d e r m i s of t h e b o t t l e nose d o l p h i n , T u r s l o p s t r u n c a t u s , had a v e r y t h i n k e r a t i n l a y e r . The s t a g e s of t h e h a i r c y c l e and t h e s t r u c t u r e and development o f t h e h a i r of Phoca v i t u l i n a a r e s i m i l a r t o t h o s e d e s c r i b e d f o r mammals of o t h e r o r d e r s (Montagna and H a r r i s o n , 1957; Montagna and V a n S c o t t , 1958; B u l l o u g h and Laurence, 1958). The q u i e s c e n t f o l l i c l e of t e l o g e n ( t h e " r e s t i n g " or c l u b h a i r phase) d e v e l o p s i n t o t h e a c t i v e h a i r p r o d u c i n g s t r u c t u r e of anagen (phase of growth and development). When 10 h a i r c e a s e s t o grow, t h e h i g h l y d e v e l o p e d h a i r b u l b s and dermal p a p i l l a e of anagen q u i c k l y r e g r e s s i n c a t a g e n ( r e g r e s -s i o n p h a s e ) , t h e c l u b h a i r i s e s t a b l i s h e d and t h e f o l l i c l e e n t e r s t e l o g e n and remains q u i e s c e n t u n l e s s t h e h a i r i s p u l -l e d out (Montagna and H a r r i s o n , 1957) or u n t i l t h e a n n u a l c y c l e b e g i n s anew ( B u l l o u g h and Laurence, 1958). Harbour s e a l h a i r grows i n groups composed of one l a r g e b r i s t l e - l i k e o v e r h a i r (guard h a i r , p r i m a r y h a i r , a x i a l h a i r ) and a v a r y i n g number of s h o r t e r , f i n e r u n d e r h a i r s ( s e condary h a i r s , downy h a i r s , wool h a i r s ) a l l of w h i c h emerge t h r o u g h a s i n g l e s u r f a c e o p e n i n g ( F i g u r e 2) (Montagna and H a r r i s o n , 1957; S o k o l o v , 1959, 1960a; S c h e f f e r , 1964). As i n a l l o t h e r p i n n i p e d s t h e e r e c t o r p i l i muscle i s absent ( S o k o l o v , 1959, 1960a, 1960b; K i n g , 1964). Sokolov (1959, 1960a, 1960b) d e s c r i b e d P. v. l a r g h a f o l l i c l e groups as c o n t a i n i n g one a x i a l , one or two i n t e r m e d i a t e ( l a r g e u n d e r h a i r s ) and 3 - 5 wool h a i r s . He c o n s i d e r e d t h e number of u n d e r h a i r s t o be s p e c i e s s p e c i f i c and u s e f u l as a taxonomic c h a r a c t e r ( S o k o l o v , 1960a, 1960b). The o v e r h a i r f o l l i c l e s l a n t s p o s t e r i a d from t h e b u l b t o t h e s u r f a c e of t h e s k i n . U n d e r h a i r f o l l i c l e s a re l o c a t e d a d j a c e n t and p o s t e r i o r t o t h e guard h a i r f o l l i c l e (Montagna and H a r r i s o n , 1957; S o k o l o v , 1960a; S c h e f f e r , 1964). Sokolov (1960a, 1960b) commented t h a t u n l i k e O t a r i i d a e , i n Odobenidae and P h o c i d a e th e h a i r s l a c k e d a m e d u l l a . 11 F i g u r e 2. O v e r h a i r and u n d e r h a i r f o l l i c l e s , 20X. Note t h e s i n g l e p i l a r y c a n a l i n which a l l h a i r s j o i n b e f o r e emerging from t h e s k i n s u r f a c e . Figure 3 . F o l l i c l e groups (hair bundles) and sebaceous glands, 2 0 X . Note the f o l l i c l e group, in the center of the picture, which shows an overhair, four underhairs, and a horseshoe shaped group of sebaceous glands. 13 D e t a i l e d i n v e s t i g a t i o n of t h e sebaceous g l a n d s by Montagna and H a r r i s o n (1957) and Sokolov (1959, 1960a, 1960b) r e s u l t e d i n apparent disagreement as t o t h e i r numbers and r e l a t i o n s h i p t o t h e h a i r b undle ( f o l l i c l e group)> Sokolov (1959, 1960a, 1960b) s t a t e d t h a t i n P. v. l a r g h a t h e r e were two sebaceous g l a n d s l o c a t e d at each h a i r b u n d l e . One g l a n d i n t h e obtuse a n g l e between t h e f o l l i c l e and t h e e p i d e r m a l s u r f a c e opened i t s duct i n t o t h e guard h a i r f o l l i c l e . The l a r g e r sebaceous g l a n d , l o c a t e d p o s t e r i o r t o t h e h a i r b u n d l e , opened i n t o t h e upper ( c o n j o i n e d ) f o l l i c l e of t h e u n d e r h a i r s . T h i s was c o n s i d e r e d t y p i c a l of p i n n i p e d s i n g e n e r a l ( S o k o l o v , 1960a, 1960b). Montagna and H a r r i s o n (1957) r e p o r t e d t h a t P. v. v i t u l i n a had s e v e r a l c l u s t e r s of sebaceous g l a n d s u r r o u n d -i n g t h e f o l l i c l e group. Each c l u s t e r c o n t a i n e d a c l o s e l y packed group of g l a n d s . Each g l a n d opened s e p a r a t e l y i n t o t h e p i l a r y c a n a l by a l o n g , narrow, duct (Montagna and H a r r i -son, 1957). I f o u n d from h o r i z o n t a l l y s e c t i o n e d s k i n samples ( p a r a l l e l t o t h e e p i d e r m a l s u r f a c e ) t h a t i n P. v. r i c h a r d i t h e sebaceous g l a n d s formed a horseshoe shaped c o l l a r around t h e f o l l i c l e group w i t h t h e o p e n i n g of t h e "horseshoe" p o i n t e d p o s t e r i a d ( F i g u r e 3 ) . Montagna and H a r r i s o n (1957) n o t e d t h a t t h e sebaceous g l a n d s produced a l a r g e q u a n t i t y of o i l y sebum wh i c h c o v e r e d the s k i n and h a i r w i t h a w a t e r p r o o f f i l m . V e r t i c a l s e c t i o n s o f s e a l s k i n f r e q u e n t l y e x h i b i t s t r i n g s of c r o s s s e c t i o n s of t h e c o n v o l u t e d sweat g l a n d s F i g u r e 4. C l u b h a i r f o l l i c l e , 52X. b u l b and p a p i l l a remnant, c o n v o l u t e d sweat g l a n d . Note c l u b h a i r t e l o g e n c r o s s s e c t i o n s of 15 ( F i g u r e 4 ) . Montagna and H a r r i s o n (1957) f o u n d t h a t a s i n g l e sweat g l a n d e n t e r e d t h e p i l a r y c a n a l j u s t below t h e d u c t s of the sebaceous g l a n d s . They d e s c r i b e d sweat i n t h e harbour s e a l as y e l l o w i s h , semi s o l i d , compact, t w i s t e d c a s t s of m a t e r i a l w h i c h d i d not r e a d i l y mix w i t h sebaceous sebum. Montagna and H a r r i s o n (1957) f e l t t h a t t h e v i s c o u s , condensed s e c r e t i o n o f t h e sweat g l a n d s might be e x c r e t o r y i n n a t u r e , a waste p r o d u c t r e s p o n s i b l e f o r t h e c h a r a c t e r i s t i c odor of th e s e a l . K i n g (1964) commented t h a t t h e sweat g l a n d of p h o c i d s e n l a r g e d d u r i n g moult and b r e e d i n g p e r i o d s and might f u n c t i o n as s e x u a l s c e n t g l a n d s . S okolov (1960a) noted t h a t sweat g l a n d s were absent from h a i r l e s s a r e a s . In S c h e f f e r ' s (1964) m a t e r i a l t h e number of second-a r y h a i r s per group v a r i e d f rom 1 - 6 . H i s d a t a , based a p p a r e n t l y on a s t u d y of f i v e specimens, showed a r e d u c t i o n i n h a i r d e n s i t y , an i n c r e a s e i n t h e mean b a s a l d i a m e t e r of p r i m a r y h a i r s and a r i s e i n t h e number of u n d e r h a i r s w i t h i n c r e a s i n g age ( S c h e f f e r , 1964). The r e s u l t s of my c o m p a r i -son of h o r i z o n t a l s e c t i o n s t h r o u g h t h e sebaceous g l a n d l e v e l o f two s k i n s , an a d u l t female c o l l e c t e d May 1, 1964 near S i t k a , A l a s k a , and a female p u p - o f - t h e - y e a r c o l l e c t e d on 30 June 1965 i n t h e Queen C h a r l o t t e I s l a n d s , agree w i t h S c h e f f e r ' s (1964) f i n d i n g s . The a d u l t guard h a i r s v a r i e d from 66 t o 120 m i c r a i n diameter w i t h a mean of 88.3. The guard h a i r s o f t h e pup showed l i t t l e v a r i a t i o n from a mean diameter of 16 49.5 m i c r a . There was a c o r r e s p o n d i n g d i f f e r e n c e i n t h e u n d e r h a i r d i a m e t e r s . The a d u l t u n d e r h a i r s v a r i e d from 18 -45 m i c r a w i t h a mean dia m e t e r of 35 m i c r a . Secondary h a i r s of t h e pup ranged from 12 - 25.5 m i c r a w i t h an average diam-e t e r of 17 m i c r a . The mean d i s t a n c e between o v e r h a i r s was t a k e n as a measurement of f o l l i c l e group d e n s i t y . I n t h e a d u l t t h i s was 294 m i c r a ; i n t h e pup i t was 76 m i c r a . In a s t u d y o f h o r i z o n t a l s e c t i o n s t h r o u g h f o l l i c l e groups of 13 male and 23 female P. v. r i c h a r d i p e l t s (Appendices A, B) I found t h a t t h e average number of under-h a i r s per f o l l i c l e group was 3.19. The range was 1 - 6 . I n males t h e range was 1 - 6 w i t h a mean o f 3.24. The range i n fema l e s was 2 - 5 w i t h an average of 3.15. H a i r d i a m e t e r s were measured from t h e sebaceous g l a n d l e v e l ( w i t h e x c e p t i o n s n o t e d i n Appendix B ) . The range i n o v e r h a i r d i a m e t e r s was 33 t o 150 m i c r a , w h i c h compares c l o s e l y w i t h S c h e f f e r (1964). F o r t h e e n t i r e sample t h e mean o v e r h a i r diameter was 76.5 m i c r a . The d i a m e t e r s of male guard h a i r s ranged from 60 - 150 m i c r a w i t h a mean o f 85.5 m i c r a . F o r females t h e range was 33 - 120 m i c r a w i t h an average of 73.3. V a r i a t i o n s i n guard h a i r d i a m e t e r s w i t h i n i n d i v i d u a l s were n o t e d . I n one specimen two d i f f e r e n t o v e r -h a i r s measured 79.5 and 120 m i c r a at t h e sebaceous g l a n d l e v e l . I n t h e same specimen, u n d e r h a i r s of t h r e e f o l l i c l e g roups, two viewed at t h e sebaceous g l a n d l e v e l and one at Anagen, e a r l y , 20X. Note t h e o v e r h a i r , e x t e n d -i n g f r om t h e anagen b u l b i n t h e lower r i g h t , which i s growing a l o n g s i d e and p a s t a club h a i r . 