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Status, range, and habitat of the cascade mantled ground squirrel, Spermophilus Saturatus, in British… Leung, Maria Ching-Yee 1991

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S T A T U S , R A N G E , AND H A B I T A T OF T H E CASCADE MANTLED GROUND S Q U I R R E L , SPERMOPHILUS S A T U R A T U S . I N . B R I T I S H COLUMBIA b y MARIA C H I N G - Y E E LEUNG B . S c . ( H o n s . ) , T h e U n i v e r s i t y o f G u e l p h , 1987 A T H E S I S SUBMITTED IN P A R T I A L F U L F I L L M E N T OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF S C I E N C E i n THE F A C U L T Y OF GRADUATE S T U D I E S ( D e p a r t m e n t o f A n i m a l S c i e n c e ) We a c c e p t t h e t h e s i s a s c o n f o r m i n g t o t h e r e q u i r e d s t a n d a r d THE U N I V E R S I T Y OF B R I T I S H COLUMBIA A p r i l 1991 @ M a r i a C h i n g - Y e e L e u n g In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of The University of British Columbia Vancouver, Canada Date 30 April DE-6 (2/88) A B S T R A C T I n t h i s s t u d y , I i n v e s t i g a t e d t h r e e a s p e c t s p e r t a i n i n g t o t h e c o n s e r v a t i o n o f t h e C a s c a d e m a n t l e d g r o u n d s q u i r r e l ( S p e r m o p h i l u s s a t u r a t u s ) . T h e s e w e r e : 1) I t s t a x o n o m i c r e l a t i o n s h i p t o i t s s i b l i n g s p e c i e s , S . l a t e r a l i s ; 2) I t s r a n g e a n d d i s t r i b u t i o n i n B r i t i s h C o l u m b i a ; a n d 3) A p r e l i m i n a r y d e s c r i p t i o n o f m a j o r c o m p o n e n t s i n S . s a t u r a t u s h a b i t a t i n B r i t i s h C o l u m b i a . T h e t a x o n o m i c r e l a t i o n s h i p o f S . s a t u r a t u s t o S . l a t e r a l i s was d e t e r m i n e d b y c o m p a r i n g s k u l l c h a r a c t e r i s t i c s a n d a l l e l e f r e q u e n c i e s o f b l o o d e n z y m e s . T h e s k u l l o f S . s a t u r a t u s i s l a r g e r a n d b r o a d e r t h a n i t s t h r e e n e i g h b o u r i n g S . l a t e r a l i s s u b s p e c i e s . S . s a t u r a t u s h a d two a l l e l e s t h a t w e r e a b s e n t i n S . l a t e r a l i s a n d d i d n o t show s e g r e g a t i o n i n a n y o f t h e l o c i w h e r e a s S . l a t e r a l i s d i d show p o l y m o r p h i s m . Some S . l a t e r a l i s w e r e more c l o s e l y r e l a t e d t o S . s a t u r a t u s t h a n t h e y w e r e t o o t h e r S . l a t e r a l i s a c c o r d i n g t o s t a n d a r d c o e f f i c i e n t s o f g e n e t i c d i s t a n c e ( R o g e r s ' s i m i l a r i t y a n d N e i ' s d i s t a n c e ) , b u t t h e s e came f r o m t h e l o c a t i o n s f u r t h e s t f r o m S . s a t u r a t u s . T h e t a x o n o m i c s t a t u s f o r S . s a t u r a t u s i s c o n f i r m e d f r o m t h e m o r p h o l o g i c a l d i f f e r e n c e s , b u t i s m a i n l y v a l i d a t e d b y t h e p r e s e n c e o f two u n i q u e a l l e l e s , t o g e t h e r w i t h g e o g r a p h i c i s o l a t i o n f r o m S . l a t e r a l i s . T h e r a n g e a n d d i s t r i b u t i o n o f S . s a t u r a t u s w e r e d e t e r m i n e d f r o m museum r e c o r d s , f r o m s u r v e y f o r m s , b y r e c o r d i n g o p p o r t u n i s t i c o b s e r v a t i o n s , b y c o n d u c t i n g s t a t i o n a r y c o u n t s / w a l k i n g t r a n s e c t s , a n d b y l i v e - t r a p p i n g . S . s a t u r a t u s w e r e f o u n d f u r t h e r n o r t h a n d w e s t t h a n p r e v i o u s l y r e p o r t e d , b u t n o t a s f a r e a s t a s i n d i c a t e d b y museum r e c o r d s . S . s a t u r a t u s a l s o a p p e a r e d t o b e uncommon t h r o u g h o u t i t s r a n g e i n B r i t i s h C o l u m b i a . D r a i n a g e s y s t e m s t h a t d e l i n e a t e i t s r a n g e , a n d d i s p l a c e m e n t b y i n v a d i n g c o m p e t i t o r s p e c i e s s h o u l d b e e x a m i n e d a s p r o b a b l e f a c t o r s l i m i t i n g t h e i r d i s p e r s a l . S u r v e y m e t h o d s u s e d w e r e e v a l u a t e d a n d I make r e c o m m e n d a t i o n s f o r t h e i r a p p l i c a t i o n i n f u t u r e s u r v e y s . T h e p e r c e n t c o v e r o f d i f f e r e n t p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s ( e . g . r o c k s ) was v i s u a l l y e s t i m a t e d a t f i v e n a t u r a l l o c a t i o n s (no f o o d s u p p l e m e n t e d ) a n d f o u r f o o d s u p p l e m e n t e d l o c a t i o n s w h e r e S . s a t u r a t u s w e r e p r e s e n t . T h e s e c o v e r t y p e s w e r e p l a c e d i n t h e p e r c e i v e d f u n c t i o n a l c o m p o n e n t s o f " n a t u r a l f o o d s p e c i e s " , " b u r r o w i n g s i t e s " , " p e r c h e s " a n d " p r o t e c t i v e c o v e r " . T h i s g r o u p i n g i n t o f u n c t i o n a l c o m p o n e n t s p r o v i d e s a p r e l i m i n a r y d e s c r i p t i o n o f S . s a t u r a t u s h a b i t a t . A l t h o u g h t h i s s t u d y p r o v i d e s i n f o r m a t i o n o n t h e t a x o n o m i c s t a t u s , r a n g e , d i s t r i b u t i o n , a n d h a b i t a t o f S . s a t u r a t u s i n B r i t i s h C o l u m b i a a n d r e s u l t s i n a r e c o m m e n d a t i o n f o r d e s i g n a t i n g S . s a t u r a t u s a " v u l n e r a b l e s p e c i e s " , f u r t h e r r e s e a r c h i s n e e d e d on t h e s e t o p i c s t h r o u g h o u t i t s f u l l r a n g e , i n c l u d i n g W a s h i n g t o n S t a t e . T h e s e d a t a a r e n e c e s s a r y t o d e t e r m i n e w h e t h e r S . s a t u r a t u s i n B r i t i s h C o l u m b i a a r e r e p r e s e n t a t i v e o f t h e s p e c i e s a n d w h a t e f f o r t s a r e n e e d e d f o r t h e i r c o n s e r v a t i o n . T A B L E OF CONTENTS i v ABSTRACT i i T A B L E OF CONTENTS i v L I S T OF T A B L E S v i L I S T OF F I G U R E S . v i i ACKNOWLEDGEMENTS v i i i CHAPTER 1: G e n e r a l I n t r o d u c t i o n T h e C a s c a d e M a n t l e d G r o u n d S q u i r r e l 1 O b j e c t i v e s 4 CHAPTER 2: Taxonomy o f t h e S p e r m o p h i l u s s a t u r a t u s -S . l a t e r a l i s S p e c i e s C o m p l e x I n t r o d u c t i o n 6 M e t h o d s E l e c t r o p h o r e t i c A n a l y s i s . . . 8 M o r p h o l o g i c a l A n a l y s i s 10 R e s u l t s E l e c t r o p h o r e t i c A n a l y s i s 12 M o r p h o l o g i c a l A n a l y s i s 15 D i s c u s s i o n 19 CHAPTER 3: R a n g e a n d D i s t r i b u t i o n o f S p e r m o p h i l u s s a t u r a t u s i n B r i t i s h C o l u m b i a I n t r o d u c t i o n 24 M e t h o d s 25 R e s u l t s 27 D i s c u s s i o n R a n g e a n d D i s t r i b u t i o n 34 A s s e s s m e n t o f S u r v e y M e t h o d s 41 CHAPTER 4: V a r i a t i o n i n M a j o r H a b i t a t C o m p o n e n t s o f S p e r m o p h i l u s s a t u r a t u s I n t r o d u c t i o n 46 M e t h o d s 47 R e s u l t s 51 D i s c u s s i o n 53 CHAPTER 5: G e n e r a l . D i s c u s s i o n a n d R e c o m m e n d a t i o n s 56 L I T E R A T U R E C I T E D 60 A p p e n d i x 1. T h e museum s p e c i m e n s e x a m i n e d 68 A p p e n d i x 2 . T h e s u r v e y f o r m 74 A p p e n d i x 3 . O r g a n i z a t i o n s a n d p e r s o n s t o whom s u r v e y f o r m s w e r e d i s t r i b u t e d 7 5 A p p e n d i x 4 . L o c a t i o n s w h e r e S . s a t u r a t u s was s i g h t e d 76 A p p e n d i x 5 . L o c a t i o n s s u r v e y e d w h e r e no S . s a t u r a t u s was s i g h t e d 7 8 A p p e n d i x 6. T r a p p i n g e f f o r t , l o c a t i o n s , a n d m a m m a l i a n s p e c i e s c a p t u r e d 82 A p p e n d i x 7 . T h e p e r c e n t c o v e r o f p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s f o r h e r b , s h r u b a n d t r e e l a y e r 83 v i L I S T OF T A B L E S T a b l e 2 . 1 . T a b l e 2 . 2 . T a b l e 2 . 3 . T a b l e 2 . 4 . T a b l e 2 . 5 . T a b l e 3 . 1 . T a b l e 4 . 1 . T a b l e 4 . 2 . T a b l e 5 . 1 . A l l e l i c v a r i a t i o n a t e i g h t p o l y m o r p h i c l o c i f o r S . s a t u r a t u s a n d S . l a t e r a l i s a t d i f f e r e n t l o c a t i o n s . 13 M e a s u r e s o f g e n e t i c r e l a t e d n e s s f o r S . s a t u r a t u s a n d S . l a t e r a l i s . 14 T h e mean , s t a n d a r d d e v i a t i o n , a n d s a m p l e s i z e f o r t e n c r a n i a l m e a s u r e m e n t s o f S . s a t u r a t u s a n d t h r e e S . l a t e r a l i s s u b s p e c i e s 16 D i f f e r e n c e b e t w e e n s e x a n d s p e c i e s f o r t e n s k u l l m e a s u r e m e n t s u s i n g A n a l y s i s o f C o v a r i a n c e . 17 D i f f e r e n c e s b e t w e e n S . s a t u r a t u s a n d t h r e e S . l a t e r a l i s s u b s p e c i e s f o r e a c h c r a n i a l c h a r a c t e r i s t i c s 18 P e r c e n t o f t o t a l f i r s t s i g h t i n g s o c c u r r i n g w i t h i n d i f f e r e n t 10 m i n u t e p e r i o d s a t f o u r l o c a t i o n s . . . 32 T h e g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s i n c l u d e d i n t h e f u n c t i o n a l c o m p o n e n t s o f S . s a t u r a t u s h a b i t a t 50 P e r c e n t c o v e r o f f u n c t i o n a l c o m p o n e n t s 52 R e c o m m e n d a t i o n s f o r t h e c o n s e r v a t i o n o f S p e r m o p h i l u s s a t u r a t u s 59 v i i .. L I S T OF F I G U R E S F i g u r e 1 . 1 . T h e r a n g e o f S p e r m o p h i l u s s a t u r a t u s a n d S . l a t e r a l i s . . . 2 F i g u r e 2 . 1 . Map s h o w i n g l o c a t i o n s o f b l o o d s a m p l i n g 9 F i g u r e 2 . 2 . L a t e r a l , v e n t r a l a n d d o r s a l v i e w s o f S . s a t u r a t u s s k u l l s h o w i n g t e n d i m e n s i o n s t h a t w e r e m e a s u r e d 11 F i g u r e 3 . 1 . Map o f s u r v e y a r e a 28 F i g u r e 3 . 2 . A g e a n d s e x c o m p o s i t i o n o f S . s a t u r a t u s a t f o u r l o c a t i o n s 30 F i g u r e 3 . 3 . P r o b a b i l i t y o f s i g h t i n g S . s a t u r a t u s 33 F i g u r e 3 . 4 . T h e p r o p o r t i o n o f v i s i t s t o p a r t i c u l a r a l t i t u d e s i n s i d e a n d o u t s i d e t h e r a n g e o f S . s a t u r a t u s . . . 35 F i g u r e 3 . 5 . T h e p r o p o r t i o n o f v i s i t s t o p a r t i c u l a r a l t i t u d e s i n s i d e t h e r a n g e o f S . s a t u r a t u s t h a t r e s u l t e d i n s i g h t i n g s 3 6 F i g u r e 4 . 