18 t h e s t r a t u m corneum l e v e l were q u i t e u n i f o r m i n s i z e (Appendix B) . Some samples w h i c h o r i g i n a t e d d u r i n g August, 1964, e x h i b i t e d d o u b l i n g of t h e h a i r f o l l i c l e s and t h e p r e s e n c e of b o t h t h e o l d and new o v e r h a i r s . THE PRE-NATAL AND NEO-NATAL PELAGE I n Washington S t a t e , S c h e f f e r and S l i p p (1944) n o t e d t h a t t h e y l l o w i s h - w h i t e f o e t a l or lanugo p e l a g e was u s u a l l y r e p l a c e d by a s h o r t s p o t t e d c o a t p r i o r t o p a r t u r i t i o n . Montagna and H a r r i s o n (1957) wrote t h a t t h e common s e a l of t h e Wash shed i t s "wooly j u v e n i l e f u r " i n u t e r o . Bonnot (1951) s t a t e d t h a t , i n C a l i f o r n i a , t h e p r e - n a t a l c o a t was shed b e f o r e b i r t h . Ognev (1935) s t a t e d of P. v. v i t u l i n a t h a t " w h i t e l i n g s are never e n c o u n t e r e d among common s e a l s " . S c h e f f e r (1958) commented t h a t t h e r e seemed t o be no r e c o r d of P. v. r i c h a r d i newborn young i n t h e embryonic p e l a g e . E x c e p t i o n s do o c c u r . K i n g (1964) commented t h a t t h e embryonic h a i r was o c c a s i o n a l l y shed i m m e d i a t e l y a f t e r b i r t h . F i s h e r (1952) r e p o r t e d f i n d i n g a pup, on t h e E c s t a l l R i v e r , B.C., which s t i l l p o s s e s s e d an e x c e e d i n g l y l o o s e c o a t of f o e t a l h a i r some 15 - 20 minutes a f t e r p a r t u r i t i o n . I n 1965 lanugo h a i r was n o t e d on p o s t -n a t a l pups at Humboldt Bay, C a l i f o r n i a , on 10 May; Boundary Bay, B.C., on 23, 24 and 27 J u l y ; V i c t o r i a , B.C., on 6 August; and P o r t Moody, B.C. ( F i g u r e 6) on 10 August. Comparison of 19 t a n ned s k i n samples from a P. v. l a r g h a w h i t e c o a t f o e t u s from B r i t i s h C o l u m b i a ( b o t h s k i n s from Hansen Fur Company, L a n g l e y , B.C.) i n d i c a t e s t h a t t h e g r e a t e r t h i c k n e s s and w o o l i e r t e x -t u r e of t h e P. v. l a r g h a p e l a g e r e f l e c t s d i f f e r e n c e s i n t h e h a i r o f t h e two forms. Under t h e b i n o c u l a r m i c r o s c o p e t h e lanugo h a i r of r i c h a r d i ( F i g u r e 8a) appeared composed of s t r a i g h t o v e r h a i r s s u r r o u n d e d and i n t e r s p e r s e d w i t h much t h i n n e r u n d e r h a i r s . The o v e r h a i r s appeared about f i v e t i m e s t h e d i a m e t e r of t h e u n d e r h a i r s . The l o n g e s t h a i r s measured 20 mm i n l e n g t h . In l a r g h a ( F i g u r e 8b) t h e h a i r s were gen-e r a l l y wavy. The o v e r h a i r s were about h a l f t h e d i a m e t e r of t h e f o e t a l guard h a i r s i n r i c h a r d i . The u n d e r h a i r s were about one f i f t h t h e d i a m e t e r of t h e o v e r h a i r s and n e a r l y as l o n g . The l o n g e s t h a i r s measured 30 mm. In b o t h s u b s p e c i e s t h e h a i r s appeared m i l k i l y opaque and w i t h o u t v i s i b l e p i g -ment ( S t u t z , 1966). I have examined a p p r o x i m a t e l y 100 P. v. r i c h a r d i f o e t a l s k i n s . These f e l l r o u g h l y i n t o two c a t e g o r i e s . F i r m h a i r , " d i r t y " w h i t e or p a l e y e l l o w i s h - g r e y , w i t h a f a i n t l y d a r k e r a r e a a l o n g t h e d o r s a l m i d l i n e , which r e s i s t e d t u g g i n g and remained on t h e p e l t appeared c h a r a c t e r i s t i c of a f o e t u s at about h a l f t e rm ( e s t i m a t e d ) . As t h e f o e t u s neared p a r t u r i -t i o n t h e h a i r would become p r o g r e s s i v e l y l o o s e r u n t i l i t c o u l d be s c r a p e d from t h e s k i n w i t h my thumb. Near-term f o e t a l s k i n s r a r e l y e x h i b i t e d any lanugo h a i r . 20 F i g u r e 6. Phoca v i t u l i n a r i c h a r d i female pup c a p t u r e d at P o r t Moody, B.C., on 10 August, 1965. Weight 17 l b s . U m b i l i c u s 3 1/2 i n c h e s l o n g , p i n k - r e d . F o e t a l (lanugo) h a i r c o a t prominent except on t o p of head, f a c e , and f o r e f U p p e r s . P h o t o -graphed at s e r v i c e y a r d , S t a n l e y P a r k , Vancouver, B.C., 13 August, 1965. F i g u r e 7. Tanned s k i n o f Phoca v i t u l i n a l a r g h a pup from B e r i n g Sea. Depth of t h e w h i t e w o l l y c o a t 30 mm. Note t h e s h o r t , s p o t t e d a d u l t t y p e coat v i s i b l e i n t h e t h r e e s m a l l p l u c k e d a r e a s . Figure 8. Tanned skin slices viewed in cross section. Magnification about 3X. a. Phoca vitulina richardi foetus. b. Phoca vitulina largha whitecoat pup. 23 Moult of t h e embryonic h a i r b oth i n u t e r o and when p o s t - n a t a l l y p r e s e n t f o l l o w s a r e g u l a r p a t t e r n i n P. v. r i c h a r i as d e t e r m i n e d by e x a m i n a t i o n and photography o f a s e r i e s of samples. Such a s e r i e s has not been a v a i l a b l e f o r P. v. l a r g h a . I n r i c h a r d i ( F i g u r e 9, b-e) t h e lanugo h a i r b e g i n s t o shed on th e a n t e r i o r d o r s a l s u r f a c e . As t h e moult c o n t i n u e s i n t h e a n t e r o - d o r s a l a r e a t h e p o s t e r i o r d o r s a l r e g i o n b e g i n s t o shed. The moult i s completed on t h e a n t e r i o r and p o s t e r i o r d o r s a l s u r f a c e s and t h e f l a n k s b e f o r e t h e f o e t a l h a i r i s shed from t h e v e n t r a l r e g i o n . The m i d d o r s a l l i n e i n t h e t h o r a c i c and lumbar r e g i o n s i s t h e f i n a l a r e a t o moult t h e f o e t a l h a i r i n r i c h a r d i . I n b o t h r i c h a r d i and l a r g h a t h e f o e t a l h a i r i n i t i a l l y c o n c e a l s t h e p o s t - n a t a l c o a t w h i c h i s exposed by moult or removal of t h e lanugo h a i r ( F i g u r e 7) ( S t u t z , 1966). The s h o r t , s p o t t e d , c o a t t h a t l i e s beneath t h e lanugo h a i r of t h e f o e t a l p e l a g e o r d i n a r i l y r e a c h e s i t s f u l l growth p r i o r t o p a r t u r i t i o n . Montagna and H a r r i s o n (1957) f o u n d t h a t newborn s e a l s from t h e Wash had no d e v e l o p i n g f o l -l i c l e s and were i n t e l o g e n ( t h e c l u b h a i r phase o f t h e moult c y c l e ) h a v i n g completed t h e i r b i r t h c o a t s i n u t e r o . M i c r o -s c o p i c e x a m i n a t i o n of s k i n samples of specimens from B r i t i s h C o l u m b i a and a d j a c e n t w a t e r s i d e n t i f i e d as p u p s - o f - t h e - y e a r r e v e a l e d t h a t a l l pups were i n t e l o g e n when t h e m a j o r i t y of specimens of o t h e r age c l a s s e s were i n anagen ( t h e phase of a c t i v e growth and development of new h a i r ) . F i g u r e 9 . The f o e t a l moult p a t t e r n i n P. v. r i c h a r d i . A. O u t l i n e d r awing of t y p i c a l s e a l p e l t a f t e r r e m o v a l from t h e c a r c a s s . Key: a, a n t e r i o r ; f , o p e n i n g at a n t e r i o r f l i p p e r s i t e ; m d, mid-d o r s a l l i n e ; p, p o s t e r i o r ; v, v e n t r a l m i d l i n e . B. The l i n e d a r e a , p l u s the a n t e r i o r f l i p p e r s and t h e f a c e , i s t h e f i r s t p a r t of t h e f o e t a l p e l t t o moult t h e lanugo h a i r . C. D. L a t e r s t a g e s of t h e f o e t a l moult. Note t h a t t h e v e n t r a l and m i d - d o r s a l areas r e t a i n t h e i r lanugo h a i r u n t i l l a t e i n t h e moult. E. The c r o s s - h a t c h e d p o r t i o n of t h e lumbar m i d - d o r s a l r e g i o n i s t h e l a s t a r e a t o shed t h e f o e t a l h a i r . 25 MOULT OF THE ADULT PELAGE S e v e r a l w r i t e r s have r e f e r r e d t o or p a r t l y d e s c r i b e d moult i n Phoca v i t u l i n a . M u r i e (1959) r e p o r t e d moult i n h a r -bour s e a l s on June 17, 1925 at Izembek Bay i n s o u t h w e s t e r n A l a s k a . Ognev (1935) quoted M i l l a i s , 1904 t o i n d i c a t e t h a t i n P. v. v i t u l i n a moult o c c u r r e d i n August and September. S c h e f f e r and S l i p p (1944) r e p o r t e d t h a t a male P. v. r i c h a r d i kept i n an outdoor aquarium began t o moult i n t h e l a s t week i n August and had n e a r l y c o mpleted t h e moult by September 25, 1942. On October 7, 1942 t h e y o b s e r v e d v a r i o u s moult s t a g e s s i m u l t a n e o u s l y i n p r o g r e s s among a h e r d of 60 s e a l s on t h e N i s q u a l l y F l a t s , s o u t h e r n Puget Sound, Washington ( S c h e f f e r and S l i p p , 1944). F i s h e r (1952) n o t e d t h a t an a d u l t female c o l l e c t e d August 8, 1946 on t h e Skeena R i v e r , B r i t i s h Colum-b i a , was i n an e a r l y s t a g e of moult. B e l k i n (1964) r e p o r t e d t h e f i r s t s i g n s of moult i n Phoca i n s u l a r i s (Phoca v i t u l i n a ? ) i n e a r l y J u l y and specimens w h i c h had completed t h e moult were t a k e n i n August. Montagna and H a r r i s o n (1957), r e f e r -r i n g t o s e a l s from t h e Wash, Engla n d , s t a t e d t h a t moult t o o k p l a c e d u r i n g t h e summer months. Specimens c o l l e c t e d i n l a t e August e x h i b i t e d anagen ( a c t i v e growth phase) or c a t a g e n ( b u l b r e g r e s s i o n p h a s e ) . Newborn pups and s e a l s c o l l e c t e d i n w i n t e r and s p r i n g e x h i b i t e d t e l o g e n ( " r e s t i n g f o l l i c l e " phase) and p o s s e s s e d o n l y c l u b h a i r f o l l i c l e s (Montagna and H a r r i s o n , 1957). 