1 . A s c h e m a t i c d i a g r a m s h o w i n g t h e p l a c e m e n t o f a s e t o f f i v e p l o t s u s e d f o r s a m p l i n g h a b i t a t . . . . 49 v i i i ACKNOWLEDGEMENTS D u r i n g t h i s p r o j e c t , I b e n e f i t t e d f r o m t h e h e l p o f many i n d i v i d u a l s a n d o r g a n i z a t i o n s . I w o u l d l i k e t o t h a n k C h r i s Dodd ( B . C . M i n i s t r y o f E n v i r o n m e n t ) f o r i n i t i a t i n g my i n t e r e s t i n t h i s p r o j e c t , a n d Bob Moody ( B . C . C o n s e r v a t i o n F o u n d a t i o n ) , Bob S c h e e r , R i k Simmons a n d B i l l M e r i l l e e s ( B . C . M i n i s t r y o f P a r k s ) f o r t h e i r i n t e r e s t a n d i n v o l v e m e n t . I am g r a t e f u l t o Wendy G o e d e m o n d t , R o l a n d A l t m a n n , Don R e i d a n d D a r i n B e n n e t t f o r t h e i r company a n d f i e l d a s s i s t a n c e . H o s p i t a l i t y a n d s h e l t e r p r o v i d e d b y B . C . P a r k s s t a f f R a n d y H a n a h , Wayne S o b o o l , Doug Crow a n d T o d H a u g h t o n , a n d t h e M e r r i t t S n o w m o b i l e C l u b was w a r m i n g l y a p p r e c i a t e d . I t h a n k T e r r y G a l l o w a y , Tom W e l l s a n d B r u c e B e n n e t t f o r t h e i r e x p e r t i s e . My g r a t i t u d e e x t e n d s t o J e a n D ' A n g e l o a n d L i l l i a n D u n b a r ( R o y a l I n l a n d H o s p i t a l ) , R i c k S m i t h , C h r i s H a r v e y - C l a r k e , B e r n i e May ( C L E E G ) , K e n M o r g a n , G . J . K e n a g y a n d C a t h l e e n N i c h o l s f o r t h e i r h e l p i n t h e c o l l e c t i o n o f e l e c t r o p h o r e t i c d a t a . I a l s o e x p r e s s my t h a n k s t o D i c k C a n n i n g s ( U . B . C . Cowan V e r t e b r a t e M u s e u m ) , J o h n R o d i l s k y (The B u r k e M u s e u m ) , D a v e N a g o r s e n ( R o y a l B . C . M u s e u m ) , K e v i n P u l l e n ( C o n n e r M u s e u m ) , E l l e n K r i t z m a n a n d D e n n i s P a u l s o n ( S l a t e r M u s e u m ) , N a t i o n a l Museums o f C a n a d a , a n d T e d U n d e r h i l l f o r p r o v i d i n g museum s p e c i m e n s a n d h i s t o r i c a l a c c o u n t s . F i n a l l y , I w o u l d l i k e t o e x p r e s s my a p p r e c i a t i o n t o my s u p e r v i s o r K i m C h e n g a n d t h e o t h e r members o f my s u p e r v i s o r y c o m m i t t e e , A l t o n H a r e s t a d , J a m i e S m i t h a n d Tom S u l l i v a n , f o r t h e i r g u i d a n c e a n d e n c o u r a g e m e n t . A n d o f c o u r s e , I mus t a c k n o w l e d g e t h o s e b r i g h t - e y e d b u s h y - t a i l e d c r e a t u r e s who e n t e r t a i n e d , i n t r i g u e d , a n d f a s c i n a t e d me, a n d w i t h o u t whose p r e s e n c e , t h i s p r o j e c t w o u l d n e v e r h a v e b e e n . F i n a n c i a l s u p p o r t f o r t h e p r o j e c t was p r o v i d e d b y T h e H a b i t a t C o n s e r v a t i o n F u n d , B . C . M i n i s t r y o f E n v i r o n m e n t , W o r l d W i l d l i f e F u n d , B . C . M i n i s t r y o f P a r k s , a n d B . C . C o n s e r v a t i o n F o u n d a t i o n . 1 CHAPTER 1: G E N E R A L INTRODUCTION CASCADE MANTLED GROUND S Q U I R R E L T h e C a s c a d e m a n t l e d g r o u n d s q u i r r e l , S p e r m o p h i l u s s a t u r a t u s , i s t h e l a r g e s t member o f t h e s u b g e n u s C a l l o s p e r m o p h i l u s , w h i c h a l s o i n c l u d e s S . l a t e r a l i s a n d S . m a d r e n s i s , a g r o u p o f s t o c k y g o l d e n b r o w n s q u i r r e l s w i t h two l a t e r a l w h i t e s t r i p e s b o r d e r e d b y n a r r o w d a r k s t r i p e s r u n n i n g f r o m s h o u l d e r s t o rump ( H o w e l l 1938 ; B a n f i e l d 1974; H a l l 1 9 8 1 ) . S . s a t u r a t u s i n h a b i t s t h e C a s c a d e M o u n t a i n s o f W a s h i n g t o n S t a t e a n d B r i t i s h C o l u m b i a , a r a n g e t h a t i s t h o u g h t t o be d i s j u n c t f r o m i t s m o r e w i d e s p r e a d s i b l i n g s p e c i e s , S . l a t e r a l i s ( F i g . 1 . 1 ) . A l t h o u g h u s u a l l y c o n s i d e r e d a s e p a r a t e s p e c i e s , t h e g e n e t i c c h a r a c t e r i s t i c s o f S . s a t u r a t u s h a v e n o t b e e n c o m p a r e d t o S . l a t e r a l i s p r i o r t o t h i s s t u d y . The n o r t h e r n l i m i t o f S . s a t u r a t u s i s a l s o unknown (Cowan a n d G u i g e t 1 9 6 0 ) . S . s a t u r a t u s i s known t o o c c u p y k r u m h o l t z , t a l u s s l o p e s , c l o s e d c o n i f e r o u s f o r e s t s , a n d o p e n meadows w h e r e u n d e r g r o u n d b u r r o w s c a n b e b u i l t ( R e i c h e l 1986; T r o m b u l a k 1 9 8 8 ) . D u r i n g s p r i n g , h e r b s a r e t h e m a i n f o o d w h i l e f u n g i a r e t h e m a i n f o o d i n f a l l ( K e n a g y 1 9 8 7 ) . F r o m m i d - A u g u s t t o l a t e S e p t e m b e r , h i b e r n a t i o n b e g i n s a n d l a s t s a b o u t 7 . 5 m o n t h s ( T r o m b u l a k 1 9 8 8 ) . F i r s t b r e e d i n g may n o t o c c u r u n t i l a f t e r a n i n d i v i d u a l h a s s u r v i v e d two w i n t e r s . A l l a d u l t m a l e s ( i . e . 2 y e a r s o r o l d e r ) 2 F i g . 1 . 1 . T h e r a n g e o f S p e r m o p h i l u s s a t u r a t u s a n d S . l a t e r a l i s . T h e s u b s p e c i e s o f S . l a t e r a l i s n e i g h b o u r i n g S . s a t u r a t u s (1) a r e S . l a t e r a l i s t e s c o r u m (2) , S . l a t e r a l i s c o n n e c t e n s (3) a n d S . l a t e r a l i s c h r y s o d e i r u s ( 4 ) . M o d i f i e d f r o m H a l l ( 1 9 8 1 ) . 3 d e v e l o p s c r o t a l t e s t e s a n d a l l a d u l t f e m a l e s , ( i . e . 2 y e a r s o r o l d e r ) b r e e d . W h i l e y e a r l i n g m a l e s may become s c r o t a l , f r o m 54% t o 100% o f y e a r l i n g f e m a l e s b r e e d . T h e a v e r a g e l i t t e r . s i z e i s f o u r a n d o n l y one l i t t e r i s r a i s e d e a c h y e a r ( K e n a g y a n d B a r n e s 1 9 8 8 ) . M i c h e n e r (1983) p l a c e d g o l d e n - m a n t l e d g r o u n d s q u i r r e l s i n t h e l e a s t o f f i v e s o c i a l c a t e g o r i e s f o r g r o u n d d w e l l i n g s q u i r r e l s . W i t h i n s u b g e n u s C a l l o s p e r m o p h i l u s , t h e o n l y c o n s i s t e n t c o h e s i v e a n d r e c o g n i t i v e b e h a v i o u r s o b s e r v e d a r e mock f i g h t i n g , s m e l l i n g , a n d b o d y c o n t a c t w h i c h o c c u r r e d among l i t t e r m a t e s o r b e t w e e n y o u n g a n d m o t h e r i n S . l a t e r a l i s ( F e r r o n 1 9 8 5 ) . S . s a t u r a t u s i s l i k e l y a s a s o c i a l . T h e m o s t s i m i l a r s p e c i e s t o S . s a t u r a t u s i n e c o l o g y i s t h e y e l l o w p i n e c h i p m u n k , E u t a m i a s a m o e n u s . b u t t h e i r d i e t s a r e d i f f e r e n t e n o u g h t h a t t h e y t o l e r a t e e a c h o t h e r w i t h i n 5 cm w h i l e f o r a g i n g ( B r o a d b o o k s 1 9 5 8 ) . P r e d a t o r s o f S . s a t u r a t u s i n c l u d e r a p t o r s , m u s t e l i d s , f e l i d s , a n d c a n i d s ( T r o m b u l a k 1 9 8 8 ) , b u t none o f t h e s e p r e d a t o r s a r e known t o d e p e n d p r i m a r i l y o n S . s a t u r a t u s f o r f o o d . I n summary , p r e v i o u s l y a v a i l a b l e i n f o r m a t i o n s u g g e s t t h a t S . s a t u r a t u s c a n l i v e i n a v a r i e t y o f h a b i t a t s , f e e d s o n a v a r i e t y o f p l a n t s p e c i e s , h a s no m a j o r c o m p e t i t o r s , a n d i s n o t t h e m a j o r p r e y i t e m f o r a n y o f i t s p r e d a t o r s . H o w e v e r , m o s t o f t h i s i n f o r m a t i o n i s g e n e r a l a n d more d e t a i l o n i t s l i f e h i s t o r y i s n e e d e d t o a s s e s s i t s p o p u l a t i o n v u l n e r a b i l i t y . 4 O B J E C T I V E S C o n s e r v a t i o n i m p l i e s t h e m a i n t e n a n c e o f v i a b l e p o p u l a t i o n s o f a s p e c i e s i n t h e i r n a t u r a l s t a t e a n d t h e m a i n t e n a n c e o f gene p o o l s f o r t h e f u t u r e ( P a s s m o r e 1974; F r a n k e l a n d S o u l e 1 9 8 1 ; F r a n k e l 1 9 8 4 ) . S . s a t u r a t u s i s a n o n - g a m e s p e c i e s , s o t h e p r i m a r y j u s t i c a t i o n f o r i t s c o n s e r v a t i o n i s e t h i c a l r a t h e r t h a n e c o n o m i c a l . On e t h i c a l g r o u n d s , a l l n a t u r a l o b j e c t s h a v e i n t r i n s i c v a l u e ( C a l l i c o t t 1986; V a n D e V e e r a n d P i e r c e 1986 ; N o r t o n 1987) . T h e g e n e t i c d i v e r s i t y r e p r e s e n t e d b y s p e c i e s a n d t h e h a b i t a t s t h e y r e q u i r e a r e among t h e c r i t e r i a u s e d t o d e v e l o p management s c h e m e s f o r t h e c o n s e r v a t i o n o f s p e c i e s ( G i l p i n a n d S o u l e 1986 ; COSEWIC u n p u b l . ) . B i o d i v e r s i t y i n c l u d e s t h e number o f d i f f e r e n t s p e c i e s a n d t h e g e n e t i c v a r i a t i o n w i t h i n t h e i r p o p u l a t i o n s . When d e v e l o p i n g a c o n s e r v a t i o n p l a n f o r a s i n g l e s p e c i e s , i n f o r m a t i o n o n i t s g e n e t i c c o m p o s i t i o n i s n e e d e d , a l o n g w i t h i n f o r m a t i o n o n i t s d i s t r i b u t i o n a n d o n t h e m a j o r c o m p o n e n t s i n i t s h a b i t a t . Among t h e o b j e c t i v e s o f t h e M i n i s t r y o f E n v i r o n m e n t W i l d l i f e B r a n c h i n B r i t i s h C o l u m b i a , i s t h e m a i n t a i n e n a n c e o f r e p r e s e n t a t i v e s p e c i e s d i v e r s i t y i n e a c h o f t h e p r o v i n c e ' s b i o p h y s i c a l z o n e s a n d t o e n s u r e w i l d l i f e a v a i l a b i l i t y t o s o c i e t y . I n 1979 , t h e G o v e r n m e n t o f B r i t i s h C o l u m b i a f o r m e d a p r e l i m i n a r y p l a n t o d e s i g n a t e t h r e a t e n e d a n d e n d a n g e r e d s p e c i e s i n t h e p r o v i n c e (Munro a n d Low 1 9 7 9 ) . S i n c e t h e n , r e g u l a t i o n s f o r n o n - g a m e mammals w e r e i m p r o v e d b y a n O r d e r - i n - C o u n c i l i n 5 M a y , 1 9 9 0 , w h e r e b y a l l n a t i v e , n o n - g a m e mammals a c q u i r e d f u l l p r o t e c t i o n o n c r o w n l a n d u n d e r t h e o f f i c i a l d e s i g n a t i o n " w i l d l i f e " . T h e l a c k o f k n o w l e d g e o n n o n - g a m e s p e c i e s l e d t o a l i s t i n g o f s p e c i e s m o s t l i k e l y t o be t h r e a t e n e d o r e n d a n g e r e d . Among t h e h i g h e s t p r i o r i t y v e r t e b r a t e s (Red L i s t ) a w a i t i n g d e s i g n a t i o n , i s t h e C a s c a d e m a n t l e d g r o u n d s q u i r r e l . T h e p u r p o s e o f t h i s s t u d y i s t o p r o v i d e i n f o r m a t i o n on t h r e e m a j o r g a p s i n k n o w l e d g e n e c e s s a r y f o r t h e c o n s e r v a t i o n o f S . s a t u r a t u s i n B r i t i s h C o l u m b i a : 1. T h e t a x o n o m y o f t h e S . s a t u r a t u s - S . l a t e r a l i s s p e c i e s c o m p l e x . 