26 E x t e r n a l s i g n s of moult such as f a d i n g p e l a g e , b r i t t l e h a i r , and s h e d d i n g h a i r a r e n o t e d over much of t h e year on t h e B r i t i s h C olumbia c o a s t and a d j o i n i n g r e g i o n s . "Shedder" s k i n s c o l l e c t e d at A n a c o r t e s , Washington, on J a n u -a r y 10, 1965 (Hansen, p e r s o n a l communication) and a m o u l t i n g a d u l t male at Howe Sound on 21 November 1965 ( B i g g , p e r s o n a l communication) are t h e e a r l i e s t and l a t e s t moult d a t e s r e p o r t e d . U n e x p l a i n e d a n o m a l i e s have been r e p o r t e d such as a p o p u l a t i o n (?) of s e a l s at Smith I s l a n d , Washington, which p r o v i d e d prime p o s t - m o u l t p e l t s i n l a t e October when t h e s e a l s of nearby A n a c o r t e s were r e p o r t e d t o be "100% s h e d d e r s " (March and Hansen, p e r s o n a l c o m m u n i c a t i o n ) . An e x a m i n a t i o n of s k i n s c o m m e r c i a l l y o b t a i n e d i n d i c a t e s t h a t d e g e n e r a t i o n o f h a i r f r om prime c o n d i t i o n has become n o t i c e a b l e i n some s k i n s by F e b r u a r y . H a i r on v e n t r a l a r e a s b e g i n s t o t u r n p a l e and b e g i n s t o f e e l c o a r s e r t h a n h a i r on t h e dorsum. B l a c k h a i r s on t h e d o r s a l s u r f a c e d e v e l o p brown or w h i t i s h t i p s . The w h i t e r i n g s of c h a r a c t e r i s t i c spot p a t t e r n s become p a l e t a n and t h e b l a c k c e n t e r s o f r o s e t t e s b e g i n t o brown. As t h e p r o c e s s c o n t i n u e s bare (worn?) a r e a s may appear i n t h e " a r m p i t s " at t h e bases of t h e a n t e r i o r f l i p p e r s . A c t u a l moult may t a k e p l a c e i n almost any month of t h e y e a r . Samples from 261 s k i n s , randomly but u n e v e n l y d i s t r i b u t e d a c r o s s t h e year (Appendices C, D) were a n a l y z e d m i c r o s c o p i c a l l y t o d e t e r m i n e t h e s t a g e of f o l l i c l e 27 development of each i n d i v i d u a l and t h e p e r c e n t a g e of s p e c i -mens i n each s t a g e of t h e moult c y c l e f o r each month. The r e s u l t s ( F i g u r e 10; T a b l e I) i n d i c a t e d t h a t anagen o c c u r r e d as e a r l y as F e b r u a r y , i n v o l v e d most of t h e p o p u l a t i o n i n J u l y - September and was s t i l l apparent i n some i n d i v i d u a l s i n November. From December t h r o u g h A p r i l almost a l l s p e c i -mens were i n t e l o g e n . T a b l e I . Moult p r o g r e s s i o n by months, p e r c e n t a g e s . ANAGEN Male Female T o t a l s 5 J a n . 0 .00 0 .00 0 .00 Feb. 0 .00 7 .70 5.60 Mar. 0 .00 0 .00 0.00 Apr . 15 .40 5 .55 13.00 May 27 .30 25 .00 20 .08 Jun. 0 .00 5 . 27 3.44 J u l . 0 .00 28 .60 44.45 Aug. 88 .88 88 .88 88.88 Sep. 100 .00 75 .00 83.00 Oct . 0 , .00 0 .00 0.00 Nov . 0 .00 20 .00 10.00 Dec. 0 .00 0 .00 0 .00 CATAGEN Not d e t e c t e d i n t h e sample. TELOGEN Ja n . 100 .00 100 .00 100.00 Feb. 100 .00 92 .30 94.40 Mar . 100 .00 100 .00 100.00 Apr . 84 .60 94 .45 87 .00 May 100 , .00 94 .73 96.56 J u l . 100 .00 71 .40 55.55 Aug. 11 . 12 11 .12 11.12 Sep. 0 .00 25 .00 17 .00 Oct . 100 .00 100 .00 100.00 Nov. 100 .00 80 .00 90.00 Dec. 100 .00 100 .00 100.00 * i n c l u d e s unsexed specimens. f» 7» to (V 39 /* 0 r ?• <» ro /• Jo it n J ^ y»f >1 J J A S 0 * 2> JFAIAAIJJASOMJ) Te/oe,t7t An»aen n n J f / H ^ J J H i ? V P / / / f i r s / rttf9$Y/l?/<r( JLCI. t o J f i S i A M J J A S O A / J 7i/OAT*, Mt/ ftmjt/t. F i g u r e 10. H i s t o g r a m o f t h e a n n u a l m o u l t i n P h o c a v i t u l i n a r i c h a r d i to oo 29 R ather s u r p r i s i n g l y , from t h e specimens sampled, no l a t i t u d i n a l d i f f e r e n c e s i n t h e moult p e r i o d were n o t e d . Specimens from A l a s k a ' s G l a c i e r Bay and from t h e G u l f I s l a n d s of B r i t i s h C olumbia were i n s i m i l a r s t a g e s at t h e same t i m e i n s p i t e of d i f f e r e n c e s i n season l e n g t h s and amounts of d a i l y l i g h t . The f a i l u r e of t h e sample t o i n d i c a t e sex and age d i f f e r e n c e s i n moult t i m i n g ( w i t h t h e e x c e p t i o n t h a t p u p s - o f - t h e - y e a r were a l l i n t e l o g e n ) was e q u a l l y s u r p r i s i n g . S c h e f f e r and S l i p p (1944) n o t e d t h a t Havinga had w r i t t e n of P. v. v i t u l i n a of H o l l a n d i n 1933 t h a t t h e y e a r l i n g s m o u l t e d i n J u l y w h i l e t h e a d u l t s r e p l a c e d t h e i r p e l a g e i n August or September. At Tugidak I s l a n d , T r i n i t y I s l a n d s , A l a s k a Mr. Henry Hansen noted on May 25, 1965 t h a t young harbour s e a l s i d e n t i f i e d as " y e a r l i n g s " were c o n s p i c u o u s l y s h e d d i n g . At th e same date and l o c a t i o n t h e a d u l t s showed no outward s i g n s o f moult (Hansen, p e r s o n a l communication). I n o t h e r s p e c i e s , sex and age d i f f e r e n c e s have been r e c o r d e d f o r d a t e s o f moult i n t h e harp s e a l , P a g o p h i l u s g r o e n l a n d i c u s ( F i s h e r , u n p u b l i s h e d ) and t h e e l e p h a n t s e a l , Mirounga l e o n i n a ( C a r r i c k e t a l , 1962; L i n g , 1965) among t h e P h o c i d a e and i n t h e n o r -t h e r n f u r s e a l , C a l l o r h i n u s u r s i n u s ( S c h e f f e r , 1961; S c h e f f e r and Johnson, 1963) and t h e South A f r i c a n f u r s e a l , A r c t o -c e p h a l u s p u s i l l u s (Rand, 1956) among t h e O t a r i i d a e . The l e n g t h of t i m e t h a t P. v. r i c h a r d i remains i n each of t h e s t a g e s of t h e moult c y c l e has not been d e t e r m i n e d . 30 S c h e f f e r and S l i p p (1944) n o t e d from e x t e r n a l e v i d e n c e t h a t moult i n a c a p t i v e specimen was completed i n about f i v e weeks. Montagna and H a r r i s o n (1957) found t h a t p l u c k e d h a i r f o l l i c l e s r e q u i r e d about t h r e e weeks f o r e s t a b l i s h m e n t of t h e b u l b and a f u r t h e r t h r e e weeks f o r t h e new h a i r t o r e a c h t h e e p i d e r m a l s u r f a c e ( F i g u r e 5 ) . E x a m i n a t i o n o f "shedder" s k i n s from which t h e o l d h a i r s can be shaken, b r u s h e d or p u l l e d r e v e a l s t h a t t h e s e c u r e l y f a s t e n e d h a i r of t h e new growth i s about h a l f t h e l e n g t h or l e s s , of t h e o l d h a i r . Shedding must, t h e r e f o r e , be o c c u r r i n g d u r i n g anagen. The mechanism of s h e d d i n g i s not u n d e r s t o o d . The t h i c k e n e d e p i d e r m a l c o l l a r w h i c h s u r r o u n d s t h e f o l l i c l e below t h e o r i f i c e (Montagna and H a r r i s o n , 1957) l o o s e n s and t h e b r u s h end of t h e c l u b h a i r ( F i g u r e 4) p u l l s l o o s e w i t h some o f t h e a d j a c e n t c e l l s a t t a c h e d ( L i n g , p e r s o n a l c o m m u n i c a t i o n ) . At t h i s t i m e t h e d e v e l o p i n g h a i r s of t h e new g e n e r a t i o n are f i r m l y a t t a c h e d t o t h e b u l b s from w h i c h t h e y are growing. Catagen, t h e s t a g e of r e g r e s s i o n of t h e h a i r b u l b and dermal p a p i l l a (Montagna and V a n S c o t t , 1958; B u l l o u g h and Laure n c e , 1958) i s of v e r y s h o r t d u r a t i o n i n p h o c i d s ( L i n g , p e r s o n a l communication) and has not been i d e n t i f i e d i n t h e specimens s t u d i e d . The a c t i v e h a i r b u l b and p a p i l l a o f anagen do not r e g r e s s i n P. v. r i c h a r d i t o t h e same e x t e n t as i n s p e c i e s i l l u s t r a t e d or d e s c r i b e d i n t h e l i t e r a t u r e o f h a i r development (Montagna and V a n S c o t t , 1958; B u l l o u g h and F i g u r e 11. T y p i c a l moult p r o g r e s s i o n i n t h e harbour s e a l o f B r i t i s h C olumbia. A. E a r l y moult. D i a g o n a l shaded areas around eye, nose, mouth and v e n t r a l and d o r s a l s u r f a c e s are s h e d d i n g o l d h a i r and e x p o s i n g s h o r t new h a i r . L i g h t l y shaded r e g i o n of f l a n k i s zone of c o n s p i c u o u s l y f a d e d and worn h a i r . B. A l a t e r s t a g e of moult. New h a i r has r e p l a c e d o l d h a i r i n t h e d i a g o n a l l y shaded a r e a s . Shedding i s i n p r o g r e s s i n t h e l i g h t l y shaded a r e a and s p r e a d i n g l a t e r a l . 32 Laurence, 1958; Rook, 1965) and te n d s t o p e r s i s t i n reduced but r e c o g n i z a b l e form t h r o u g h o u t t e l o g e n ( F i g u r e 4 ) . The a d u l t moult i s p a t t e r n e d , as i n many o t h e r mammals ( S c h e f f e r , 1961; C r o w c r o f t , 1957; Morejohn and Howard, 1956; B a s s e t t and L l e w e l l y n , 1948) and d i f f e r s i n sequence ( F i g u r e 11) from t h e i n t r a u t e r i n e moult of t h e f o e t u s ( F i g u r e 9) ( S t u t z , 1966). My o b s e r v a t i o n s i n d i c a t e t h a t t h e g e n e r a l p a t t e r n o f t h e moult agrees w i t h t h e d e s c r i p t i o n of a c a p t i v e s e a l by S c h e f f e r and S l i p p (1944). The f i r s t a r e a s t o r e p l a c e th e h a i r are around t h e body openings (anus, eyes, p e n i a l o p e n i n g , e a r s ) . Moult of t h e body i t s e l f b e g i n s a l o n g t h e v e n t r a l m i d l i n e and on t h e a n t e r i o r (neck) and p o s t e r i o r ( t a i l ) d o r s a l m i d l i n e a r e a s and p r o g r e s s e s as a wave of development and s h e d d i n g u n t i l t h e moult i s complete ( F i g u r e 11). The e x a c t p a t t e r n of moult i s s u b j e c t t o much i n d i v i d -u a l v a r i a t i o n . Some s k i n s , c l a s s i f i e d by p e l t b u yers as " t o t a l s h e d d e r s " , moult h a p h a z a r d l y and may have v a r i o u s s i z e d b a l d p a t c h e s i n whi c h r e g e n e r a t i n g h a i r i s not apparent. These may r e f l e c t p a t h o l o g i c a l c o n d i t i o n s . Roest (1964) r e p o r t e d on a n e a r l y h a i r l e s s r i b b o n s e a l , H i s t r i o p h o c a f a s c i a t a cap-t u r e d on t h e s o u t h e r n C a l i f o r n i a c o a s t i n November, 1962. Roest (1964) s u g g e s t e d i l l n e s s as a p o s s i b l e cause of b a l d -ness i n t h i s specimen. S c h e f f e r (1958) mentioned b a l d n e s s i n 33 r i n g e d s e a l s , Pusa h i s p i d a , but w i t h o u t e x p l a n a t i o n . McLaren (1958) r e p o r t e d on b a l d n e s s i n r i n g e d s e a l s i n t h e e a s t e r n C a nadian a r c t i c but o f f e r e d no e x p l a n a t i o n . A specimen of Phoca v i t u l i n a v i t u l i n a , c a p t u r e d when about two weeks o l d , was t o o t h l e s s and almost c o m p l e t e l y b a l d and d i d not d e v e l o p h a i r or t e e t h d u r i n g t h r e e y e a r s of c a p t i v i t y ( K i n g , 1964) . The coat o f t h e p o s t - n a t a l pup i n harbour s e a l s , except i n Phoca v i t u l i n a l a r g h a ( W i l k e , 1954; K i n g , 1964), i s n o r m a l l y s i m i l a r i n c o l o u r and p a t t e r n t o t h e t y p i c a l a d u l t p e l t ( S c h e f f e r and S l i p p , 1944; F i s h e r , 1952; Montagna and H a r r i s o n , 1957). An i n d i c a t i o n t h a t t h e p a t t e r n p e r s i s t s t h r o u g h s u c c e s s i v e m o u l t s i s i n d i c a t e d by F i g u r e s 12 and 13 w h i c h were s u p p l i e d by Dr. R.J. H a r r i s o n of t h e London Hos-p i t a l M e d i c a l C o l l e g e , England. In t h e s e photographs, t a k e n i n 1960 and 1962 t h e d i s t i n c t i v e p a t t e r n o f s p o t s on t h e " c h e s t " a n t e r i o r t o t h e l e f t p e c t o r a l f l i p p e r a r e c l e a r l y v i s i b l e , and c l e a r l y i d e n t i c a l . Figure 12. Three year old female common seal, Phoca  v i t u l i n a v i t u l i n a photographed November, 1960; Compare the spot pattern v i s i b l e anterior to the l e f t front f l i p p e r with Figure 13. (Photo courtesy of Professor R.J. Harrison, London Hospital Medical College.) F i g u r e 13. F i v e year o l d female P. v. v i t u l i n a photo-graphed August, 1962. T h i s i s t h e same s e a l i l l u s t r a t e d i n F i g u r e 12. Note t h e unchanged spot p a t t e r n a n t e r i o r t o t h e l e f t f r o n t f l i p -p e r . (Photograph by c o u r t e s y of P r o f e s s o r R. J . H a r r i s o n , London H o s p i t a l M e d i c a l C o l l e g e . ) 36 PELAGE PATTERNS AND COLOURS DESCRIPTION Harbour s e a l s a re d e s c r i b e d and i l l u s t r a t e d w i d e l y i n t h e l i t e r a t u r e . Imaizumi (1961) p o r t r a y e d a v e r y d a r k s e a l w i t h w h i t e r i n g s s u r r o u n d i n g b l a c k s p o t s w h i c h he i d e n t i f i e d as Phoca v i t u l i n a k u r i l e n s i s of t h e K u r i l e I s l a n d s . Another i l l u s t r a t i o n , o f a v e r y l i g h t c o l o u r e d s e a l w i t h dark s p o t s he r e f e r r e d t o Phoca v i t u l i n a l a r g h a . Ognev (1935) d e s c r i b e d P. v. l a r g h a as u s u a l l y h a v i n g a l i g h t e r background c o l o u r t h a n P. v. v i t u l i n a o f t h e e a s t e r n A t l a n t i c . The dark c o l o u r d e s c r i b e d as t y p i c a l of t h e shore pupping form o f t h e K u r i l e I s l a n d s was used by B e l k i n (1964) as a taxonomic d e v i c e f o r s e p a r a t i n g Phoca i n s u l a r i s (Phoca v i t u l i n a k u r i l e n -s i s ? ) from P. v. l a r g h a • S i m i l a r l y , c o l o u r was a key c h a r a c -t e r i s t i c i d e n t i f y i n g P. v. g e r o n i m e n s i s as a form d i s t i n c t f rom P. v. r i c h a r d i (Schwarz, 1942; S c h e f f e r and S l i p p , 1944). Ognev (1935) quoted M i l l a i s , 1904 c o n c e r n i n g c o l o u r s and p e l a g e p a t t e r n s of harbour s e a l s of t h e e a s t e r n A t l a n t i c and went i n t o g r e a t d e t a i l i n an e f f o r t t o d e s c r i b e c o l o u r and p a t t e r n t y p e s of P a c i f i c harbour s e a l s . A l t h o u g h t h e r e i s g r e a t v a r i a t i o n between i n d i v i d u a l s k i n s i t appears t h a t no c o m b i n a t i o n of s p o t s or c o l o u r s i s e x c l u s i v e t o e i t h e r ocean. No w r i t t e n system of p e l a g e c l a s s i f i c a t i o n may be adequate f o r so v a r i e d a s p e c i e s . The c l a s s i f i c a t i o n i n d i -37 c a t e d i n T a b l e I I i s based on my p e r s o n a l o b s e r v a t i o n s and i s an e f f o r t t o s i m p l i f y t h e d e s c r i p t i o n s of t h e m a j o r i t y of s k i n s c o l l e c t e d from t h e P a c i f i c c o a s t of N o r t h A m e r i c a . T a b l e I I . P e l a g e t y p e s of harbour s e a l , Phoca v i t u l i n a , s k i n s c o l l e c t e d on t h e P a c i f i c c o a s t . G e n e r a l D e s c r i p t i o n B l a c k or v e r y dark brown on b o t h dorsum and v e n t e r . Marks w h i t e , d i s t i n c t . B l u i s h - g r e y dorsum, v a r y i n g from almost w h i t e t o v e r y dark. F l a n k s and v e n t e r p a l e r t h a n dorsum, u s u a l l y w h i t e . Spots on f l a n k s and v e n t e r may be b l u i s h - g r e y or brown. Spots on dorsum may be b l a c k , brown or w h i t e . Marks are d i s t i n c t . M o t t l e d shades of i n d i s t i n c t p a t t e r n and vague ma r k i n g s . P a l e shades of m i n g l e d g r e y - b e i g e - w h i t e w i t h dorsum d a r k e r i n n e u t r a l shade, f r e q u e n t l y w i t h o u t marks on v e n t e r ; or dark b l u i s h - b r o w n shade of d o r -sum w i t h l i g h t e r v e n t e r and i n d i s t i n c t mot-t l e d m arkings. To t h e t h r e e major n a t u r a l c o l o u r t y p e s , b l a c k , common and muddy s h o u l d be added a f o u r t h or a r t i f a c t c o l -our t y p e . The s o - c a l l e d " A l a s k a g o l d e n s e a l s k i n s " are never f o u n d on l i v i n g s e a l s , wet or d r y . They r e s u l t from i m p r o p e r l y p r o c e s s e d raw s e a l s k i n s where t h e h a i r has been smeared w i t h b l u b b e r g r e a s e and a l l o w e d t o d r y (Dunbar, 1949). The y e l l o w c o l o u r can be p r e v e n t e d by c a r e f u l l y r emoving a l l b l u b b e r from t h e h i d e and washing t h e p e l t s i n d e t e r g e n t s t o remove f a t t h a t has been rubbed ont o t h e h a i r . 1. B l a c k 2. Common 3. Muddy 38 " O x i d i z e d " ( y e l l o w ) s k i n s b r i n g l owered market p r i c e s s i n c e European p r o c e s s o r s dye them t o c o n c e a l t h e f a u l t y c o l o u r . COLOUR PATTERN - SIZE RELATIONSHIPS IN COMMERCIAL SKINS The a r e a s of i n d i v i d u a l s S k i n s , t o t h e n e a r e s t 1/4 square f o o t were measured i n t h r e e s e r i e s of p e l t s at Hansen Fur Company. A l l s k i n s were s t r e t c h e d t o t h e i r maximum s i z e and d r i e d b e f o r e measuring. No s e x i n g or a g i n g were a p p l i e d t o t h e s e s k i n s . Sample A, 472 s k i n s , common c o l o u r p a t t e r n , a l l a r e a s , undated. Sample B, 33 s k i n s , common c o l o u r p a t t e r n , T r i n i t y I s l a n d s , A l a s k a , J u l y , 1955. Sample C, 33 s k i n s , b l a c k c o l o u r p a t t e r n , a l l a r e a s , undated. T a b l e I I I . Comparison of samples. Range, Sq. F t . Mean, Sq. F t . S t a n d a r d D e v i a t i o n A. 5.00 - 17.75 11.173 2.8 B. 7.00 - 17.25 12.666 1.3 C . 9.00 - 16.25 13.015 1.6 A p p l i c a t i o n of S t u d e n t ' s T Te s t ( L i , 1964) i n d i -c a t e d no s t a t i s t i c a l l y s i g n i f i c a n t d i f f e r e n c e between t h e means o f samples B and C (at t h e 5% s i g n i f i c a n c e l e v e l T 39 e q u a l l e d -0.306 w i t h 64 degrees of freedom). Between samples A and C t h e r e was a s i g n i f i c a n t d i f f e r e n c e i n t h e means (at t h e 5% s i g n i f i c a n c e l e v e l , T e q u a l l e d -3.71 w i t h i n f i n i t e d e g r e s s of freedom). There was no s i g n i f i c a n t d i f f e r e n c e between samples A and B. At t h e 5% s i g n i f i c a n c e l e v e l T e q u a l l e d -0.961 w i t h i n f i n i t e degrees of freedom. THE GEOGRAPHICAL DISTRIBUTION OF COLOUR AND PATTERN Dunbar (1949) and M a n s f i e l d (1963) w r i t i n g of P. v. c o n c o l o r o f e a s t e r n Canada commented t h a t harbour s e a l s were much d a r k e r i n t h e n o r t h e r n p a r t s of t h e i r range t h a n t h e i r c o n s p e c i f i c s f u r t h e r s o u t h . T h i s t r e n d does not appear among P a c i f i c harbour s e a l s . The harbour s e a l of s o u t h e r n C a l i f o r n i a and t h e n o r t h w e s t e r n c o a s t of Mexico was f o r m e r l y c l a s s i f i e d as Phoca v i t u l i n a g e r o n i m e n s i s ( M i l -l e r and K e l l o g g , 1955) on t h e b a s i s of i t s dark c o l o u r (Schwarz, 1942) . I n g l e s (1957) e x h i b i t s a photograph of a b l a c k harbour s e a l from near La J o l l a , C a l i f o r n i a . I have examined b o t h v e r y d a r k and v e r y l i g h t s k i n s f rom Puget Sound, Queen C h a r l o t t e I s l a n d s and s e v e r a l A l a s k a n l o c a -t i o n s . S c h e f f e r and S l i p p (1944) remarked upon v e r y l i g h t c o l o u r e d s e a l s from Washington S t a t e p o i n t s and a n e a r l y e n t i r e l y b l a c k female from t h e F r a s e r R i v e r . T a b l e IV i n d i c a t e s t h e c o l o u r p a t t e r n s and T a b l e IV. D i s t r i b u t i o n of p e l a g e t y p e s by a r e a . A r e a A B C D E F G T o t a l s Number % 50 55. 