2 . T h e r a n g e a n d d i s t r i b u t i o n o f S . s a t u r a t u s i n B r i t i s h C o l u m b i a . 3 . T h e m a j o r h a b i t a t c o m p o n e n t s u s e d b y S . s a t u r a t u s . By i n t e g r a t i n g new i n f o r m a t i o n o n t h e s e t h r e e t o p i c s w i t h e x i s t i n g i n f o r m a t i o n on S . s a t u r a t u s a n d S . l a t e r a l i s , I p r o p o s e r e c o m m e n d a t i o n s p e r t a i n i n g t o t h e c o n s e r v a t i o n o f S . s a t u r a t u s . p a r t i c u l a r l y i n B r i t i s h C o l u m b i a . 6 CHAPTER 2: TAXONOMY OF THE SPERMOPHILUS SATURATUS - S . L A T E R A L I S S P E C I E S COMPLEX INTRODUCTION M a y r (1970) d e f i n e s b i o l o g i c a l s p e c i e s a s " g r o u p s o f i n t e r b r e e d i n g n a t u r a l p o p u l a t i o n s t h a t a r e r e p r o d u c t i v e l y i s o l a t e d f r o m o t h e r s u c h g r o u p s " , b u t a p p l y i n g t h i s d e f i n i t i o n t o d i s t i n g u i s h c l o s e l y - r e l a t e d g r o u p s i s o f t e n d i f f i c u l t . F o r e x a m p l e , g e o g r a p h i c a l l y i s o l a t e d g r o u p s o f i n d i v i d u a l s a r e n a t u r a l l y r e p r o d u c t i v e l y i s o l a t e d , b u t may s t i l l b e c a p a b l e o f p r o d u c i n g v i a b l e a n d f e r t i l e y o u n g when g i v e n t h e o p p o r t u n i t y t o i n t e r b r e e d . N e v e r t h e l e s s , t h e e s s e n c e o f t h e s p e c i e s c o n c e p t i s g e n e t i c c o h e s i v e n e s s , a n d c o n s i s t e n t d i f f e r e n c e s b e t w e e n g r o u p s s e p a r a t e t h e m i n t o s p e c i e s . T h u s , t h e number o f s p e c i e s p a r t i a l l y r e f l e c t s g e n e t i c d i v e r s i t y . G e n e t i c d i v e r s i t y may e n s u r e f u t u r e s p e c i a t i o n ( F r a n k e l 1 9 8 4 ) , a n d m i n i m i z e c h a n c e e x t i n c t i o n s t h a t a r e d u e t o l a c k o f a d a p t a b i l i t y t o c h a n g i n g e n v i r o n m e n t a l c o n d i t i o n s ( S o u l e 1 9 8 0 ; L a n d e 1 9 8 8 ) . B e c a u s e r e p r o d u c t i v e i s o l a t i o n i s d e p e n d e n t o n t h e n a t u r e o f t h e d i f f e r e n c e s b e t w e e n p o p u l a t i o n s a n d n o t j u s t t h e g e n e t i c d i f f e r e n c e s a c c u m u l a t e d ( B i c k m a n 1 9 8 3 ) , q u a n t i f y i n g o v e r a l l d i v e r s i t y b y m e a s u r i n g i n t e r - a n d i n t r a - s p e c i f i c v a r i a t i o n i s a more p r e c i s e m e a s u r e m e n t o f g e n e t i c d i v e r s i t y t h a n i s number o f s p e c i e s . 7 T h e p h y l o g e n y o f N e a r c t i c S c i u r i d a e , i n c l u d i n g C a l l o s p e r m o p h i l u s . h a s b e e n d e s c r i b e d b y H a f n e r ( 1 9 8 4 ) , b u t t h e g e n e t i c v a r i a t i o n w i t h i n a n d b e t w e e n S . l a t e r a l i s a n d S . s a t u r a t u s h a d n o t b e e n q u a n t i f i e d . B e f o r e t h e a d v e n t o f t e c h n i q u e s t o m e a s u r e g e n e t i c v a r i a t i o n , S . s a t u r a t u s was g i v e n s p e c i e s s t a t u s b a s e d o n i t s l a r g e r b o d y s i z e a n d l e s s d i s t i n c t l a t e r a l s t r i p e s c o m p a r e d t o S . s a t u r a t u s ( H o w e l l 1 9 3 8 ) . H e r e , I r e a s s e s s t h i s s t a t u s b y m e a s u r i n g i n t e r a n d i n t r a - s p e c i f i c v a r i a t i o n u s i n g e l e c t r o p h o r e s i s a n d m o r p h o l o g i c a l m e a s u r e m e n t s . I c h o s e t o u s e e l e c t r o p h o r e t i c a n a l y s i s t o r e - e x a m i n e t h e t a x o n o m i c s t a t u s o f t h e s e s i b l i n g s p e c i e s b e c a u s e i t p r o v i d e s more d i r e c t g e n e t i c i n f o r m a t i o n t h a n m o r p h o l o g i c a l d i m e n s i o n s ( G e i s t 1 9 8 9 ) , w h i c h s h o u l d o n l y be u s e d a s s u p p l e m e n t a l e v i d e n c e . A c o m p a r i s o n o f a l l e l e f r e q u e n c i e s b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s w i l l h e l p t o d e c i d e w h e t h e r S . s a t u r a t u s s h o u l d be t r e a t e d a s a s p e c i e s s e p a r a t e f r o m S . l a t e r a l i s . T h e i n t e r p r e t a t i o n o f a l l e l e f r e q u e n c i e s i n r e l a t i o n t o p o p u l a t i o n m o d e l s i s b e s t d o n e b y s t a r t i n g f r o m t h e s m a l l e s t t o t h e l a r g e s t p o p u l a t i o n u n i t ( R i c h a r d s o n e t a l . 1 9 8 6 ) . I e x a m i n e t h e v a r i a t i o n w i t h i n c a p t u r e l o c a t i o n s , t h e n t h e i n t r a - s p e c i f i c v a r i a t i o n among l o c a t i o n s , a n d f i n a l l y , s u b s p e c i f i c o r i n t e r -s p e c i f i c d i f f e r e n c e s . 8 METHODS E l e c t r o p h o r e t i c A n a l y s i s S p e r m o p h i l u s s a t u r a t u s a n d S p e r m o p h i l u s l a t e r a l i s ( s u b s p e c i e s t e s c o r u m ) w e r e c a p t u r e d w i t h Tomahawk #2 01 l i v e t r a p s a t f o u r a n d s i x l o c a t i o n s r e s p e c t i v e l y ( F i g . 2 . 1 ) . T h e number o f i n d i v i d u a l s s a m p l e d f r o m e a c h l o c a t i o n r a n g e d f r o m 3 t o 2 3 . A p p r o x i m a t e l y 1 .0 m l o f b l o o d was c o l l e c t e d f r o m e a c h a n i m a l b y r u p t u r i n g t h e o r b i t a l p l e x u s w i t h h e p a r i n i z e d c a p i l l a r y t u b e s w h i l e t h e a n i m a l was a n e s t h e t i z e d a f t e r i n h a l i n g m e t h o x y f l o u r i n e ( P o o l 1 9 8 7 ) . T h e b l o o d was i m m e d i a t e l y c e n t r i f u g e d t o s e p a r a t e r e d b l o o d c e l l s f r o m s e r u m . E a c h f r a c t i o n was p l a c e d i n a v i a l , f r o z e n i n l i q u i d n i t r o g e n o r d r y i c e a n d s t o r e d i n a - 7 0 °C f r e e z e r f o r up t o s i x m o n t h s u n t i l e l e c t r o p h o r e t i c a n a l y s i s . T h e g r o u n d s q u i r r e l s w e r e a l l o w e d t o r e c o v e r f o r 20 m i n u t e s i n a c o v e r e d c o n t a i n e r a n d t h e n r e l e a s e d a t t h e i r c a p t u r e s i t e . 1 S t a n d a r d h o r i z o n t a l s t a r c h g e l e l e c t r o p h o r e s i s t e c h n i q u e s (May e t a l . 1979) w e r e a p p l i e d t o t h e b l o o d s a m p l e s b y C o r n e l l U n i v e r s i t y ' s L a b o r a t o r y o f E c o l o g i c a l a n d E v o l u t i o n a r y G e n e t i c s ( C L E E G ) u n d e r t h e s u p e r v i s i o n o f B . P . M a y . A c t i v i t y was r e c o r d e d f o r 24 p r e s u m p t i v e g e n e l o c i : g l u c o s e p h o s p h a t e i s o m e r a s e ( G P I ) , n u c l e o s i d e p h o s p h o r y l a s e ( N P ) , p h o s p h o g l u c o n a t e d e h y d r o g e n a s e ( P G D ) , p e p t i d a s e l e u c y l - g l y c y l - g l y c i n e ( P E P - L G G ) , e s t e r a s e ( E S T - 1 ) , p e p t i d a s e w i t h g l y c y l - l e u c i n e - 1 ( P E P - G L - 1 ) , p e p t i d a s e w i t h g l y c y l l e u c i n e - 2 ( P E P - G L - 2 ) , g e n e r a l p r o t e i n - 2 (PRO - 2 ) , p e p t i d a s e l e u c y l a l a n i n e ( P E P - L A ) , s u p e r o x i d e F i g . 2 . 1 . Map o f s o u t h w e s t e r n C a n a d a a n d n o r t h w e s t e r n U n i t e d S t a t e s s h o w i n g l o c a t i o n s o f b l o o d s a m p l i n g . F o r S . s a t u r a t u s : I r o n M o u n t a i n 5 0 ° 0 3 ' N 1 2 0 ° 4 6 , W ( 1 ) , B r o o k m e r e Dump 4 9 ° 4 9 ' N 1 2 0 ° 5 2 ' W ( 2 ) , T h y n n e M o u n t a i n 4 9 ° 4 9 ' N 1 2 0 ° 5 2 ' W ( 3 ) , a n d L a k e W e n a t c h e e 4 7 ° 5 4 ' N 1 2 0 ° 4 5 ' W ( 4 ) . F o r S . l a t e r a l i s : M o u n t B a l d y 4 9 ° 1 0 I N 1 1 9 ° 1 5 , W ( A ) , R o s s l a n d 4 9 ° 0 5 ' N 1 1 7 ° 4 9 ' W (B) , S p h i n x M o u n t a i n 4 9 ° 3 7 ' N 1 1 6 ° 3 9 ' W (C) , w e s t o f S p r a y L a k e s 5 0 ° 5 0 ' N 1 1 5 ° 2 0 ' W (D) , L i m e s t o n e M o u n t a i n 5 0 ° 5 1 ' N 1 1 5 ° 1 0 ' W (E) , a n d P e t e r L o u g h e e d P r o v i n c i a l P a r k 5 0 ° 5 8 ' N 1 1 5 ° 2 0 ' W (F) . 10 d i s m u t a s e ( S O D ) , a d e n o s i n e d e a m i n a s e ( A D A ) , m a n n o s e p h o s p h a t e i s o m e r a s e ( M P I ) , t r i o s e p h o s p h a t e i s o m e r a s e ( T P I ) , m a l a t e d e h y d r o g e n a s e (MDH), l a c t a t e d e h y d r o g e n a s e ( L D H ) , g l u c o s e - 6 -p h o s p h a t e d e h y d r o g e n a s e ( G 6 P D H ) , m e t h y l u m b e l l i f e r y l p h o s p h a t e ( M U P ) , a l p h a - m a n n o s i d a s e ( a - M A N ) , c a t a l a s e ( C A T ) , d i a p h o r a s e ( D I A ) , p e p t i d a s e p h e n y l - a l a n y l - p r o l i n e ( P E P - P A P ) , l e u c i n e a m i n o p e p t i d a s e ( L A P ) , p h o s p h o g l u c o m u t a s e ( P G M ) , g e n e r a l p r o t e i n -1 (PRO-1) . T h e f r e q u e n c y o f a l l e l e s a t t h e l o c i f o r e a c h l o c a t i o n w e r e q u a n t i f i e d , t e s t e d f o r H a r d y - W e i n b e r g e q u i l i b r i u m a n d c o m p a r e d u s i n g t h e g e n e t i c m e a s u r e m e n t s o f R o g e r s ' (1972) s i m i l a r i t y a n d N e i ' s (1978) u n b i a s e d d i s t a n c e a s c a l c u l a t e d b y B I O S Y S - 1 ( S w o f f o r d a n d S e l a n d e r 1 9 8 9 ) . G u i d e l i n e s f o r u s i n g R o g e r s ' s i m i l a r i t y a n d N e i ' s d i s t a n c e t o i n t e r p r e t t a x a n o m i c s t a t u s c a n b e f o u n d i n A v i s e (1976) a n d N e i ( 1 9 7 6 ) . M o r p h o l o g i c a l A n a l y s i s B e f o r e c o l l e c t i n g v a l u e s f o r s t a t i s t i c a l a n a l y s e s , a s e t o f 25 s k u l l s w e r e m e a s u r e d t w i c e b y two o b s e r v e r s ( m y s e l f a n d a n a s s i s t a n t ) t o a s s u r e t h e r e p e a t a b i l i t y o f t h e m e a s u r e m e n t s a n d m i n i m i z e o b s e r v e r b i a s . T h e n e a c h o b s e r v e r was a s s i g n e d t o m e a s u r e h a l f o f t h e c h a r a c t e r i s t i c s . T e n c r a n i a l c h a r a c t e r i s t i c s ( F i g . 