56 6 18.75 17 23.61 7 50 .00 7 41.18 4 33.33 5 50 .00 96 BLACK Number % 14 15.55 12 32.50 33 45.83 2 14.29 8 47.06 4 33.33 3 30 .00 76 COMMON Number % 26 28.29 14 43.75 22 30.56 5 35.71 2 11.76 4 33.33 2 20 .00 75 MUDDY T o t a l Numbers 90 32 72 14 17 12 10 247 Are a A. Queen C h a r l o t t e I s l a n d s , B r i t i s h C o l u m b i a B. G l a c i e r Bay, A l a s k a C. G e o r g i a S t r a i t , B r i t i s h Columbia D. K n i g h t I n l e t , B r i t i s h Columbia E. B e r i n g Sea F. Skeena - P r i n c e Rupert, B r i t i s h C o l u m b i a G. Juneau, A l a s k a 41 g e o g r a p h i c o r i g i n s of 247 s e a l p e l t s . The p a t t e r n of d i s -t r i b u t i o n of t h e b l a c k and common s k i n s was t e s t e d f o r s i g -n i f i c a n c e w i t h an "F t e s t " a n a l y s i s of v a r i a n c e ( L i , 1964). The r e s u l t s i n d i c a t e d s i g n i f i c a n t d i f f e r e n c e s i n t h e d i s t r i -b u t i o n of b l a c k (F e q u a l s 4.06) and common (F e q u a l s 3.66) s k i n s but not i n t h e d i s t r i b u t i o n o f muddy p e l t s (F e q u a l s 1.05) . Duncan's m u l t i p l e range t e s t ( L i , 1964) i n d i c a t e d s t a t i s t i c a l l y s i g n i f i c a n t d i f f e r e n c e s between t h e means of t h e b l a c k components of t h e Queen C h a r l o t t e I s l a n d s , G l a c i e r Bay, and G e o r g i a S t r a i t samples. The means of common p a t -t e r n e d s k i n s d i f f e r e d s i g n i f i c a n t l y between t h e Queen Char-l o t t e I s l a n d s and S t r a i t of G e o r g i a samples, but not between t h o s e a r e a s and t h e G l a c i e r Bay sample. S e v e r a l specimens of b o t h j u v e n i l e and a d u l t p e l t s from t h e c o a s t of Maine, A t l a n t i c Ocean, were i n b l a c k and muddy p e l a g e p a t t e r n s . These were not s e p a r a b l e i n t y p e from P a c i f i c Ocean specimens. FOREIGN GROWTHS AND STAINS OF HAIR A l g a l i n c r u s t a t i o n of t h e e n t i r e a n t e r i o r of a s e a l h i d e and i t s d o r s a l s u r f a c e t o t h e t a i l ( F i g u r e 14) was prominent on t h e h a i r o f a female c o l l e c t e d 19 September 1965 at Westham I s l a n d , mouth of t h e F r a s e r R i v e r , B r i t i s h F i g u r e 14. S k e t c h of alg a e e n c r u s t e d p e l t of female harbour s e a l c o l l e c t e d 19 September, 1965 at Westham I s l a n d , mouth of t h e F r a s e r R i v e r , B r i t i s h Columbia. The l i n e d a r e a was t h i c k l y e n c r u s t e d w i t h P l e u r o c a p s a sp. T h i s specimen was i n a c t i v e moult and o b v i o u s l y s h e d d i n g i t s " o l d " h a i r . 43 Columbia. The growth was i d e n t i f i e d as a b l u e - g r e e n a l g a e , Cyanophyta, Chamaesiphonales, P l e u r o c a p s a sp. by Drs. R.F. S c a g e l and J.R. S t e i n of t h e Department of B i o l o g y and Botany, U n i v e r s i t y of B r i t i s h Columbia. Westham I s l a n d i s l o c a t e d i n t h e mouth of t h e F r a s e r R i v e r i n a r e g i o n of v e r y low s u r f a c e s a l i n i t y and moderate s a l i n i t y at lower depths (Waldichuk, 1957; T a b a t a and Le B r a s s e u r , 1958). F r i t s c h (1945) d e s c r i b e d P l e u r o c a p s a as a genus of w e l l a e r a t e d ocean w a t e r s , s w i f t r u n n i n g streams and t h e l i t t o r a l a reas of l a k e s . S i n c e t h e p a r t i c u -l a r s p e c i e s was not i d e n t i f i e d no d e c i s i o n can be r e a c h e d as t o t h e p r o b a b l e a r e a , s a l t or f r e s h , where t h e i n c r u s t a t i o n o r i g i n a t e d . S c h e f f e r and S l i p p (1944) commented t h a t a l g a e and b a r n a c l e growths on s e a l h a i r had been r e p o r t e d . S c h e f f e r (1962) remarked t h a t " e p i p h y t i c " green a l g a e ( C h l o r o p h y t a ) had been r e p o r t e d from p h o c i d s . Brown a l g a e ( E c t o c a r p u s ) , r e d a l g a e ( E r y t h r o c l a d i a ) , and gooseneck b a r n a c l e s (Lepas) were seen o c c a s i o n a l l y on f u r s e a l s ( C a l l o r h i n u s ) . The p r e s e n c e of c r e o s o t e smudged i n t o t h e h a i r was n o t e d on s k i n s of a n i m a l s which had p r o b a b l y been i n c o n t a c t w i t h wharf p i l i n g s . Large c r e o s o t e - l i k e markings were seen on two female s k i n s c o l l e c t e d from t h e w e s t s i d e of S h e l i k o f S t r a i t , A l a s k a , i n mid-May, 1965. The s t a i n s p e n e t r a t e d t h e 44 f u l l length of the hair in large blotches located on the body immediately posterior to the pectoral flippers. 45 DISCUSSION H a i r i s a key c h a r a c t e r i s t i c of t h e Mammalia (Cockrum, 1962; B l a i r , 1957) which has been l o s t or g r e a t l y r e d u c e d i n some mammals, n o t a b l y among t h e C e t a c e a ( B l a i r , 1957; H a r r i s o n and K i n g , 1965). P r o t e c t i o n a g a i n s t c o l d and concealment from p r e y or p r e d a t o r s (Romer, 1962) are f u n c t i o n s performed by h a i r . A i r t r a p p e d between t h e h a i r s and t h e s k i n i s t h e p r i n c i p a l i n s u l a t o r o f most h a i r e d mammals (Romer, 1962). The e r e c -t o r p i l i muscle, which r a i s e s h a i r t o improve entrapment of a i r , i s absent from a l l p i n n i p e d s ( S o k o l o v , 1960a; K i n g , 1964) . R e t e n t i o n of h a i r by harbour s e a l s may be c o n s i d -e r e d a c o n s e r v a t i v e t r a i t . The mammal a n c e s t r a l t o t h e P h o c i d a e , not yet i d e n t i f i e d (McLaren, 1960) u n d o u b t e d l y had h a i r and, w i t h t h e e x c e p t i o n of t h e e r e c t o r p i l i muscle, t h e s e a l has a complete mammalian h a i r system c o n s i s t i n g of o v e r -h a i r s , u n d e r h a i r s , sweat and sebaceous g l a n d s (Montagna and H a r r i s o n , 1957). Montagna and H a r r i s o n (1957) noted t h a t an i n s u l a t i n g b l a n k e t o f a i r b u b b l e s adhered t o t h e h a i r s when t h e s e a l submerged. They f e l t t h a t t h e a i r b u b b l e s , p a r t i c u -l a r l y t h o s e among t h e u n d e r h a i r s , would a s s i s t c o n s e r v a t i o n o f body heat by h e l p i n g t o keep t h e s k i n d r y (Montagna and H a r r i s o n , 1957). Hart and F i s h e r (1964)commented t h a t t h e 46 p r e s e n c e of a i r bubbles i n p h o c i d h a i r d i d not p r e v e n t water from d i r e c t l y c o n t a c t i n g t h e s k i n . Harbour s e a l h a i r f u n c t i o n s i n p r o t e c t i o n of t h e s k i n a g a i n s t m e c h a n i c a l i n j u r y and a g a i n s t s o l a r r a d i a t i o n . I have f r e q u e n t l y o b s e r v e d Phoca v i t u l i n a l y i n g exposed t o s u n l i g h t on s h o r e , f l o a t i n g l o g s , o f f s h o r e r o c k s and i n s h a l l o w t i d e p o o l s . The s k i n r e g i o n w h i c h r e c e i v e s t h e most a b r a s i o n from c o n t a c t w i t h t h e s u b s t r a t e when the s e a l i s ashore i s t h e a n t e r i o r v e n t r a l s u r f a c e . S o k o l o v (1960a) noted t h a t t h e e p i d e r m i s was t h i c k e s t on t h e m i d v e n t r a l a r e a between t h e p e c t o r a l f l i p p e r s . D u r i n g t h e a n n u a l moult t h i s i s t h e e a r l i e s t r e g i o n (except f o r body openings) t o r e p l a c e t h e h a i r . Dunbar (1949) and M a n s f i e l d (1963) have remarked t h a t A t l a n t i c harbour s e a l s are d a r k e s t i n c o l o u r at t h e n o r t h e r n end of t h e i r range. A dark c o l o u r c o u l d f u n c t i o n i n r e t e n t i o n of s o l a r h e a t , however another dark p h o c i d , Monachus s c h a u i n s l a n d i , i s f r e q u e n t l y seen l y i n g on c o r a l sand i n t h e sun on t r o p i c a l Midway and Laysan I s l a n d s ( B a i l e y , 1952). The harbour s e a l of s o u t h e r n C a l i f o r n i a and B a j a C a l i f o r n i a have so h i g h a p r o p o r t i o n of b l a c k c o l o u r p a t t e r n s k i n s t h a t t h e y were f o r m e r l y s e p a r a t e d t a x -o n o m i c a l l y from P. v. r i c h a r d i (Schwarz, 1942; S c h e f f e r and S l i p p , 1944; M i l l e r and K e l l o g g , 1955). 47 I t i s p r o b a b l e t h a t t h e c o l o u r p a t t e r n s i n Phoca  v i t u l i n a had some a d a p t i v e s i g n i f i c a n c e i n t h e p a s t . There i s no r e a s o n t o b e l i e v e t h a t , w i t h t h e e x c e p t i o n of P. v. l a r g h a ' s w h i t e c o a t pup, which i s exposed on i c e ( K i n g , 1964), t h e r e i s now any a d a p t i v e s i g n i f i c a n c e i n t h e d i s t r i b u t i o n o f p e l a g e p a t t e r n s i n t h i s s p e c i e s . The l a c k of a d a p t i v e s i g n i f i c a n c e i n t h e d i s t r i b u t i o n of p e l a g e p a t t e r n s i s a c l u e t o t h e s e p a r a t i o n of P. v. r i c h a r d i i n t o s e p a r a t e b r e e d -i n g s t o c k s . The d i s t r i b u t i o n of p e l a g e t y p e s by a r e a , d e s c r i b e d e a r l i e r i n t h i s r e p o r t , demonstrates t h a t G l a c i e r Bay and v i c i n i t y i n A l a s k a , t h e Queen C h a r l o t t e I s l a n d s and t h e S t r a i t o f G e o r g i a have p o p u l a t i o n s w h i c h d i f f e r g e n e t i c a l l y . Each of t h e s e a r e a s can be c o n s i d e r e d a d i s c r e t e e n t i t y . The narrow c o n t i n e n t a l s h e l f which i s b o r d e r e d by a huge a b y s s a l p l a i n (Uda, 1963) and t h e e c o l o g i c a l c h a r a c t e r i s t i c s of t h e s e a l are r e s p o n s i b l e f o r t h e s e p a r a t i o n of b r e e d i n g p o p u l a -t i o n s ( F i g u r e 15). S c h e f f e r and S p e r r y (1931) r e p o r t e d t h a t harbour s e a l s f e d p r i m a r i l y on f i s h e s and i n v e r t e b r a t e s of s h o a l water bays and s u r f a c e water o f f s h o r e . The s e a l s commonly f e d i n t h e t i d a l a reas about s t r e a m mouths ( S c h e f f e r and S p e r r y , 1931). S c h e f f e r and S l i p p (1944) s t a t e d t h a t t h e harbour s e a l was seldom r e p o r t e d beyond f i f t e e n m i l e s from s h o r e . S p a l d i n g (1964) r e p o r t e d t h a t t h e harbour s e a l was 48 r a r e l y seen more t h a n f i v e m i l e s from shore a l t h o u g h some were o b s e r v e d i n June 1962, w i t h o t h e r s p e c i e s , at t h e P o r t -l o c k Bank i n t h e G u l f of A l a s k a between 30 and 40 m i l e s from s h o r e . The e v i d e n c e s u p p o r t s a view t h a t t h e harbour s e a l i s a s p e c i e s of i n s h o r e and p r i m a r i l y s h a l l o w w a t e r s . The tendency t o use s h a l l o w w a t e r s , c l o s e t o s h o r e , i s t h e p r o b a b l e r e a s o n f o r Phoca v i t u l i n a ' s absence from Guadalupe I s l a n d , B a j a C a l i f o r n i a , w h i c h i s about 170 m i l e s west of t h e m a i n l a n d ( S c h e f f e r , 1958). The narrow l i n e a r h a b i t a t has a l s o d i v i d e d t h e s e a l p o p u l a t i o n i n t o s e p a r a t e gene p o o l s . G l a c i e r Bay i s l o c a t e d at t h e n o r t h e r n end of t h e " I n s i d e Passage" and opens southward i n t o t h e n o r -t h e r n end of t h e A l e x a n d e r A r c h i p e l a g o . The Queen C h a r l o t t e I s l a n d s l i e s o u t h of t h e A l e x a n d e r A r c h i p e l a g o and west of the n o r t h e r n B r i t i s h C olumbia c o a s t and are nowhere c l o s e r t h a n about 30 m i l e s t o e i t h e r t h e A l a s k a n I s l a n d s or t h e o u t e r i s l a n d s of t h e B.C. m a i n l a n d . The S t r a i t of G e o r g i a c o m p r i s e s an e n c l o s e d and r a t h e r l a k e - l i k e waterway about 120 n a u t i c a l m i l e s l o n g , an average of 18 n a u t i c a l m i l e s wide, w i t h a mean depth of 85 fathoms (Waldichuk, 1957). For a n a l y s i s of s e a l p o p u l a t i o n s I have i n c l u d e d Puget Sound and t h e waterways n o r t h t o S o i n t u l a i n t h e " S t r a i t o f Geor-g i a " . Between Olympia, Washington and S o i n t u l a , B.C., GUc'tr Jtr»,t of F i g u r e 15. The B r i t i s h C o lumbia and s o u t h -e a s t e r n A l a s k a c o a s t s . S k e t c h map of key l o c a t i o n s . F i g u r e 16. B l a c k s k i n . Male c o l l e c t e d near S k i d e g a t e , Queen C h a r l o t t e I s l a n d s , l a t e A p r i l , 1965. F i g u r e 17. Common s k i n . Male c o l l e c t e d from Puget Sound, e a r l y May, 1965. F i g u r e 18. Muddy s k i n . Female from t h e Queen C h a r l o t t e I s l a n d s . C o l l e c t e d l a t e J u l y , 1964. 5 3 t h e r e are no n a t u r a l b a r r i e r s or n a t u r a l d e f i l e s r e s t r i c t i n g or c o n s t r i c t i n g the f r e e f l o w of genes i n t h e Phoca v i t u l i n a p o p u l a t i o n . Even though t h e r e i s no t r u e m i g r a t i o n i n h a r -bour s e a l s , t h e wide o v e r l a p of home ranges c o u p l e d w i t h a p r e f e r e n c e f o r c e r t a i n t y p e s of h a b i t a t at v a r i o u s seasons ( S p a l d i n g , 1964) would keep t h e s e a l p o p u l a t i o n of t h e S t r a i t of G e o r g i a a r e a c o n t i n u o u s l y mixed and p r e v e n t t h e i s o l a t i o n of b r e e d i n g groups from t h e remainder of t h e r e g i o n a l popu-l a t i o n . The Queen C h a r l o t t e I s l a n d s form a g e o g r a p h i c a l and e c o l o g i c a l u n i t which p r o b a b l y does have some i n f l u e n c e on t h e p o p u l a t i o n of t h e nearby m a i n l a n d s h o r e . T r a n s f e r of i n d i v i d u a l s between t h e c o a s t a l i s l a n d f r i n g e and t h e Queen C h a r l o t t e I s l a n d s i s p r o b a b l y i r r e g u l a r and i n s m a l l numbers s i n c e t h e average d i s t a n c e would be about f i f t y m i l e s . The t r a n s f e r of g e n e t i c m a t e r i a l between t h e n o r t h and s o u t h a l o n g t h e c o a s t of t h e n o r t h e r n B r i t i s h Columbia mainland and t h e west c o a s t s of Vancouver I s l a n d and Washing-t o n S t a t e must t h e n be l i m i t e d t o a r e l a t i v e t r i c k l e a l o n g t h e narrow edge of t h e c o n t i n e n t a l s h e l f . Areas such as G l a c i e r Bay, t h e Queen C h a r l o t t e I s l a n d s and S t r a i t of Geor-g i a would t h e r e f o r e be r e l a t i v e l y i s o l a t e d c e n t e r s of l a r g e b r e e d i n g p o p u l a t i o n s s e p a r a t e d by s t r e t c h e s of narrow zones of harbour s e a l h a b i t a t . These narrow zones would act as n a t u r a l d e f i l e s and reduce t h e i n c i d e n c e of gene t r a n s f e r s 54 a c r o s s t h e i r a r e a s . F i s h e r ( p e r s o n a l communication) has d e s c r i b e d t h e harbour s e a l of t h e w e s t e r n A t l a n t i c as almost u n i f o r m i n c o l o u r p a t t e r n as c o n t r a s t e d w i t h t h e v a r i a t i o n n o t e d a l o n g t h e P a c i f i c c o a s t . On t h e N o r t h American A t l a n t i c c o a s t t h e e x i s t e n c e of p l e n t i f u l areas s u i t a b l e f o r harbour s e a l havens p l u s a wide, s h a l l o w , c o n t i n e n t a l s h e l f and e x t e n s i v e o f f -s h o re banks p e r m i t s u n r e s t r i c t e d m i x i n g of s t o c k s from t h e e n t i r e c o a s t and p r e v e n t s t h e development of p o p u l a t i o n s of v i s i b l y d i f f e r e n t g e n e t i c makeup. The p o s s i b l e p h y l o g e n e t i c s i g n i f i c a n c e o f e i t h e r i n u t e r o or p o s t - n a t a l s h e d d i n g of t h e w h i t e f o e t a l p e l a g e i s s u g g e s t e d by t h e development of g e o g r a p h i c v a r i a t i o n s w i t h i n a w i d e s p r e a d s p e c i e s h a v i n g r a t h e r weakly s e p a r a b l e sub-s p e c i e s (Ognev, 1935; S c h e f f e r , 1958; K i n g , 1964). White h a i r c o l o u r might have some p r o t e c t i v e f u n c t i o n i f newborn harbour s e a l s , l i k e newborn harp s e a l s , P a g o p h i l u s groen-l a n d i c u s , were temporary non-swimmers who were exposed on i c e (Ognev, 1935; F i s h e r , 1955; K i n g , 1964). F i s h e r (1952) and S c h e f f e r and S l i p p (1944) have commented on t h e swimming a b i l i t y of newborn harbour s e a l s . F i s h e r (1952) noted a pup b o r n on an E c s t a l l R i v e r sandbar, i n an a r e a where n e a r l y a l l p o s t p a r t u m pups wear t h e s p o t t e d c o a t , t h a t was n e a r l y i n v i s i b l e because i t p o s s e s s e d a matted c o a t of y e l l o w i s h -w h i t e lanugo h a i r t h a t b l e n d e d w i t h t h e sand. The c o n d i t i o n s 55 of non-swimming pups exposed on i c e are not met i n any h a r -bour s e a l s u b s p e c i e s except P. v. l a r g h a (Ognev, 1935; W i l k e , 1954; K i n g , 1964). W i l k e (1954) w o r k i n g o f f Hokkaido w i t h Japanese s e a l e r s found t h a t P. v. l a r g h a pups were bor n on f l o a t i n g i c e between l a t e J a n u a r y and m i d - A p r i l and were sub-j e c t t o p r e d a t i o n w h i l e t h e y were on t h e i c e . These pups were i n v a r i o u s s t a g e s of moult i n l a t e A p r i l ( W i l k e , 1954). The p r e s e n c e of a w h i t e p r e - n a t a l c o a t i n a l l t h e r a c e s and t h e o c c a s i o n a l b i r t h of w h i t e pups i n P. v. r i c h a r d i as f a r s o u t h as C a l i f o r n i a i n d i c a t e t h a t w h i t e p o s t - n a t a l pups were once u n i v e r s a l i n Phoca v i t u l i n a s t o c k s . I t i s p r o b a b l e t h a t P. v. l a r g h a d u r i n g t h e P l e i s t o c e n e was l o n g s e p a r a t e d from t h e o t h e r r a c e s of Phoca v i t u l i n a and r e t a i n e d i t s w h i t e j u v e n i l e p o s t - n a t a l p e l a g e due t o r e t e n t i o n or a d o p t i o n of p a c k - i c e pupping areas where t h e w h i t e pup has a s u r v i v a l advantage over t h e c o l o u r e d pup ( S t u t z , 1966). The r e c e n t l y d e s c r i b e d s y m p a t r i c s p e c i e s or sub-s p e c i e s Phoca ( v i t u l i n a ? ) i n s u l a r i s ( B e l k i n , 1964) of t h e K u r i l e I s l a n d s b e a r s a s p o t t e d pup on shore i n May, i s r e p r o d u c t i v e l y i s o l a t e d from P. v. l a r g h a and seems much more l i k e harbour s e a l s i n g e n e r a l . The o b s e r v e d d i f f e r e n c e s i n the w h i t e c o a t h a i r of P. v. r i c h a r d i and P. v. l a r g h a p l u s t h e e x i s t e n c e of Phoca " i n s u l a r i s " i n d i c a t e t h a t i f P. v. l a r g h a i s i n d e e d a s u b s p e c i e s of Phoca v i t u l i n a i t i s a v e r y d i f f e r e n t s e a l from t h e r e s t of t h e s p e c i e s ( S t u t z , 1966). 