2 . 2 ) , m e a s u r e d w i t h v e r n i e r c a l i p e r s t o t h e n e a r e s t 0 . 1 mm, w e r e c h o s e n b e c a u s e t h e y c o u l d b e c o m p a r e d w i t h o t h e r d a t a a n d a r e c o n s i d e r e d s t a n d a r d c r a n i a l m e a s u r e m e n t s ( H o w e l l 1 9 3 8 ; H a l l 1 9 8 1 ) . T h e t a x o n o m i c g r o u p s c o m p a r e d w e r e S . s a t u r a t u s a n d i t s n e i g h b o u r i n g s i b l i n g s p e c i e s ( F i g . 1 . 1 ) , a m o u n t i n g t o t h r e e s u b s p e c i e s , S . l a t e r a l i s t e s c o r u m , S . 11 F i g . 2.2. L a t e r a l , v e n t r a l a n d d o r s a l v i e w s o f S . s a t u r a t u s s k u l l s h o w i n g t e n d i m e n s i o n s t h a t w e r e m e a s u r e d : d i a s t e m a l e n g t h (1) , o c c i p i t a l n a s a l l e n g t h ( 2 ) , p a l a t a l l e n g t h ( 3 ) , z y g o m a t i c b r e a d t h ( 4 ) , m a s t o i d a l b r e a d t h ( 5 ) , i n t e r o r b i t a l b r e a d t h ( 6 ) , n a s a l l e n g t h ( 7 ) , m a x i l l a r y t o o t h row ( 8 ) , c r a n i a l b r e a d t h ( 9 ) , a n d p o s t o r b i t a l c o n s t r i c t i o n ( 1 0 ) . l a t e r a l i s c h r y s o d e i r u s . a n d S . l a t e r a l i s c o n n e c t e n s . Where s k u l l s w e r e i n c o m p l e t e o r damaged a n d c e r t a i n d i m e n s i o n s u n m e a s u r a b l e , t h e s k u l l was e x c l u d e d f o r c o m p a r i s o n s o f t h a t c h a r a c t e r i s t i c . T h e c a t a l o g u e n u m b e r , c o l l e c t i o n l o c a t i o n a n d museum f o r e a c h s k u l l a r e l i s t e d i n A p p e n d i x 1. I n o r d e r t o d i s t i n g u i s h s h a p e a s w e l l a s s i z e d i f f e r e n c e s , a n a n a l y s i s o f c o v a r i a n c e was u s e d w i t h z y g o m a t i c b r e a d t h a s t h e c o v a r i a t e . R E S U L T S E l e c t r o p h o r e t i c A n a l y s e s E i g h t o f 24 l o c i e x a m i n e d w e r e p o l y m o r p h i c ( T a b l e 2 . 1 ) . B l o o d s a m p l e s t a k e n f r o m S . s a t u r a t u s i n d i c a t e d t h a t t h e s e l o c i w e r e f i x e d f o r a p a r t i c u l a r a l l e l e , w h i l e t h o s e f r o m S . l a t e r a l i s showed s e g r e g a t i o n a n d v a r i e d b e t w e e n p o p u l a t i o n s , a v e r a g i n g 0 . 9 3 5 ( r a n g e : 0 . 8 8 0 - 0 . 9 8 5 ) a c c o r d i n g t o R o g e r s ' s i m i l a r i t y (1972) a n d 0 . 0 2 8 ( r a n g e : 0 . 0 0 0 - 0 . 0 8 3 ) a c c o r d i n g t o N e i ' s d i s t a n c e ( 1 9 7 8 ) . F r e q u e n c i e s f o r t h e p o l y m o r p h i c l o c i a r e p r e s e n t e d i n T a b l e 2 . 1 . Two a l l e l e s (PGD, P R O - 2 ) w e r e p r e s e n t i n S . s a t u r a t u s a n d a b s e n t i n S . l a t e r a l i s ( T a b l e 2 . 1 ) . H o w e v e r , t h e d i f f e r e n c e i n a l l e l e f r e q u e n c i e s b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s c h a n g e d f r o m one S . l a t e r a l i s l o c a t i o n t o a n o t h e r ( T a b l e 2 . 2 ) . T h e r e was g r e a t e r d i f f e r e n c e b e t w e e n some l o c a t i o n s o f S . l a t e r a l i s t h a n b e t w e e n S . l a t e r a l i s a n d S . s a t u r a t u s a c c o r d i n g t o R o g e r s ' s i m i l a r i t y a n d N e i ' s 13 T a b l e 2 . 1 . A l l e l i c v a r i a t i o n a t e i g h t p o l y m o r p h i c l o c i f o r S. s a t u r a t u s a n d S . l a t e r a l i s a t d i f f e r e n t l o c a t i o n s . S e e F i g . 2 . 1 . f o r l o c a t i o n c o d e s . Locations of S. saturatus Locations of S. lateralis Locus GPI Allele A B 1,2,3,4 1.000 0.000 0.556 0.444 B 0.583 0.417 0.833 0.167 D 1.000 0.000 1.000 0.000 1.000 0.000 NP A B 1.000 0.000 0.833 0.167 1.000 0.000 1.000 0.000 0.857 0.143 0.875 0.125 0.885 0.115 PGD A B 0.000 1.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 PEP-LGG A B 1.000 0.000 1.000 0.000 0.833 0.167 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 EST-1 A B 1.000 0.000 0.000 1.000 0.000 .1.000 0.333 0.667 0.500 0.500 1.000 0.000 0.769 0.231 PEP-GL-1 A B 1.000 0.000 0.889 0.111 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 PEP-GL-2 A B 1.000 0.000 0.500 0.500 0.300 0.700 0.500 0.500 1.000 0.000 1.000 0.000 1.000 0.000 PRO-2 A B C 0.000 0.000 1.000 0.944 0.056 0.000 0.750 0.250 0.000 0.500 0.500 0.000 0.571 0.429 0.000 0.286 0.714 0.000 0.500 0.500 0.000 No. of samples •48 13 * 48 is the sum of samples from: 1. Iron Mtn (7); 2. Brookmere Dump (23); 3. Thynne Mtn (11); 4. Wenatchee Lake (7). 14 T a b l e 2 . 2 . M e a s u r e s o f g e n e t i c r e l a t e d n e s s f o r S . s a t u r a t u s (Ss ) a n d S . l a t e r a l i s ( S I ) . R o g e r s ' (1972) s i m i l a r i t y a p p e a r s a b o v e t h e d i a g o n a l a n d N e i ' s (1978) u n b i a s e d d i s t a n c e a p p e a r s b e l o w t h e d i a g o n a l . S e e F i g . 2 . 1 . f o r l o c a t i o n c o d e s . *S. saturatus 3. lateralis Location 1,2,3,4 A B C D E F 1,2,3,4 (Ss) - 0.825 0.826 0.867 0.895 0.916 0.908 A (SI) 0.158 - 0.964 0.944 0.919 0.885 0.903 B(SI) 0.160 0.001 - 0.950 0.912 0.880 0.899 C (SI) 0.105 0.009 0.003 - 0.956 0.930 0.949 D (SI) 0.088 0.034 0.039 0.005 0.967 0.985 E (SI) 0.078 0.083 0.083 0.026 0.011 - 0.985 F (SI) 0.079 0.054 0.057 0.013 0.000 0.002 -* Ail S. saturatus were lumped because they were all the same. u n b i a s e d d i s t a n c e ( T a b l e 2 . 2 ) . On a v e r a g e , h o w e v e r , t h e d i f f e r e n c e b e t w e e n t h e two s p e c i e s was 0 . 8 7 3 f o r R o g e r s ' s i m i l a r i t y a n d 0 . 1 1 1 f o r N e i ' s u n b i a s e d d i s t a n c e . D e v i a t i o n f r o m H a r d y - W e i n b e r g e q u i l i b r i u m was n o t d e t e c t e d (P > 0 . 0 5 ) f o r a n y l o c a t i o n s o f S . l a t e r a l i s . I t was n o t p o s s i b l e t o t e s t f o r H a r d y - W e i n b e r g e q u i l i b r i u m i n t h e S . s a t u r a t u s s a m p l e d b e c a u s e t h e y h a d no a p p a r e n t g e n e t i c v a r i a t i o n . M o r p h o l o g i c a l A n a l y s e s S i n c e m a l e s a n d f e m a l e s w e r e s i g n i f i c a n t l y (P < 0 . 0 5 ) d i m o r p h i c f o r s i x m e a s u r e m e n t s ( p a l a t a l l e n g t h , p o s t o r b i t a l c o n s t r i c t i o n , o c c i p i t o n a s a l l e n g t h , m a s t o i d a l b r e a d t h , n a s a l l e n g t h , c r a n i a l b r e a d t h ) , s e x e s w e r e a n a l y z e d s e p a r a t e l y ( T a b l e 2 . 3 a n d T a b l e 2 . 4 ) . T h e z y g o m a t i c b r e a d t h was a s i g n i f i c a n t c o v a r i a t e (P < 0 . 0 5 ) f o r a l l c r a n i a l m e a s u r e m e n t s e x c e p t f o r m a x i l l a r y t o o t h row l e n g t h f o r b o t h s e x e s . I n a d d i t i o n , p o s t o r b i t a l c o n s t r i c t i o n was n o t a s i g n i f i c a n t c o v a r i a t e f o r f e m a l e s ( T a b l e 2 . 4 ) . A l t h o u g h l a r g e r s k u l l s a r e g e n e r a l l y l a r g e r f o r a l l m e a s u r e m e n t s , s i g n i f i c a n t d i f f e r e n c e s f r o m t h e a n a l y s i s o f c o v a r i a n c e i n d i c a t e s t h a t t h e s k u l l p r o p o r t i o n s d i f f e r b e t w e e n S . s a t u r a t u s a n d t h e s u b s p e c i e s o f S . l a t e r a l i s ( T a b l e 2 . 5 ) . S . s a t u r a t u s a r e l a r g e r t h a n S . l a t e r a l i s f o r a l l c r a n i a l m e a s u r e m e n t s a n d w i t h i n S . l a t e r a l i s , t h e s u b s p e c i e s S . l a t e r a l i s t e s c o r u m i s c o n s i s t e n t l y t h e l a r g e s t ( T a b l e 2 . 3 ) . 16 T a b l e 2 . 3 . T h e mean (x) , s t a n d a r d d e v i a t i o n (sd) , a n d s a m p l e s i z e (N) f o r t h e t e n c r a n i a l m e a s u r e m e n t s o f S . s a t u r a t u s ( S s ) , S . l a t e r a l i s t e s c o r u m ( S i t ) , S . l a t e r a l i s c h r y s o d e i r u s ( S l c h ) , a n d S . l a t e r a l i s c o n n e c t e n s ( S l c o ) . Character Male Female Diastema length Occipitonasal length Palatal length Zygomatic breadth Mastoidal breadth Ss Sit Slch Slco Ss Sit Slch Slco X 11.62 10.98 10.3 10.55 11.32 10.88 10.21 10.4 sd 0.63 0.55 0.49 0.57 0.60 0.57 0.52 0.51 N 43 27 13 13 37 15 38 11 X 45.82 44.24 42.38 42.48 45.09 43.31 41.69 41.84 sd 1.54 1.09 1.11 1.21 1.22 1.52 1.13 1.03 N 42 26 13 13 36 13 37 10 X 21.09 20.22 19.58 19.40 20.77 20.08 19.12 19.16 sd 0.65 0.73 0.62 0.59 0.66 0.84 0.66 0.57 N 43 27 13 13 37 15 38 11 X 28.08 27.33 25.98 26.11 27.54 26.94 25.96 25.76 sd 1.36 0.8 1.11 1.2 1.11 0.94 0.97 1.02 N 43 27 13 13 36 14 38 11 X 19.85 19.68 19.08 19.14 19.50 19.25 18.73 18.88 sd 0.56 0.59 0.55 0.59 0.54 0.67 0.59 0.44 N 43 26 13 12 37 13 37 11 Interorbital constriction Nasal length Cranial breadth Postorbital constriction x sd N X 11.17 10.59 9.92 9.65 11 10.32 9.77 9.52 sd 0.54 0.54 0.34 0.59 0.72 0.49 0.45 0.73 N 43 27 13 37 14 38 11 X 14.98 14.24 13.55 12.99 14.83 13.82 13.28 12.62 sd 0.67 0.62 0.57 0.66 0.56 0.61 0.64 0.63 N 42 27 13 13 36 13 38 10 w X 9.18 8.76 8.62 8.52 9.29 8.65 8.53 8.45 sd 0.25 0.43 0.31 0.34 0.35 0.24 0.25 0.35 N 43 27 13 13 37 15 38 11 X 19.79 19.48 19.34 19.34 19.45 19.31 19.01 18.90 sd 0.41 0.64 0.54 0.45 0.52 0.53 0.57 0.40 N 43 27 13 11 37 15 37 11 13.3 0.42 43 12.78 0.56 27 12.96 0.47 13 12.62 0.75 13 12.97 0.64 37 12.65 0.34 15 12.45 0.45 38 12.59 0.59 11 T a b l e 2 . 4 . D i f f e r e n c e s b e t w e e n s e x a n d s p e c i e s ( w i t h i n sex) f o r t e n s k u l l m e a s u r e m e n t s u s i n g A n a l y s i s o f C o v a r i a n c e w i t h z y g o m a t i c b r e a d t h a s t h e c o v a r i a t e . N o t e t h a t t h e s t a t i s t i c s f o r z y g o m a t i c b r e a d t h a r e f r o m A n a l y s i s o f V a r i a n c e . S i g n i f i c a n c e d i f f e r e n c e s a r e i n d i c a t e d b y "s" (P < 0 . 0 5 ) , a n d n o n s i g n i f i c a n t d i f f e r e n c e s a r e i n d i c a t e d by " n s " . Sex Female Male Species Covariate Species Covariate F F F F F Diastema length 1.23 ns 8.28 s 98.84 s 6.42 s 102.06 s Occipital nasal length 13.39 s 28.58 s 128.11 s 14.55 s 180.69 s Palatal length 5.68 s 16.27 s 88.80 s 12.04 s 79.82 s Zygomatic breadth 3.38 ns 17.93 s - - 16.26 s - -Mastoidal breadth 11.63 s 1.04 ns 125.81 s 0.35 ns 129.34 s Interorbital constriction 2.58 ns 12.81 s 17.65 s 14.07 s 38.48 s Nasal length 6.03 s 26.26 s 24.96 s 17.27 s 45.35 s Maxillary tooth row 0.39 ns 30.51 s 1.02 ns 12.76 s 0.64 ns Cranial breadth 11.40 s 0.13 ns 44.89 s 0.67 ns 35.19 s Postorbital constriction 12.32 s 5.72 s 1.40 ns 12.30 s 8.52 s 18 T a b l e 2 . 5 . D i f f e r e n c e s b e t w e e n S . s a t u r a t u s ( S s ) , S . l a t e r a l i s  t e s c o r u m ( S i t ) , S . l a t e r a l i s c h r v s o d e i r u s ( S l c h ) , a n d S . l a t e r a l i s c o n n e c t e n s ( S l c o ) f o r e a c h c r a n i a l c h a r a c t e r i s t i c a c c o r d i n g t o L e a s t S i g n i f i c a n t D i f f e r e n c e s m u l t i p l e r a n g e t e s t . H o r i z o n t a l l i n e s i n d i c a t e n o n s i g n i f i c a n t d i f f e r e n c e s (P > 0 . 0 5 ) . S e x e s w e r e a n a l y z e d s e p a r a t e l y . Character Male Female Ss Sit Slch Slco Ss Slco Sit Slco Diastema length _ m ' Ss Sit Slco Slch Occipitonasal length Palatal length Zygomatic breadth Mastoidal breadth Ss Sit Slch ' Slco Interorbital constriction Nasal length Maxillary tooth row Cranial breadth' Postorbital constriction 19 D I S C U S S I O N T h e d a t a p r e s e n t e d s u p p o r t t h e t a x o n o m i c s t a t u s o f S . s a t u r a t u s a s a s p e c i e s . T h e two a l l e l e s (PGD, P R O -2 ) t h a t a r e p r e s e n t i n t h e S . s a t u r a t u s s a m p l e d a n d a b s e n t i n t h e S . l a t e r a l i s s a m p l e d , s u g g e s t s t h a t S . s a t u r a t u s i s g e n e t i c a l l y d i s t i n c t . R o g e r s ' s i m i l a r i t y a n d N e i ' s d i s t a n c e d e t e c t e d a l a r g e e n o u g h d i f f e r e n c e t o m e r i t g i v i n g S . s a t u r a t u s s p e c i e s s t a t u s , e s p e c i a l l y when c o m p a r i n g t o S . l a t e r a l i s f r o m t h e l o c a t i o n s c l o s e s t t o S . s a t u r a t u s . A l t h o u g h S . l a t e r a l i s f r o m t h e e a s t e r n m o s t l o c a t i o n s ( i . e . f u r t h e s t f r o m S . s a t u r a t u s ) w e r e g e n e t i c a l l y l e s s d i s t i n c t f r o m S . s a t u r a t u s t h a n t h e c l o s e s t S . l a t e r a l i s s a m p l e d , t h e g e o g r a p h i c d i s t a n c e b e t w e e n S . s a t u r a t u s a n d t h e e a s t e r n m o s t S . l a t e r a l i s was t o o f a r f o r them t o i n t e r b r e e d . T h e g e n e t i c c o e f f i c i e n t s s h o u l d b e i n t e r p r e t e d w i t h c a u t i o n b e c a u s e o f t h e w e i g h t i n g o f i n t e r - a n d i n t r a - s p e c i f i c v a r i a t i o n a n d t h e c o e f f i c i e n t s ' u n d e r l y i n g a s s u m p t i o n s o f r a n d o m m a t i n g a n d i n f i n i t e p o p u l a t i o n s i z e , n e i t h e r o f w h i c h h a v e b e e n v a l i d a t e d . W h i l e t h e number o f b l o o d s a m p l e s c o l l e c t e d was c o n s i d e r e d a d e q u a t e f o r t h e o v e r a l l c o m p a r i s o n b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s . t h e s a m p l e s i z e f o r i n d i v i d u a l l o c a t i o n s w i t h i n s p e c i e s was s m a l l . F u r t h e r m o r e , t h e s e l o c a t i o n s d i d n o t r e p r e s e n t t h e f u l l g e o g r a p h i c r a n g e o f t h e two s p e c i e s . N e v e r t h e l e s s , d i f f e r e n c e s i n a l l e l i c f r e q u e n c i e s f o u n d b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s s u p p o r t t h e t r e a t m e n t o f S . s a t u r a t u s a s a s p e c i e s s e p a r a t e f r o m S . l a t e r a l i s . T h e p a t t e r n o f g e n e t i c c h a n g e a c r o s s t h e r a n g e o f S . l a t e r a l i s f i t s t h e i s o l a t i o n - b y - d i s t a n c e m o d e l i n w h i c h t h e g e n o t y p e s o f a d j a c e n t i n d i v i d u a l s a r e more c l o s e l y r e l a t e d t h a n i n d i v i d u a l s f u r t h e r away ( R i c h a r d s o n e t a l . 1 9 8 6 ) . M o r p h o l o g i c a l c h a r a c t e r i s t i c s , i n c l u d i n g p e l a g e c o l o u r a n d p a t t e r n c a n d i s t i n g u i s h S . s a t u r a t u s f r o m S . l a t e r a l i s . W a r l s b e r g (1990) f o u n 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 p e l a g e c h a r a c t e r i s t i c s b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s f r o m A r i z o n a b y q u a n t i f y i n g t h e s o l a r h e a t g a i n o f t h e i r c o a t l a y e r s a n d s k i n . A l t h o u g h S . s a t u r a t u s g e n e r a l l y h a s l e s s d e f i n e d l a t e r a l s t r i p e s t h a n S . l a t e r a l i s ( H a l l 1 9 8 1 ) , d u r i n g t h i s s t u d y , I o b s e r v e d S . l a t e r a l i s w i t h i n d i s t i n c t l a t e r a l s t r i p e s i n P e t e r L o u g h e e d P r o v i n c i a l P a r k , A l b e r t a ( F i g u r e 2 . 1 l o c a t i o n E ) . T h e o c c u r r e n c e o f i n d i s t i n c t s t r i p e s o n S . s a t u r a t u s a n d t h e s e S . l a t e r a l i s i s c o n s i s t e n t w i t h t h e g e n e t i c c o e f f i c i e n t s w h i c h i n d i c a t e d t h a t t h e S . l a t e r a l i s f r o m P e t e r L o u g h e e d P r o v i n c i a l P a r k a r e more c l o s e l y r e l a t e d t o S . s a t u r a t u s t h a n t h e S . l a t e r a l i s f r o m o t h e r l o c a t i o n s s a m p l e d . T h e i d e n t i c a l g e n e t i c p r o f i l e s o f a l l s a m p l e d S . s a t u r a t u s c a n f i t t h e s i n g l e p a n m i c t i c u n i t m o d e l , t h e i s o l a t i o n - b y -d i s t a n c e m o d e l , o r t h e d i s c r e t e s u b p o p u l a t i o n s m o d e l ( R i c h a r d s o n e t a l . 1986) a n d a r e t h u s n o t u s e f u l i n d e t e r m i n i n g t h e a c c e s s i b i l i t y o f i n d i v i d u a l S . s a t u r a t u s f r o m o n e l o c a t i o n t o a n o t h e r . T h e l a c k o f g e n e t i c v a r i a t i o n c a n n o t b e e x p l a i n e d b y h a b i t a t l o s s f o r t h e same r e a s o n a s n o t e d i n t h e s p o t t e d o w l , S t r i x o c c i d e n t a l i s ( B a r r o w c l o u g h a n d G u t i e r r e z 1 9 9 0 ) . T h e number o f g e n e r a t i o n s r e q u i r e d t o r e d u c e g e n e t i c v a r i a t i o n b y r e d u c i n g p o p u l a t i o n s i z e e x c e e d s t h e l e n g t h o f t i m e s i n c e h a b i t a t l o s s b e g a n ( N e i e t a l . 1 9 7 5 ) . B e s i d e s , h a b i t a t f r a g m e n t a t i o n w o u l d c a u s e g e n e t i c h o m o g e n e i t y i n i s o l a t e d l o c a t i o n s , b u t w o u l d a l l o w v a r i a b i l i t y b e t w e e n l o c a t i o n s . T h e a p p a r e n t g e n e t i c u n i f o r m i t y o f S . s a t u r a t u s may b e d u e t o f o u n d e r e f f e c t s ( B a r t o n a n d C h a r l e s w o r t h 1 9 8 4 ) . T h e l o c a t i o n s s a m p l e d a r e o n l y r e p r e s e n t a t i v e o f r e c e n t n o r t h e r n e x t e n s i o n s o f t h e r a n g e o f S . s a t u r a t u s . Low h e t e r o z y g o s i t y o f S . s p i l o s o m a o n P a d r e I s l a n d c o m p a r e d t o m a i n l a n d p o p u l a t i o n s h a s a l s o b e e n a t t r i b u t e d t o f o u n d e r e f f e c t s ( C o t h r a n e t a l . 1 9 7 7 ) . I n b r e e d i n g d e p r e s s i o n i s s o m e t i m e s a s s o c i a t e d w i t h a l a c k o f g e n e t i c v a r i a t i o n ( F r a n k l i n 1980) a n d h a s b e e n a c a u s e o f c o n c e r n f o r s e v e r a l e n d a n g e r e d s p e c i e s i n c l u d i n g c h e e t a h s A c i n o n y x i u b a t u s ( O ' B r i e n e t a l . 1983) a n d g o l d e n l i o n t a m a r i n s , L e o n t o p i t h e c u s r o s a l i a ( R a l l s a n d B a l l o u 1 9 8 6 ) . T h e r i s k o f l o w s u r v i v o r s h i p d u e t o t h e e x p o s u r e o f d e l e t e r i o u s r e c e s s i v e a l l e l e s a n d d e m o g r a p h i c s t o c h a s t i c i t y ( G i l p i n a n d S o u l e 1986 ; L a n d e 1988) o c c u r s when g e n e t i c v a r i a t i o n i s r e d u c e d . T h e d a n g e r h a s o n l y a r i s e n w h e r e mate c h o i c e h a s become r e s t r i c t e d t o more c l o s e l y r e l a t e d i n d i v i d u a l s ( R a l l s e t a l . 1986) t h a n i n t h e p a s t . No e v i d e n c e e x i s t s o n t h e e x t e n t o f m a t i n g among c l o s e k i n i n S . s a t u r a t u s . Enzyme e l e c t r o p h o r e s i s a l s o may n o t h a v e a f i n e e n o u g h r e s o l u t i o n f o r i d e n t i f y i n g g e n e t i c v a r i a t i o n t h a t may b e d e t e c t e d b y DNA s e q u e n c i n g t e c h n i q u e s ( C h a m b e r s a n d B a y l e s s 1 9 8 3 ; A y a l a a n d K i g e r 1984) w h i c h c o u l d p r o v i d e more i n s i g h t on t h e g e n e t i c c h a r a c t e r i s t i c s o f S . s a t u r a t u s o v e r i t s r a n g e . I f S . s a t u r a t u s i s h i g h l y i n b r e d , y e t h a s a g e n o t y p e f l e x i b l e e n o u g h t o c o p e w i t h f l u c t u a t i n g c l i m a t i c c o n d i t i o n s i n i t s r a n g e , t h e n o u t b r e e d i n g d e p r e s s i o n m i g h t a c t u a l l y b e a g r e a t e r r i s k t o t h e s q u i r r e l s t h a n i n b r e e d i n g . T h e b r e a k d o w n o f c o a d a p t e d a l l e l e s c o u l d l e a d t o a l e s s e r a b i l i t y t o c o p e i n i t s e n v i r o n m e n t ( S h e i l d s 1 9 8 2 ) . D i l u t i o n o f g e n e p o o l s h a s b e e n o f c o n c e r n w h e r e h y b r i d s a r e p r o d u c e d i n s p e c i e s s u c h a s r e d w o l v e s ( C a n i s n i g e r ) a n d c o y o t e s ( C a n i s l a t r a n s ) ( P a r k e r 1 9 8 8 ) , a n d wood b i s o n ( B i s o n b i s o n a t h a b a s c a e ) a n d p l a i n s b i s o n ( B i s o n  b i s o n b i s o n ) (Lowe e t a l . 1 9 9 0 ) . W h i l e e l e c t r o p h o r e t i c d a t a a r e l i m i t e d t o c o m p a r i s o n s b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s t e s c o r u m , m o r p h o l o g i c a l d a t a e x t e n d t h e c o m p a r i s o n s t o o t h e r S . l a t e r a l i s s u b s p e c i e s . M o r p h o l o g i c a l l y , I f o u n d more d i f f e r e n c e s e x i s t e d b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s s u b s p e c i e s t h a n among t h e S . l a t e r a l i s s u b s p e c i e s ( T a b l e 2 . 