56 RECOMMENDATIONS A c l o s e d season on harbour s e a l s i n B r i t i s h Colum-b i a i s recommended f o r 15 May t h r o u g h November 15 a n n u a l l y . T h i s i s t h e p e r i o d when t h e m a j o r i t y of harbour s e a l s a r e i n a c t i v e moult and p o s s e s s s k i n s of no comme r c i a l v a l u e . Department of F i s h e r i e s R e g u l a t i o n s s h o u l d e s t a b -l i s h s e a l i n g zones. 1. S t r a i t of G e o r g i a . 2. West c o a s t of Vancouver I s l a n d . 3. M a i n l a n d c o a s t , n o r t h of K n i g h t I n l e t . 4. Queen C h a r l o t t e I s l a n d s . The f o u r r e g i o n s l i s t e d are areas of d i s c r e t e popu-l a t i o n s . Serious r e d u c t i o n of t h e s t o c k s i n any of t h e s e zones would r e s u l t i n l i m i t e d i m m i g r a t i o n f rom o t h e r a r e a s , p r o b a b l y not at a r a t e s u f f i c i e n t t o m a i n t a i n t h e s i z e of a t h r e a t e n e d o r d e p l e t e d p o p u l a t i o n . S h o u l d h i g h p e l t p r i c e s b r i n g heavy h u n t i n g p r e s s u r e on t h e s e a l , s e p a r a t e r e g u l a -t i o n s might be r e q u i r e d f o r c o n s e r v a t i o n o f t h e s t o c k s i n d i f f e r e n t zones. The g e n e t i c background f o r c o l o u r v a r i a t i o n s i n Phoca v i t u l i n a i s c o m p l e t e l y unknown. Any l o n g term program of maintenance and o b s e r v a t i o n of c a p t i v e s e a l s s h o u l d i n c l u d e a b r e e d i n g program, e s t a b l i s h m e n t of a h e r d book, and photo-g r a p h i c r e c o r d s of each s e a l ( p r e f e r a b l y t a k e n at a s t a n d a r d a n g l e at about t h e same date a n n u a l l y ) , t o e s t a b l i s h t h e g e n e t i c p a t t e r n . 58 SUMMARY Phoca v i t u l i n a uses a v a r i e t y of s h a l l o w c o a s t a l and f r e s h w a t e r h a b i t a t s , i s n o n - m i g r a t o r y and r a r e l y f o u n d more t h a n f i f t e e n m i l e s from s h o r e . The s e a l p o s s e s s e s a complete mammalian h a i r system ( l e s s t h e e r e c t o r p i l i muscle) c o n s i s t i n g of guard h a i r s , u n d e r h a i r s , sebaceous and sweat g l a n d s . The h a i r p r o v i d e s p r i m a r i l y m e c h a n i c a l p r o t e c t i o n f o r t h e s k i n r a t h e r t h a n i n s u l -a t i o n a g a i n s t heat l o s s . A f o e t a l moult, n o r m a l l y i n t r a u t e r i n e (except i n P. v. l a r g h a , which may not be a harbour s e a l ) d i f f e r s i n p a t t e r n sequence from t h e a d u l t moult. The a d u l t moult o c c u r s d u r i n g August - October f o r most specimens. There i s some moult a c t i v i t y among a d u l t s d u r i n g almost e v e r y month of t h e y e a r . D i s t r i b u t i o n of two c o l o u r p a t t e r n s , b l a c k and com-mon, i n d i c a t e l o c a t i o n s of d i s c r e t e p o p u l a t i o n s whose e x i s t -ence has management i m p l i c a t i o n s . 59 ACKNOWLEDGEMENTS I am g r a t e f u l t o t h e many b i o l o g i s t s and o t h e r s who have a s s i s t e d me i n t h i s s t u d y . I am p a r t i c u l a r l y g r a t e -f u l f o r a s s i s t a n c e i n problems of h a i r development from Dr. V. B. S c h e f f e r of t h e U.S. F i s h and W i l d l i f e S e r v i c e and Dr. J.K. L i n g of t h e A u s t r a l i a n N a t i o n a l U n i v e r s i t y . S p e c i a l mention must be made of Mr. Henry Hansen, f u r buy e r , L a n g l e y , B.C., who made a l l of h i s p e r s o n a l o b s e r v a t i o n s , s e a l s k i n s , and f a c i l i t i e s a v a i l a b l e f o r s t u d y and who a s s i s t e d i n t h e a c q u i s -i t i o n o f specimens at h i s p e r s o n a l expense; t h e Canada D e p a r t -ment of F i s h e r i e s whose p e r s o n n e l p r o v i d e d much needed t r a n s -p o r t a t i o n , s u p p o r t and a s s i s t a n c e i n t h e f i e l d ; Dr. G.W. Eaton of t h e F a c u l t y of A g r i c u l t u r e , U.B.C, f o r major a s s i s t -ance i n s t a t i s t i c a l a n a l y s i s of d a t a ; and Dr. H.D. F i s h e r who p a t i e n t l y s u p p l i e d a d v i c e and d i r e c t i o n and sometimes seemed a l i t t l e w o r r i e d . 60 LITERATURE CITED B a i l e y , A.M. 1952. The H a w a i i a n monk s e a l . Museum P i c t o r -i a l No. 7, Denver Museum of N a t u r a l H i s t o r y , C o l o r a d o . 32 p. B a s s e t t , C.F. and L.M. L l e w e l l y n . 1948. The m o l t i n g and f u r growth p a t t e r n i n t h e a d u l t s i l v e r f o x . Amer. M i d i . Nat. 39 : 597-601. B e l k i n , A.N. 1964. A new s p e c i e s o f s e a l - Phoca i n s u l a r i s sp. n. - from t h e K u r i l e I s l a n d s . D o k l . Akad. Nauk. SSSR 158 : 1217-1219. B l a i r , W.F. 1957. Mammals, p. 615-774 i n B l a i r , W.F.; A.P. B l a i r ; P. B r o d k o r b ; F.R. C a g l e ; and G.A. Moore. V e r t e -b r a t e s o f t h e U n i t e d S t a t e s . M c G r a w - H i l l Book Co., New York. 819 p. Bennot, P. 1951. 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W i l k e , F. 1954. S e a l s of n o r t h e r n Hokkaido. J . Mammal. 35 : 218-224. A P P E N D I X Appendix A. H o r i z o n t a l s e c t i o n s t h r o u g h f o l l i c l e groups, w i t h h a i r measurements, male. S l i d e g Q £ H a i r d i a m e t e r s i n m i c r a Av. # Range No. p a Over- U n d e r h a i r s Under- Under- Remarks h a i r 1 2 3 4 5 6 h a i r s h a i r s 2a 10/5/64 82^ 5 2 | 45 30 30 3, .28 3-4 average h a i r s 39a 10/5/64 93 30 19| 12 3 2-4 17 a 10/5/64 63 33 30 28§ 21 16| 3 3 21a 10/5/64 60 2 2 i 2. .7 2-3 23b 10/5/64 d i s t o r t i o n s 3. .8 3-4 26b 10/5/64 d i s t o r t i o n s 3. .4 3-4 12a 13/5/64 126 45 39 1 3 j 7 3 2-4 11a 27/5/64 150 54 30 24 18 3. . 14 2-4 .369 mm h a i r s 112b 111b 129b 130b 4/8/64 8/8/64 14/8/64 18/8/64 66 d i s t o r t e d d i s t o r t e d 57 39 1 9 i 1 6 j l\ l\ not measured 133a 18/8/64 6 7 | 30 21 21 3.4 4.3 3.3 3.1 2-4 2- 5 3- 6 3-4 2-4 d u p l i c a t e o v e r h a p r s pigment H a i r s d u p l i c a t e o v e r - and u n d e r h a i r s New h a i r s d e e p l y m e l a n i n s t a i n s OS Appendix B. H o r i z o n t a l s e c t i o n s t h r o u g h f o l l i c l e groups, w i t h h a i r measurements, f e m a l e s S l i d e H a i r d i a m e t e r s i n m i c r a Av. # Range No. Spec.Date o v e r - U n d e r h a i r s Under- Under- Remarks h a i r 1 2 3 4 5 6 h a i r s h a i r s 10 a 1/5/64 66 45 39 36 10 a 1/5/64 120 45 33 18 3 ,2 3-4 10 a 1/5/64 7 9 | 45 33 21 l b 10/5/64 66 37§ 33 15 3 ,3 13a 10/5/64 90 4 6 | 30 15 3 .8 3-5 18a 10/5/64 8 2 j 42 30 21 9 4. . 1 4-5 22a 10/5/64 63 36 24 21 12 2 .7 1-4 27 a 10/5/64 78 24 16| 12 3 2-5 36a 10/5/64 7 6 | 45 36 21 12 10| 4 . 1 4-5 40b 10/5/64 60 36 27 24 1 3 | 3 .7 2-4 37 a 12/5/64 7 6 | 19^ 15 16^ 2 .2 1-3 6b 13/5/64 6 7 | 45 22£ 1 3 j 4 8b 16/5/64 69 3 7 | 30 21 15 9 8b 16/5/64 7 9 j 45 2 2 j 22§ 15 240b 30/6/65 4 9 j 2 5 | 16| 1 3 j 12 3. .03 2-4 135a 1/8/64 8 2 | 54 45 33 21 4 3-5 136a 4/8/64 d i s t o r t e d 3. .75 2-5 137 a 4/8/64 33 21 15 7 2. .6 2-3 118b 6/8/64 67 i 48 24 12 3 1-5 118b 6/8/64 45 102a 6/8/64 78 39 2 5 j 24 7 2. .9 2-4 121a 6/8/64 90 16| 15 12 3. 2 2-4 104a 8/8/64 111 45 36 30 3. 2 2-4 106b 8/8/64 99 24 1 6 i 1 3 j 9 3. 6 2-4 124b 12/8/64 42 2 5 | 24 18 3. 5 3-4 127 a 14/8/64 78 22 | d i s t o r t e d 3 2-4 131a 18/8/64 33 2 2 j 3 2-4 e p i d e r m i s s u r f a c e two s e c t i o n s from below s u r f a c e 294 m i c r a between o v e r h a i r s BIB specimen. 6 r s . 10 mos. age average d i s t a n c e 6 o v e r h a i r s , 76 m i c r a . Pup of t h e year d u p l i c a t e o v e r h a i r s new o v e r h a i r round o l d o v e r h a i r f l a t t e n e d 05 67 Appendix C. Dates of moult s t a g e s , male. S l i d e No. .Spec.Date Age Stage A r e a B24B 3/1/65 0: 6 t e l o g e n Ganges Harbour G12B 12/1/65 4: 6 t e l o g e n J e r v i s I n l e t B45B 14/1/65 11: 6 t e l o g e n G a l i a n o S. G15B 15/1/65 4: 6 t e l o g e n J e r v i s I n l e t G17B 18/1/65 7: 6 t e l o g e n P r i n c e s s L o u i s e I n l e t B46B 25/1/65 5: 6 t e l o g e n P o r t l a n d I s l a n d B47B 26/1/65 6: 6 t e l o g e n P o r t l a n d I s l a n d B26B 15/2/65 0:6-8 t e l o g e n G u l f I s l a n d s H4B 18/2/65 4: 7 t e l o g e n Toba I n l e t H7B 20/2/65 3: 7 t e l o g e n P r y c e Channel H8B 20/2/65 14: 7 t e l o g e n P r y c e Channel B36B 28/2/65 2: 7 t e l o g e n Mayne I s l a n d B40B 6/3/65 9: 8 t e l o g e n Tumbo I s l a n d 178b 7/3/65 ' t e l o g e n C r o f t e n B42B 7/3/65 0: 8 t e l o g e n B e l l e I . C h a i n H15B 12/3/65 4: 8 t e l o g e n K n i g h t I n l e t H16B 12/3/65 9: 8 t e l o g e n K n i g h t I n l e t H14B 12/3/65 3: 8 t e l o g e n K n i g h t I n l e t H17B 12/3/65 7: 8 t e l o g e n K n i g h t I n l e t H40B 17/3/65 4: 8 t e l o g e n K n i g h t I n l e t H28B 19/3/65 0: 8 t e l o g e n K n i g h t I n l e t H34B 20/3/65 3: 8 t e l o g e n K n i g h t I n l e t H35B 20/3/65 6: 8 t e l o g e n K n i g h t I n l e t H37B 20/3/65 6: 8 t e l o g e n K n i g h t I n l e t H29B 20/3/65 1: 8 t e l o g e n K n i g h t I n l e t H38B 20/3/65 3: 8 t e l o g e n K n i g h t I n l e t H24B 22/3/65 19: 8 t e l o g e n K n i g h t I n l e t B52B 10/4/65 3: 9 t e l o g e n Tumbo I s l a n d B50B 10/4/65 0: 9 t e l o g e n B e l l e I . C h a i n B51B 10/4/65 0: 9 t e l o g e n B e l l e I . C h a i n B53B 10/4/65 3: 9 t e l o g e n B e l l e I . C h a i n B54B 10/4/65 t e l o g e n Tumbo I s l a n d B55B 11/4/65 2: 9 anagen Cowichan Bay B57B 11/4/65 0: 9 t e l o g e n Cowichan Bay C7B 16/4/65 0: 9 t e l o g e n N i t i n a t Lake C16B 17/4/65 0: 9 t e l o g e n N i t i n a t Lake 298a 25/4/65 t e l o g e n Q.C.I. 308b 25/4/65 t e l o g e n Q.C.I. 273a 25/4/65 ear l y Q.C.I. 290 a 29/4/65 t e l o g e n Q.C.I. Appendix C (cont'd.) S l i d e No. Spec.Date Age Stage A r e a E7B 3/5/65 1: 10 anagen S