5 ) . I n g e n e r a l , t h e r e was a s i g n i f i c a n t s i z e g r a d i e n t f o r t h e c h a r a c t e r i s t i c s m e a s u r e d , w h i c h , i n o r d e r o f d e c r e a s i n g s i z e , w e r e : S . s a t u r a t u s . S . l a t e r a l i s t e s c o r u m , S . l a t e r a l i s c h r y s o d e i r u s a n d S . l a t e r a l i s  c o n n e c t e n s . T h e l a t t e r two showed no c o n s i s t e n t s i z e d i f f e r e n c e , p o s s i b l y d u e t o i n a d e q u a t e s a m p l e s i z e . H o w e v e r , t h e s i g n i f i c a n t d i f f e r e n c e s b e t w e e n S . l a t e r a l i s t e s c o r u m a n d t h e o t h e r s u b s p e c i e s s u p p o r t s i t s s u b s p e c i f i c s t a t u s . H o w e l l (1938) f o u n d S . l a t e r a l i s c o n n e c t e n s t o b e s l i g h t l y l a r g e r t h a n S . l a t e r a l i s c h r y s o d e i r u s . T h e same g e n e r a l t r e n d i n c r a n i a l c h a r a c t e r i s t i c s f o r t h e s p e c i e s a n d s u b s p e c i e s a n d m a l e s b e i n g l a r g e r t h a n f e m a l e s was r e p o r t e d b y H o w e l l ( 1 9 3 8 ) . W i t h z y g o m a t i c b r e a d t h b e i n g a s i g n i f i c a n t c o v a r i a t e , l a r g e r s k u l l s a p p e a r e d t o b e b r o a d e r d u e t o a w i d e r z y g o m a t i c a r c h , w h i c h i s p a r t i c u l a r l y n o t i c e a b l e i n S . s a t u r a t u s . I n c o n c l u s i o n , t h e p r e s e n c e o f two a l l e l e s t h a t o n l y a p p e a r i n S . s a t u r a t u s , t h e m o r p h o l o g i c a l d i f f e r e n c e s b e t w e e n S . s a t u r a t u s a n d S . l a t e r a l i s . t o g e t h e r w i t h t h e d i s j u n c t r a n g e o f S . s a t u r a t u s t h a t i s o l a t e s i t r e p r o d u c t i v e l y f r o m S . l a t e r a l i s , s u g g e s t t h a t S . s a t u r a t u s s h o u l d b e t r e a t e d a s a d i s t i n c t s p e c i e s , s e p a r a t e f r o m S . l a t e r a l i s . U n t i l m o r e c o n c l u s i v e d a t a a r e a v a i l a b l e , t h e two s h o u l d b e r e g a r d e d a s s i b l i n g s p e c i e s w h i c h a r e s i m i l a r i n m o r p h o l o g y , b u t w i t h p r o b a b l e r e p r o d u c t i v e i s o l a t i o n ( B i c k m a n 1 9 8 3 ) . B e c a u s e i t i s i m p o r t a n t t o c o n s e r v e g e n e t i c d i v e r s i t y ( F r a n k e l a n d S o u l e 1 9 8 1 ; F r a n k e l 1 9 8 4 ) , i t i s a p p r o p r i a t e t o r e c o g n i z e t h e u n i q u e g e n e t i c v a r i a t i o n f o u n d i n S . s a t u r a t u s a n d t o p r e s e r v e i t s e v o l u t i o n a r y l i n e a g e . CHAPTER 3: RANGE AND D I S T R I B U T I O N OF SPERMOPHILUS SATURATUS IN B R I T I S H COLUMBIA INTRODUCTION S . s a t u r a t u s i s u n i q u e t o t h e C a s c a d e m o u n t a i n s o f s o u t h w e s t e r n B r i t i s h C o l u m b i a a n d W a s h i n g t o n S t a t e , a l o n g w i t h o t h e r t a x o n o m i c g r o u p s s u c h a s t h e C a s c a d e f r o g (Rana c a s c a d a e ; N u s s b a u m e t a l . 1983) a n d t h e C a s c a d e f o x ( V u l p e s v u l p e s  c a s c a d e n s i s ; H a l l 1 9 8 1 ) . L i t t l e i s known a b o u t t h e s q u i r r e l ' s N o r t h e r n l i m i t i n B r i t i s h C o l u m b i a ( T r o m b u l a k 1988) e x c e p t f o r d a t a p r e s e n t e d i n H a l l (1981) a n d H o w e l l (1938) . T o h e l p d e t e r m i n e t h e c o n s e r v a t i o n s t a t u s o f S . s a t u r a t u s i n B r i t i s h C o l u m b i a , I i n i t i a t e d a s u r v e y i n B r i t i s h C o l u m b i a i n 1989 t o i m p r o v e t h e k n o w l e d g e o n t h e r a n g e a n d d i s t r i b u t i o n a n d t o c h e c k f o r a n y r e c e n t c h a n g e s i n t h e r a n g e w h i c h may h a v e b e e n i n c u r r e d a s a r e s u l t o f h a b i t a t a l t e r a t i o n . S . s a t u r a t u s f a l l s i n t o a b o d y s i z e a n d f o o d h a b i t c a t e g o r y o v e r l a p p i n g some o t h e r s y m p a t r i c s c i u r i d s . A s a r e s u l t , i t s f e e d i n g s i g n s a n d f e c a l p e l l e t s a r e d i f f i c u l t t o d i s t i n g u i s h f r o m o t h e r s p e c i e s a n d h e n c e c a n n o t b e u s e d t o d e t e r m i n e t h e s q u i r r e l ' s p r e s e n c e a n d a b u n d a n c e . B e c a u s e t h e s q u i r r e l may h i b e r n a t e up t o e i g h t m o n t h s o f t h e y e a r ( K e n a g y a n d B a r n e s 1 9 8 8 ) , s u r v e y i n g i n , t h e f i e l d i s r e s t r i c t e d t o summer. H o w e v e r , S . s a t u r a t u s i s e a s i l y d i s t i n g u i s h e d f r o m o t h e r mammals i n t h e r e g i o n b y i t s p e l a g e c o l o u r a t i o n a n d s i z e ( B u r t a n d G r o s s e n h e i d e r 1976) w h i c h , t o g e t h e r w i t h i t s d i u r n a l b e h a v i o u r , a n d h i g h t o l e r a n c e t o human p r e s e n c e ( B a n f i e l d 1 9 7 4 ) , s u g g e s t t h e p o t e n t i a l f o r v i s u a l - t y p e s u r v e y s a n d may a l s o make s i g h t i n g s b y n o n - p r o f e s s i o n a l n a t u r a l i s t s a s o u r c e o f r e l i a b l e i n f o r m a t i o n . S u r v e y i n g S . s a t u r a t u s i n B r i t i s h C o l u m b i a may n o t o n l y a i d i n d e v e l o p i n g a w i l d l i f e management p l a n , b u t c a n a l s o p r o v i d e a f r a m e w o r k f o r o t h e r s u r v e y s w i t h s i m i l a r o b j e c t i v e s i n v o l v i n g a n i m a l s w i t h s i m i l a r b e h a v i o u r p a t t e r n s . METHODS B e f o r e b e g i n n i n g t h e f i e l d s u r v e y f o r S . s a t u r a t u s a p r e l i m i n a r y r a n g e a n d d i s t r i b u t i o n map was c o n s t r u c t e d f r o m d a t a p r e s e n t e d i n H a l l (1981) a n d f r o m c o l l e c t i o n l o c a t i o n s o f s p e c i m e n s s t o r e d i n museums ( A p p e n d i x 1 ) . New i n f o r m a t i o n was s o u g h t b y d i s t r i b u t i n g s u r v e y f o r m s ( A p p e n d i x 2) t o i n t e r e s t e d i n d i v i d u a l s , r e g i o n a l n a t u r a l i s t c l u b s a n d o t h e r r e l e v a n t o r g a n i z a t i o n s ( A p p e n d i x 3 ) . F i e l d s u r v e y s f o r t h e g r o u n d s q u i r r e l s w e r e i m p l e m e n t e d d u r i n g t h e a c t i v e p o r t i o n o f t h e s q u i r r e l ' s c i r c a n n u a l c y c l e b e t w e e n May a n d S e p t e m b e r i n b o t h 1989 a n d 1 9 9 0 . D u r i n g t h e s e p e r i o d s , I c o n d u c t e d l i v e - t r a p p i n g , s t a t i o n a r y c o u n t s o r w a l k i n g t r a n s e c t s , a n d o p p o r t u n i s t i c o b s e r v a t i o n s ( e . g . r o a d s i g h t i n g s ) . O b s e r v a t i o n s w e r e a i d e d b y 8 x 21 mm b i n o c u l a r s , a 20 x 60 mm s p o t t i n g s c o p e , a n d a h i g h p i t c h e d w h i s t l e . s o u n d e d i n t e r m i t t e n t l y . T h e minimum o b s e r v a t i o n t i m e , i n c l u d i n g c h a n g i n g v i e w p o i n t s , a t e a c h p l a c e was one h o u r . A l t i t u d e was r e c o r d e d f o r e a c h l o c a t i o n . T h e r e c o r d e d l o c a t i o n s w e r e d i v i d e d i n t o t h r e e c a t e g o r i e s : 1) w h e r e S . s a t u r a t u s was s i g h t e d , 2) w h e r e i t was n o t s i g h t e d w i t h i n i t s r a n g e , a n d 3) o u t s i d e i t s r a n g e . W i t h i n e a c h c a t e g o r y , t h e number o f v i s i t s t o d i f f e r e n t l o c a t i o n s was t a l l i e d f o r e a c h a l t i t u d i n a l i n t e r v a l ( 0 - 2 5 0 0 m i n 500 m i n t e r v a l s ) . E a c h a l t i t u d i n a l i n t e r v a l t h a t a t r a n s e c t c r o s s e d t h r o u g h was r e g a r d e d a s a s e p a r a t e v i s i t . I f a t r a n s e c t c r o s s e d more t h a n on e a l t i t u d i n a l i n t e r v a l a n d r e s u l t e d i n a S . s a t u r a t u s s i g h t i n g , o n l y t h e i n t e r v a l w i t h i n w h i c h t h e s i g h t i n g o c c u r r e d was i n c l u d e d i n t h e c a t e g o r y f o r v i s i t s t o p l a c e s w h e r e S . s a t u r a t u s was s i g h t e d . T h e r e m a i n i n g a l t i t u d i n a l i n t e r v a l s o n t h e same t r a n s e c t w e r e p u t i n t h e c a t e g o r y f o r v i s i t s t o p l a c e s w i t h no s i g h t i n g s w i t h i n i t s r a n g e . C o m p a r i n g a l t i t u d e s b e t w e e n t h e c a t e g o r i e s d i s t i n g u i s h e s p o s s i b l e a l t i t u d i n a l p r e f e r e n c e s o f S . s a t u r a t u s f r o m b i a s e s t o w a r d s s a m p l i n g p a r t i c u l a r a l t i t u d e s . T h e r a n g e was d e l i n e a t e d b y j o i n i n g t h e m i d p o i n t s b e t w e e n t h e o u t e r m o s t l o c a t i o n s w h e r e S . s a t u r a t u s w e r e f o u n d a n d a d j a c e n t o u t e r l o c a t i o n s w h e r e i t was . n o t f o u n d . A more c o n s e r v a t i v e e s t i m a t e t h a t may b e a b e t t e r s a f e g u a r d f o r c o n s e r v a t i o n s t r a t e g i e s , w o u l d b e t o j o i n t h e o u t e r m o s t l o c a t i o n s w h e r e S . s a t u r a t u s was s i g h t e d . When t r a p p i n g , Tomahawk #201 l i v e t r a p s w e r e b a i t e d w i t h p e a n u t b u t t e r a n d o a t s . T r a p s w e r e o p e n e d a t 0 8 : 0 0 h o u r s P S T , c l o s e d a t 1 8 : 0 0 h o u r s P S T , a n d c h e c k e d e v e r y two h o u r s . T h e s e x , w e i g h t , l e n g t h a n d age c l a s s ( a d u l t , y e a r l i n g , j u v e n i l e ) e s t i m a t e d f r o m s i z e a n d r e p r o d u c t i v e c o n d i t i o n , w e r e r e c o r d e d f o r a l l S . s a t u r a t u s c a p t u r e d . E a c h i n d i v i d u a l was i d e n t i f i e d w i t h a m e t a l e a r - t a g c a r r y i n g a u n i q u e t h r e e d i g i t number a n d a u n i q u e c o l o u r c o m b i n a t i o n o f y a r n . t i e d o n t o t h e e a r t a g t o p e r m i t i d e n t i f i c a t i o n o f i n d i v i d u a l s f r o m a d i s t a n c e . I n p r o v i n c i a l p a r k s w h e r e l e s s p e r m a n e n t a n d more c r y p t i c m a r k i n g i s p r e f e r r e d , b l a c k L ' o r e a l h a i r c o l o u r i n g was a p p l i e d t o t h e p e l a g e f o r t h e i d e n t i f i c a t i o n o f i n d i v i d u a l s . T h e p r o b a b i l i t y o f s i g h t i n g a S . s a t u r a t u s w i t h i n a c e r t a i n t i m e i n t e r v a l was c a l c u l a t e d b y c o r r e l a t i n g t h e c u m u l a t i v e p e r c e n t s i g h t i n g s a t l o c a t i o n s o f known i n h a b i t a t i o n t o c u m u l a t i v e t i m e s p e n t o b s e r v i n g a t t h e s e s i t e s . T h i s a l l o w e d me t o p r e d i c t t h e amount o f t i m e n e e d e d a t e a c h l o c a t i o n i n o r d e r t o d e t e c t S . s a t u r a t u s i f i t was t h e r e . R E S U L T S C o o r d i n a t e s f o r l o c a t i o n s w h e r e S . s a t u r a t u s was s i g h t e d a n d t h e i r c o r r e s p o n d i n g l e t t e r s i n F i g . 