t e v e s t o n 5b 5/5/64 anagen P r i n c e Rupert B65B 8/5/65 2: 10 t e l o g e n P o r t l a n d I s l a n d B63B 8/5/65 1: 10 anagen E. Sydney I s l a n d 33a 10/5/64 t e l o g e n S u l l i v a n Bay 34a 10/5/64 t e l o g e n Q.C.I. 42b 10/5/64 t e l o g e n P e t e r s b u r g 2a 10/5/64 t e l o g e n Q.C.I. 39b 10/5/64 t e l o g e n Q.C.I. 15a 10/5/64 t e l o g e n P e t e r s b u r g 12c 13/5/64 t e l o g e n U c l u e l e t B70B 15/5/65 3: 10 t e l o g e n G a b r i o l a Pass B68B 15/5/65 7: 10 t e l o g e n G a b r i o l a P ass 35b 18/5/64 2: 10 t e l o g e n G a l i a n o I s l a n d 9a 18/5/64 5: 10 anagen Wise I s l a n d 301a 24/5/65 anagen Q.C.I. 288a 24/5/65 t e l o g e n Q.C.I. 4a 27/5/64 t e l o g e n C ampbell R i v e r l i b 27/5/64 t e l o g e n Campbell R i v e r B77B 29/5/65 2: 10 t e l o g e n Pender I s l a n d B78B 30/5/65 2: 10 t e l o g e n A c t i v e Pass 65a 5/6/64 t e l o g e n Q.C.I. 70b 10/6/64 t e l o g e n G l a c i e r Bay 52b 10/6/64 t e l o g e n G l a c i e r Bay 59b 10/6/64 t e l o g e n Q.C.I. 39b 10/6/64 t e l o g e n Q.C.I. A10B 10/6/65 0: 11 t e l o g e n P o r t Moody 240b 30/6/65 new pup t e l o g e n Q.C.I. 241a 30/6/65 t e l o g e n Q.C.I. 243b 30/6/65 t e l o g e n Q.C.I. 245a 30/6/65 t e l o g e n Q.C.I. 82b 30/7/64 t e l o g e n Q.C.I. 90b 30/7/64 t e l o g e n Q.C. I. 112a 4/8/64 anagen G l a c i e r Bay 151a 4/8/64 anagen G l a c i e r Bay 141a 4/8/64 anagen G l a c i e r Bay 98a 6/8/64 anagen R i v e r s I n l e t 143b 6/8/64 anagen Q.C.I. 144a 6/8/64 anagen R i v e r s I n l e t 120a 6/8/64 anagen R i v e r s I n l e t Appendix C (cont'd.) S l i d e No. Spec.Date Age Stage A r e a 110b 8/8/64 anagen C l o - o o r e 122a 8/8/64 anagen K n i g h t I n l e t 111a 8/8/64 anagen K n i g h t I n l e t 123a 8/8/64 anagen K n i g h t I n l e t B11B 8/8/64 6: 1 anagen Sydney I s l a n d 109a 8/8/64 anagen K n i g h t I n l e t 132b 18/8/64 anagen Q.C.I. 134b 18/8/64 anagen Q.C.I. 130 a 18/8/64 anagen Q.C.I. FIB 31/8/64 0: 2 t e l o g e n K i t imat F2B 31/8/64 0: 2 t e l o g e n K i t imat F3B 2/9/64 3: 3 anagen K i l d a l e Arm mouth F5B 2/9/64 2: 3 anagen K i l d a l e Arm mouth 158a 24/10/64 t e l o g e n P o w e l l R i v e r 163a 24/10/64 t e l o g e n P o w e l l R i v e r 167b 15/11/64 t e l o g e n Q.C.I. 168a 15/11/64 t e l o g e n J e r v i s I n l e t 17 5b 28/11/64 t e l o g e n P r i n c e Rupert 182a 28/11/64 t e l o g e n P r i n c e Rupert G6B 7/12/64 3: 5 t e l o g e n J e r v i s I n l e t 70 Appendix D. Dates o f moult s t a g e s , f e m a l e . S l i d e No. Spec.Date Age Stage A r e a G13B 12/1/65 3: 6 t e l o g e n J e r v i s I n l e t G14B 14/1/65 3: 6 t e l o g e n J e r v i s I n l e t B25B 17/1/65 19: 6 t e l o g e n C r o f t e n G18B 20/1/65 3: 6 t e l o g e n J e r v i s I n l e t B48B 2/2/65 3: 7 t e l o g e n G a l i a n o I s l a n d 177a 6/2/65 t e l o g e n Johnson S t r a i t B34B 14/2/65 28: .7 v e r y e a r l y C r o f t e n anagen H1B 14/2/65 3: 7 t e l o g e n Savory I s l a n d H2B 18/2/65 0: 7 t e l o g e n Toba I n l e t H3B 18/2/65 2: 7 t e l o g e n Toba I n l e t H5B 18/2/65 0: 7 t e l o g e n Toba I n l e t H10B 22/2/65 15: 7 t e l o g e n C o r d e r o Channel H U B 24/2/65 19: 7 t e l o g e n C o r d e r o Channel H12B 24/2/65 6: 7 t e l o g e n C o r d e r o Channel H13B 24/2/65 2: 7 t e l o g e n C o r d e r o Channel 43a 26/2/64 5: 7 t e l o g e n I n d i a n Arm 45a 26/2/64 18: .7 t e l o g e n I n d i a n Arm B30B 1/3/65 t e l o g e n G a b r i o l a P ass B37B 1/3/65 1: 8 t e l o g e n G a b r i o l a Pass B38B 1/3/65 2: 8 t e l o g e n G a b r i o l a P ass B41B 6/3/65 13: 8 t e l o g e n Tumbo I s l a n d H18B 15/3/65 8: 8 t e l o g e n K n i g h t I n l e t H19B 15/3/65 3: 8 t e l o g e n K n i g h t I n l e t H20B 15/3/65 11: 8 t e l o g e n K n i g h t I n l e t H21B 16/3/65 0: 8 t e l o g e n K n i g h t I n l e t H41B 16/3/65 3: 8 t e l o g e n K n i g h t I n l e t H39B 17/3/65 0: 8 t e l o g e n K n i g h t I n l e t H23B 18/3/65 10: 8 t e l o g e n K n i g h t I n l e t H22B 18/3/65 3: 8 t e l o g e n K n i g h t I n l e t H31B 19/3/65 9: 8 t e l o g e n s K n i g h t I n l e t H30B 20/3/65 0: 8 t e l o g e n K n i g h t I n l e t H36B 20/3/65 1: 8 t e l o g e n K n i g h t I n l e t H32B 20/3/65 1: 8 t e l o g e n K n i g h t I n l e t H27B 22/3/65 6: 8 t e l o g e n K n i g h t I n l e t H25B 22/3/65 0: 8 t e l o g e n K n i g h t I n l e t H26B 22/3/65 0: 8 t e l o g e n K n i g h t I n l e t B79B 30/3/65 3: 10 t e l o g e n Stewart I . , Washington D12B 8/4/64 0: 9 t e l o g e n Squamish R i v e r mouth (Howe Sound) B56B 11/4/65 0: 9 t e l o g e n Cowichan Bay 71 Appendix D (cont'd.) S l i d e No. Spec.Date Age St age A r e a D13B 11/4/65 0: 9 t e l o g e n Squamish R i v e r mouth C6B 16/4/65 12: 9 t e l o g e n N i t i n a t Lake C12B 16/4/65 9: 9 t e l o g e n N i t i n a t Lake C8B 16/4/65 1: 9 t e l o g e n N i t i n a t Lake C9B 16/4/65 1: 9 t e l o g e n N i t i n a t Lake C14B 17/4/65 0: 9 t e l o g e n N i t i n a t Lake C15B 17/4/65 0: 9 t e l o g e n N i t i n a t Lake C17B 17/4/65 9: 9 t e l o g e n N i t i n a t Lake C18B 17/4/65 12: 9 t e l o g e n N i t i n a t Lake C19B 17/4/65 12: 9 t e l o g e n N i t i n a t Lake C13B 17/4/65 5: 9 t e l o g e n N i t i n a t Lake B60B 19/4/65 t e l o g e n Cowichan Bay 286b 29/4/65 anagen Q.C.I. 289a 29/4/65 t e l o g e n Q.C.I. 294b 29/4/65 t e l o g e n Q.C.I. 303a 29/4/65 t e l o g e n Q.C.I. 7a 5/5/64 anagen P r i n c e Rupert 87 a 8/5/64 0: 10 anagen I n d i a n Arm B66B 8/5/65 0: 10 t e l o g e n C r o f t e n B61B 8/5/65 5: 10 t e l o g e n C o a l I s l a n d l b 10/5/64 t e l o g e n W r a n g e l l 36a 10/5/64 t e l o g e n P e t e r s b u r g 31a 10/5/64 t e l o g e n Q.C.I. 29b 10/5/64 t e l o g e n P e t e r s b u r g 14a 10/5/64 anagen P e t e r s b u r g 22a 10/5/64 t e l o g e n P e t e r s b u r g 41a 10/5/64 t e l o g e n Q.C.I. 6a 13/5/64 t e l o g e n U c l u e l e t 25b 13/5/64 anagen U c l u e l e t 30b 13/5/64 t e l o g e n U c l u e l e t 8a 18/5/64 6: 10 t e l o g e n Moresby I s l a n d B72B 21/5/65 12: 10 t e l o g e n Cowichan Bay B7B 22/5/64 1: 10 t e l o g e n Ada I . o f f Nanoose P t . B7 5B 23/5/65 8: 10 t e l o g e n Swartz Bay B74B 23/5/65 9: 10 anagen T h e t i s I . 292a 24/5/65 anagen Q.C.I. 3a 27/5/64 t e l o g e n Campbell R i v e r B76B 29/5/65 4: 10 t e l o g e n S a l t s p r i n g I s l a n d B79B 30/5/65 3: 10 t e l o g e n Stewart I . , Washington 221a 24/5/65 new pup t e l o g e n B81B 5/6/65 0: 11 t e l o g e n Tumbo I s l a n d Appendix D (cont'd.) S l i d e No. Spec.Date Age Stage A r e a 53a 5/6/64 t e l o g e n Q.C.I. 50b 5/6/64 t e l o g e n Q.C.I. 54b 5/6/64 t e l o g e n Q.C. I . 56 a 5/6/64 t e l o g e n Q.C.I. 60a 5/6/64 t e l o g e n G l a c i e r Bay 71a 5/6/64 t e l o g e n ~ Q.C.I. 46b 10/6/64 t e l o g e n G l a c i e r Bay 48a 10/6/64 anagen G l a c i e r Bay 49a 10/6/64 t e l o g e n G l a c i e r Bay 67a 10/6/64 t e l o g e n G l a c i e r Bay 73a 20/6/64 t e l o g e n Juneau 78a 20/6/64 t e l o g e n Juneau 58b 20/6/64 t e l o g e n Juneau 61a 20/6/64 t e l o g e n Juneau 55b 20/6/64 t e l o g e n Juneau 51a 20/6/64 t e l o g e n Juneau A11B 26/6/64 5: 11 t e l o g e n I n d i a n Arm 240b 30/6/65 Pup t e l o g e n Q.C.I. 81b 81b 25/7/64 t e l o g e n G l a c i e r Bay 86x 25/7/64 t e l o g e n G l a c i e r Bay 83 a 30/7/64 anagen Q.C.I. 85a 30/7/64 anagen Q.C.I. 89a 30/7/64 anagen Q.C.I. 88a 30/7/64 anagen Q.C.I. 84a 30/7/64 anagen Q.C.I. 93a 1/8/64 anagen Q.C.I. 92b 1/8/64 anagen Q.C.I. 135b 1/8/64 anagen Q.C.I. 96b 1/8/64 anagen Q.C.I. 142b 2/8/64 1: 10 t e l o g e n F r a s e r : R i v e r 128b 3/8/64 1: 10 anagen S t e v e s t o n 136a 4/8/64 anagen G l a c i e r Bay 114b 4/8/64 anagen G l a c i e r Bay 115b 4/8/64 anagen S o i n t u l a 113a 4/8/64 anagen G l a c i e r Bay 95a 4/8/64 anagen G l a c i e r Bay 149b 4/8/64 anagen P r i n c e : Rupert 97 a 4/8/64 anagen G l a c i e r Bay 148a 4/8/64 anagen G l a c i e r Bay B10B 6/8/64 t e l o g e n Sechum ] Harbour 99b 6/8/64 anagen Q.C.I. 73 Appendix D (cont'd.) S l i d e No. Spec.Date Age Stage A r e a 119a 6/8/64 anagen R i v e r s I n l e t 150b 6/8/64 anagen R i v e r s I n l e t 101a 6/8/64 anagen R i v e r s I n l e t 138a 6/8/64 anagen Q.C.I. 152 a 6/8/64 anagen Q.C.I. 140b 6/8/64 anagen Q.C.I. 103b 6/8/64 anagen Q.C.I. 118a 6/8/64 anagen Q.C.I. 117 a 6/8/64 anagen Q.C.I. 104b 8/8/64 anagen K n i g h t I n l e t 145b 8/8/64 anagen K n i g h t I n l e t 105a 8/8/64 anagen K n i g h t I n l e t 106b 8/8/64 anagen K n i g h t I n l e t 147 a 14/8/64 anagen Sydney-155a 14/8/64 t e l o g e n Sydney 153b 14/8/64 anagen Sydney 126a 14/8/64 anagen Sydney 125b 14/8/64 t e l o g e n Sydney 131b 18/8/64 anagen Q.C.I. 132a 18/8/64 anagen Q.C.I. F4B 2/9/64 3 3 anagen K i l d a l e Arm mouth F10B 17/9/64 3: 3 anagen Gardner C a n a l F11B 17/9/64 0 3 t e l o g e n K i t l o p e Lake F12B 18/9/64 3 3 anagen K i t l o p e R i v e r mouth 162a 16/10/64 t e l o g e n Q.C.I. 160a 25/10/64 t e l o g e n Q.C.I. 165a 15/11/64 t e l o g e n J e r v i s I n l e t 166a 15/11/64 t e l o g e n J e r v i s I n l e t 191a 17/11/64 anagen P e l i c a n Bay 170 a 28/11/64 t e l o g e n P r i n c e Rupert 172a 28/11/64 t e l o g e n P r i n c e Rupert G7B 9/12/64 13: 5 t e l o g e n P r i n c e s s L o u i s a I n l e t G9B 11/12/64 1: 5 t e l o g e n J e r v i s I n l e t B20B 20/12/64 1: 5 t e l o g e n F u l f o r d Harbour B21B 25/12/64 6: 5 t e l o g e n C r o f t o n B23B 28/12/64 3: 5 t e l o g e n Nanaimo Appendix E. Date of moult s t a g e s , sex undetermined. Spec. Date Age Stage A r e a 16a 5/5/64 t e l o g e n F r a s e r R i v e r 19a 5/5/64 t e l o g e n S i t k a 186b 17/11/64 t e l o g e n P e l i c a n Bay 

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