3 . 1 a r e l i s t e d i n A p p e n d i x 4 , a l o n g w i t h t h e i r r e s p e c t i v e b i o g e o c l i m a t i c z o n e , a l t i t u d e , a n d m e t h o d o f s u r v e y b y w h i c h t h e s q u i r r e l was d e t e c t e d . T h e l o c a t i o n s , t h e i r r e s p e c t i v e b i o g e o c l i m a t i c z o n e R a n g e o f S p e r m o p h i l u s s a t u r a t u s i n B r i t i s h C o l u m b i a P r e s e n t r a n g e P r e v i o u s known r a n g e Kilometres 0 5 LEGEND • Major point surveyed without sighting Spermophilus saturatus ^ Museum specimen of Spermophilus  saturatus collected prior to 1989 0 Sighting ol Soermoohilus saturatus during the 1989-1990 survey -|- Sighting ol Soermoohilus lateralis during the 1989-1990 survey Fig. 3.1. Map of survey area. Location names corresponding to numbers and letters are listed in Appendices 4 and 5. Fig. 3.1. Map of survey area. Location names corresponding to numbers and letters are listed i, • Appendices 4 and 5. 00 a n d a l t i t u d e , w h e r e no S . s a t u r a t u s was d e t e c t e d b y o b s e r v a t i o n , a n d t h e i r c o r r e s p o n d i n g n u m b e r s i n F i g . 3 . 1 , a r e l i s t e d i n A p p e n d i x 5 . T h e c l a s s i f i c a t i o n s f o r b i o g e o c l i m a t i c z o n e s a r e b a s e d o n K r a j i n a ( 1 9 5 9 ) , b u t M i t c h e l l a n d G r e e n (1981) p r o v i d e a more r e c e n t v e r s i o n o f t h e c l a s s i f i c a t i o n f o r t h e K a m l o o p s r e g i o n . T h e s e b i o g e o c l i m a t i c c l a s s i f i c a t i o n f o r i n d i v i d u a l l o c a t i o n s ( A p p e n d i c e s 4 a n d 5) c a n n o t b e g e n e r a l l y c o m p a r e d b e c a u s e t h e y a r e n o t a v a i l a b l e f o r a l l l o c a t i o n s , a n d t h e s c a l e t o w h i c h t h e y a p p l y i s t o o l a r g e t o d i s c r i m i n a t e s u i t a b l e f r o m u n s u i t a b l e h a b i t a t . N i n e o f 3 5 s u r v e y f o r m s d i s t r i b u t e d w e r e r e s p o n d e d t o o f w h i c h s e v e n w e r e a c t u a l s i g h t i n g s ( A p p e n d i x 3) . T h e s e h a v e b e e n i n c l u d e d i n F i g . 3 . 1 . T r a p p i n g e f f o r t , l o c a t i o n s , a n d s m a l l mammals c a u g h t a r e l i s t e d i n A p p e n d i x 6 . T r a p p i n g d i d n o t l e a d t o t h e d i s c o v e r y o f new l o c a t i o n s o c c u p i e d b y S . s a t u r a t u s i n h a b i t a t i o n , a s t r a p p i n g was o n l y s u c c e s s f u l w h e r e S . s a t u r a t u s h a d a l r e a d y b e e n s i g h t e d . H o w e v e r , a n age a n d s e x c o m p o s i t i o n a n d minimum p o p u l a t i o n e s t i m a t e was o b t a i n e d by t r a p p i n g a t f o u r known l o c a t i o n s o f S . s a t u r a t u s i n h a b i t a t i o n ( F i g . 3 . 2 ) . T h e mean a g e a n d s e x c o m p o s i t i o n i n 1990 f o r t h e f o u r l o c a t i o n s was : a d u l t f e m a l e (31%), a d u l t m a l e (25%), y e a r l i n g f e m a l e (24%), y e a r l i n g m a l e (21%). T r a p p i n g i n c o n s e c u t i v e y e a r s t o o k p l a c e o n l y a t T h y n n e M o u n t a i n . H e r e , o f t h r e e a d u l t f e m a l e s , f o u r a d u l t m a l e s , two y e a r l i n g f e m a l e s a n d two y e a r l i n g m a l e s m a r k e d i n 1 9 8 9 , two a d u l t f e m a l e s , two a d u l t m a l e s , two y e a r l i n g f e m a l e s , a n d one Cascade Lookout Brookmere Dump Iron Mtn Thynne Mtn 1990 Thynne Mtn 1989 Location F i g . 3 . 2 . A g e a n d s e x c o m p o s i t i o n o f S . s a t u r a t u s a t f o u r l o c a t i o n s . C a s c a d e L o o k o u t a n d B r o o k m e r e Dump e a c h h a d 80 t r a p p i n g d a y s w h i l e I r o n M o u n t a i n and T h y n n e M o u n t a i n e a c h h a d 120 t r a p p i n g d a y s . S e e A p p e n d i x 6 f o r d e t a i l s . 31 y e a r l i n g m a l e w e r e r e c a p t u r e d i n 1 9 9 0 . I n a d d i t i o n , f i v e S . s a t u r a t u s w e r e c a p t u r e d f o r t h e f i r s t t i m e i n 1 9 9 0 . O f 91 l o c a t i o n s s u r v e y e d , 36 w e r e i n h a b i t e d b y S . s a t u r a t u s . T h e s e s i g h t i n g s o f S . s a t u r a t u s made i n 1989 a n d 1990 i n d i c a t e t h a t t h e i r r a n g e i s f u r t h e r n o r t h a n d w e s t t h a n d o c u m e n t e d b y museum s p e c i m e n s ( F i g . 3 . 1 ) . F o r e a c h o f t h e f o u r t r a p p i n g l o c a t i o n s , t h e p e r c e n t a g e o f " t i m e t o f i r s t s i g h t i n g " f o r e a c h t i m e c a t e g o r y was c a l c u l a t e d ( T a b l e 3 . 1 ) . A l t h o u g h o b s e r v a t i o n a l p e r i o d s d i d n o t a l w a y s l a s t u n t i l S . s a t u r a t u s was s i g h t e d , t h e "60 +" t i m e c a t e g o r y i n T a b l e 3 . 1 a s s u m e s t h a t S . s a t u r a t u s w o u l d h a v e b e e n s i g h t e d g i v e n e n o u g h t i m e . T o s u p p o r t t h i s a s s u m p t i o n , f o u r a r t i f i c i a l p o i n t s (one f o r e a c h l o c a t i o n ) w e r e c r e a t e d f o r a c u m u l a t i v e p e r c e n t s i g h t i n g v a l u e o f 100 % a t 1 0 , 0 0 0 c u m u l a t i v e m i n u t e s . T h e s e p o i n t s w e r e i n c l u d e d i n t h e d a t a f o r f i n d i n g t h e b e s t f i t c u r v e , b u t a r e n o t d e p i c t e d i n F i g . 3 . 3 . T h e a v e r a g e c u m u l a t i v e p e r c e n t s i g h t i n g s (y) f o r t h e f o u r l o c a t i o n s w e r e p l o t t e d a g a i n s t c u m u l a t i v e t i m e ( t ) . T h e b e s t f i t c u r v e , y = 7 8 ( l - e " 0 - U t ) w i t h R 2 = 0 . 9 9 , r e a c h e s a n a s y m p t o t e a t 78% ( F i g . 3 . 3 ) . A l t h o u g h R 2 i s h i g h , t h e d a t a p o i n t s a r e w i d e l y s c a t t e r e d . I f t h e f o u r a r t i f i c i a l p o i n t s a r e e x c l u d e d f r o m t h e d a t a s e t , t h e b e s t f i t c u r v e i s y = 7 0 ( l - e " ° - 2 1 t ) w i t h R 2 = 0 . 9 9 . U s i n g " c u m u l a t i v e p e r c e n t s i g h t i n g s " i m p l i e s t h a t t h e p e r c e n t s i g h t i n g s a t e a c h t i m e p o i n t i s d e p e n d e n t o n t h e p e r c e n t s i g h t i n g s a t t h e p r e v i o u s t i m e p o i n t . T h i s may b i a s t h e R 2 v a l u e u p w a r d s . T h e l o c a t i o n s a r e c l a s s e d a s p l a c e s w h e r e S . s a t u r a t u s was s i g h t e d ( i . e . s i g h t i n g ) , p l a c e s i n t h e r a n g e w h e r e no S . 32 T a b l e 3 . 1 . P e r c e n t o f t o t a l f i r s t s i g h t i n g s o c c u r r i n g w i t h i n d i f f e r e n t 10 m i n u t e p e r i o d s a t f o u r l o c a t i o n s . Iron Mountain Time category Thynne Mountain Cascade Lookout Brookmere Dump (minutes) 0 -10 20 23 73 50 11-20 0 31 0 33 21 -30 0 15 9 17 31 -40 13 0 0 0 41 - 50 0 8 0 0 51-60 13 0 0 0 60 + 53 23 18 0 Number of sightings 15 13 11 6 Minimum number of animals 7 11 14 23 F i g . 3 . 3 . P r o b a b i l i t y o f s i g h t i n g S . s a t u r a t u s a s d e t e r m i n e d b y c u m u l a t i v e p e r c e n t s i g h t i n g s o v e r t i m e s i n c e b e g i n n i n g o f o b s e r v a t i o n p e r i o d . 3 4 s a t u r a t u s was s i g h t e d ( i . e . i n s i d e r a n g e ) a n d p l a c e s o u t s i d e t h e r a n g e ( i . e . o u t s i d e r a n g e ) . T h e K o l m o g o r o v - S m i r n o v o n e -s a m p l e t e s t d e t e c t e d no d i f f e r e n c e s t o w a r d s s a m p l i n g p a r t i c u l a r a l t i t u d e s i n s i d e a n d o u t s i d e t h e r a n g e (D = 0 . 1 1 6 , P > 0 . 0 5 , N = 3 7 ; F i g . 3 . 4 . ) . W i t h i n t h e r a n g e o f S . s a t u r a t u s t h e a l t i t u d i n a l d i s t r i b u t i o n o f t h e number o f v i s i t s w i t h s i g h t i n g s o f S . s a t u r a t u s w e r e n o t d i f f e r e n t f r o m t h e a l t i t u d i n a l d i s t r i b u t i o n o f v i s i t s w i t h o u t s i g h t i n g s o f S . s a t u r a t u s ( F i g . 3 . 5 ; K o l m o g o r o v - S m i r n o v o n e - s a m p l e t e s t , D = 0 . 0 4 3 , P > 0 . 0 5 , N = 33) . D I S C U S S I O N R a n g e a n d D i s t r i b u t i o n T h e r a n g e a n d d i s t r i b u t i o n o f a s p e c i e s i s d e t e r m i n e d by t h e b i o l o g i c a l c o n s t r a i n t s o f t h e o r g a n i s m a n d t h e b i o p h y s i c a l a t t r i b u t e s o f t h e l a n d s c a p e . T h e e c o l o g i c a l n i c h e c o n c e p t i n c o r p o r a t e s b o t h o f t h e s e c h a r a c t e r i s t i c s ( S c h o e n e r 1 9 8 9 ) , a n d t h e y a r e i n s e p a r a b l e when d e c i d i n g w h i c h f a c t o r s may l i m i t t h e r a n g e a n d d i s t r i b u t i o n o f S . s a t u r a t u s . P h y s i c a l b a r r i e r s c a n i n c l u d e a n y c o m b i n a t i o n o f h a b i t a t f e a t u r e s t h a t c a n n o t b e o v e r c o m e b y t h e d i s p e r s a l c a p a b i l i t y o r p h y s i o l o g i c a l t o l e r a n c e o f t h e a n i m a l . R i v e r s a r e o b v i o u s p o t e n t i a l b o u n d a r i e s t o s m a l l t e r r e s t r i a l mammals s u c h a s S . s a t u r a t u s a n d t h r e e d r a i n a g e s y s t e m s s u r r o u n d i t s r a n g e i n 45-40-35-30-(JO > 25-1 § 20H 0) 15-10-5-Outside range Inside range 0-499 500-999 1000-1499 1500-1999 2000-2499 2500-2999 Altitudinal Range (m) F i g . 3 . 4 . T h e p r o p o r t i o n o f v i s i t s t o p a r t i c u l a r a l t i t u d e s i n s i d e (n=67) a n d o u t s i d e (n=37) w t h e r a n g e o f S . s a t u r a t u s . w F i g . 3 . 5 . T h e p r o p o r t i o n o f v i s i t s t o p a r t i c u l a r a l t i t u d e s i n s i d e t h e r a n g e o f S . s a t u r a t u s w t h a t r e s u l t e d i n s i g h t i n g s . S a m p l e s i z e : s i g h t i n g s ( 3 3 ) , n o n s i g h t i n g s ( 3 4 ) . B r i t i s h C o l u m b i a . T o t h e w e s t i s t h e F r a s e r R i v e r ; t o t h e n o r t h i s t h e N i c o l a R i v e r s y s t e m ; a n d t o t h e e a s t i s t h e O k a n a g a n s y s t e m . T h e l a t t e r a l s o a p p e a r s t o f o r m t h e w e s t e r n b a r r i e r o f S . l a t e r a l i s . r e n d e r i n g t h e r a n g e s o f t h e two s i b l i n g s p e c i e s c o m p l e t e l y d i s j u n c t i n C a n a d a . T h e two a r e n o t known t o o v e r l a p e l s e w h e r e ( H a l l 1 9 8 1 ) . G l a c i e r s , a n d s u b s e q u e n t d r a i n a g e p a t t e r n s , c o u l d h a v e i s o l a t e d members o f t h e s u b g e n u s C a l l o s p e r m o p h i l u s ( H o w e l l 1938) i n t o s e v e r a l r e f u g i a d u r i n g o r f o l l o w i n g t h e P l e i s t o c e n e . S u c h i s o l a t i o n c o u l d h a v e p r o v i d e d t h e c i r c u m s t a n c e s n e c e s s a r y f o r a l l o p a t r i c s p e c i a t i o n ( H a f f e r 1 9 8 2 ) . S . s a t u r a t u s p r o b a b l y s p r e a d n o r t h w a r d i n t o B r i t i s h C o l u m b i a f r o m t h e s o u t h e r n C a s c a d e M o u n t a i n R a n g e a f t e r t h e r e t r e a t o f t h e C o r d i l l e r a n I c e S h e e t . T h e s e g l a c i e r s c o v e r e d m o s t o f s o u t h e r n B r i t i s h C o l u m b i a a n d e x t e n d e d a s f a r a s t h e P u g e t I n l e t i n W a s h i n g t o n S t a t e 1 7 , 5 0 0 B . P . ( F u l t o n e t a l . 1 9 8 6 ) . T h e S k a g i t V a l l e y t h a t s e p a r a t e s t h e S . s a t u r a t u s n e a r W i l l i a m ' s P e a k ( L o c a t i o n AC i n F i g . 3 . 1 , a n d A p p e n d i x 4) f r o m t h e o t h e r l o c a t i o n s o f S . s a t u r a t u s i n B r i t i s h C o l u m b i a s u g g e s t s t h i s h i s t o r i c a l d i s p e r s a l . S . s a t u r a t u s a t W i l l i a m ' s P e a k w e r e p r o b a b l y c o l o n i z e d f r o m t h e s o u t h a n d n o t f r o m t h e e a s t . A l t h o u g h t h e d r a i n a g e s y s t e m s may l i m i t t h e d i s p e r s a l o f S . s a t u r a t u s . t h e y do s o o n l y o n a b r o a d s c a l e a n d o t h e r v a r i a b l e s a r e p r o b a b l y i n v o l v e d . F o r e x a m p l e , i f t e m p e r a t u r e e x p l a i n s t h e n o r t h e r n b o u n d a r y o f t h e r a n g e , t h e e l e v a t i o n l i m i t a t i o n s o f t h e a n i m a l s h o u l d d e c r e a s e a s l a t i t u d e i n c r e a s e s . D e c r e a s e d r e p r o d u c t i o n h a s b e e n a s s o c i a t e d w i t h S p e r m o p h i l u s s p e c i e s 38 l i v i n g i n c o m p r e s s e d e n v i r o n m e n t s ( B r o n s o n 1 9 7 9 ; P h i l i p s 1984) w h e r e t h e g r o w i n g s e a s o n f o r p l a n t s i s r e l a t i v e l y s h o r t . Brown (1971) p r e s e n t s d a t a f r o m t h e G r e a t B a s i n s u g g e s t i n g t h a t mammals o n m o u n t a i n t o p s a r e p o o r c o l o n i z e r s . H o w e v e r , e n v i r o n m e n t a l v a r i a b l e s t h a t may r e s t r i c t t h e d i s p e r s a l o f S . s a t u r a t u s w i l l n o t be d i s c u s s e d f u r t h e r h e r e s i n c e t h e y h a v e n o t b e e n e x p e r i m e n t a l l y t e s t e d . T h e i n c r e a s e d r a n g e e x t e n d i n g n o r t h w a r d a n d w e s t w a r d a s f o u n d d u r i n g t h i s s u r v e y p r o b a b l y r e f l e c t s e n h a n c e d human a c c e s s t o t h e s e a r e a s i n h i s t o r i c a l t i m e , a n d n o t a r a n g e e x p a n s i o n b y t h e s q u i r r e l . T h e s h i f t o f t h e e a s t e r n b o u n d a r y w e s t w a r d i n B r i t i s h C o l u m b i a ( F i g . 3 . 1 ) may o n l y b e a p p a r e n t d u e t o i n a d e q u a t e s a m p l i n g e f f o r t a r o u n d t h e e a s t e r n b o u n d a r y . I t i s more l i k e l y t h a t t h e s h i f t i s r e a l a n d r e f l e c t s t h e c o m p e t i t i v e e x c l u s i o n o f S . s a t u r a t u s b y S . c o l u m b i a n u s . R o a d s b u i l t d u r i n g t h i s c e n t u r y f a c i l i t a t e d t h e w e s t w a r d d i s p e r s a l o f S . c o l u m b i a n u s ( D a v i d N a g o r s e n , p e r s . comm.) e i t h e r b y c r e a t i n g c o r r i d o r s o f s u i t a b l e h a b i t a t o r b y v e h i c l e s u n k n o w i n g l y t r a n s p o r t i n g t h e a n i m a l s t o s u i t a b l e h a b i t a t . I n B r i t i s h C o l u m b i a , S . s a t u r a t u s h a v e r e s i d e d i n M a n n i n g P r o v i n c i a l P a r k s i n c e i t s d e s i g n a t i o n i n 1 9 4 1 , b u t S . c o l u m b i a n u s was f i r s t o b s e r v e d o n l y a f t e r t h e c o n s t r u c t i o n o f t h e h i g h w a y t r a v e r s i n g t h e p a r k ( T e d U n d e r h i l l , p e r s . c o m m . ) . H a b i t a t d i s p l a c e m e n t f a c i l i t a t e d b y human a c t i v i t i e s h a s b e e n r e p o r t e d r e c e n t l y f o r t h e mouse (Mus s p r e t o i d e s ) i n I s r a e l ( A u f f r a y e t a l 1990) a n d r o u n d t a i l c h u b ( G i l a r o b u s t a ) i n New M e x i c o ( B e s t g e n a n d P r o p s t 1 9 8 9 ) . T h e c o m p e t i t i v e e x c l u s i o n p r i n c i p l e s t a t e s t h a t i f t h e d i f f e r e n t i a t i o n b e t w e e n t h e r e a l i z e d n i c h e s o f two c o m p e t i n g s p e c i e s i s p r e c l u d e d b y t h e h a b i t a t l i m i t a t i o n s , t h e n one s p e c i e s w i l l e x c l u d e t h e o t h e r (Begon a n d M o r t i m e r 1 9 8 1 ) . Two t y p e s o f c o m p e t i t i v e e x c l u s i o n a r e p o s s i b l e , e x p l o i t a t i v e a n d i n t e r f e r e n c e . T h e e x p l o i t a b l e n i c h e c o m p o n e n t s o f S . s a t u r a t u s b y S . c o l u m b i a n u s a r e f o o d a n d s h e l t e r . B o t h a n i m a l s a r e d i u r n a l s e m i f o s s o r i a l s q u i r r e l s . T h e r e i s some o v e r l a p p i n g f o r a g e s p e c i e s , i n c l u d i n g L u p i n u s s p p . a n d T a r a x a c u m s p p . ( T r o m b u l a k 1988 ; E l l i o t a n d F l i n d e r s 1985 ; A n d r u s i a k a n d H a r e s t a d 1 9 8 7 ) . Meadows t y p i f y S . c o l u m b i a n u s h a b i t a t ( B u r t a n d G r o s s e n h e i d e r 1976) w h e r e a s S . s a t u r a t u s u s e b o t h f o r e s t a n d meadow h a b i t a t ( T r o m b u l a k 1 9 8 7 ) . E v i d e n c e o f c o m p e t i t i v e e x c l u s i o n h a s b e e n o b s e r v e d b e t w e e n S . l a t e r a l i s ( s i b l i n g s p e c i e s o f S . s a t u r a t u s ) a n d S . c o l u m b i a n u s i n R o s s l a n d , B r i t i s h C o l u m b i a . A t one s i t e w h e r e a l l S . c o l u m b i a n u s w e r e r e m o v e d , S . l a t e r a l i s s u b s e q u e n t l y i n h a b i t e d t h e a r e a ( p e r s . o b s e r v . ) . I f i n t e r f e r e n c e c o m p e t i t i o n d o e s o c c u r , t h e l a r g e r s i z e o f S . c o l u m b i a n u s ( 3 4 0 - 8 1 2 g) c o m p a r e d t o S . s a t u r a t u s ( 1 7 0 - 2 7 6 g) ( B u r t a n d G r o s s e n h e i d e r 1976) w o u l d p r o b a b l y p u t t h e l a t t e r a t a m a j o r d i s a d v a n t a g e . I n meadow h a b i t a t , S . c o l u m b i a n u s , w i t h i t s c o o p e r a t i v e a n d c o l o n i a l s o c i a l s y s t e m , c o u l d o u t n u m b e r S . s a t u r a t u s . w i t h i t s s o l i t a r y b e h a v i o u r ( M i c h e n e r 1 9 8 3 ) . O t h e r s c i u r i d s a l s o l i v e i n t h e r a n g e o f S . s a t u r a t u s . b u t p e r h a p s b e c a u s e o f t h e i r l o n g e r h i s t o r y o f c o e x i s t e n c e c o m p a r e d t o t h e r e c e n t i n t e r a c t i o n s b e t w e e n S . s a t u r a t u s a n d S . c o l u m b i a n u s . c o m p e t i t i o n f o r f o o d r e s o u r c e s i s m i n i m a l . I n p a r t i c u l a r , t h e y e l l o w p i n e c h i p m u n k ( E u t a m i a s a m o e n u s ) , t h e s c i u r i d f o u n d m o s t commonly i n t h e same h a b i t a t a s S . s a t u r a t u s , e a t s many o f t h e same f o o d s p e c i e s . N e v e r t h e l e s s , S . s a t u r a t u s a n d E . amoenus t o l e r a t e e a c h o t h e r i n c l o s e p r o x i m i t y , a n d w h i l e S . s a t u r a t u s f e e d m a i n l y on t h e l e a v e s a n d s t e m s , E . amoenus f e e d m a i n l y o n t h e s e e d s o f t h e same p l a n t s p e c i e s ( B r o a d b o o k s 1 9 5 8 ) . R e d s q u i r r e l s ( T a m i a s c i u r u s h u d s o n i c u s ) a r e s o m e t i m e s f o u n d i n t h e same a r e a a s S . s a t u r a t u s . b u t t h e y d i f f e r f r o m t h e g r o u n d s q u i r r e l b y n e s t i n g i n t r e e s i n t h e summer, a n d f e e d i n g p r i m a r i l y on c o n i f e r s e e d s ( B a n f i e l d 1 9 7 4 ) . B e s i d e s s h i f t i n g t h e r a n g e b o u n d a r y o f S . s a t u r a t u s , S . c o l u m b i a n u s may b e a f f e c t i n g t h e d i s t r i b u t i o n o f S . s a t u r a t u s D p o p u l a t i o n s w h e r e S . s a t u r a t u s e x i s t o n m a r g i n a l h a b i t a t . T h e a b s e n c e o f S . s a t u r a t u s a t s i t e s w i t h i n i t s r a n g e c o u l d b e c a u s e d b y t h i s a n d o t h e r f o r m s o f h a b i t a t a l t e r a t i o n . B e t t s (1990) a n d T o m i c h (1982) s u g g e s t t h a t t h e p r e s e n t d i s t r i b u t i o n o f o t h e r S p e r m o p h i l u s s p e c i e s i s a f f e c t e d b y p a s t e p i d e m i c s o f p l a g u e ( Y e r s i n i a p e s t i s ) . A t t h e t u r n o f t h e c e n t u r y , p l a g u e t h a t o r i g i n a t e d f r o m Y u n n a n p r o v i n c e i n C h i n a , was i n t r o d u c e d i n t o w i l d p o p u l a t i o n s o f mammals i n U n i t e d S t a t e s f r o m n o n - n a t i v e r a t s v i a f l e a s ( M i t c h e l l 1 9 6 6 ) . T h e p l a g u e b a c i l l u s was f o u n d i n S . l a t e r a l i s , t h e s i b l i n g s p e c i e s o f S . s a t u r a t u s , i n w e s t e r n U n i t e d S t a t e s . D i e o f f s o f g r o u n d s q u i r r e l c o l o n i e s w e r e o b s e r v e d a n d a l t h o u g h t h i s was n o t s e e n w i t h S . l a t e r a l i s o r S . s a t u r a t u s . t h e y a r e s u s c e p t i b l e h o s t s ( S t a r k e t a l . 1 9 6 6 ) . T h e f l e a s i d e n t i f i e d f r o m S . s a t u r a t u s d u r i n g t h e p r e s e n t s u r v e y w e r e O r o p s y l l a i d a h o e n s i s w h i c h may m a i n t a i n s y l v a t i c p l a g u e i n g r o u n d s q u i r r e l p o p u l a t i o n s ( L e w i s e t a l . 1 9 8 8 ) . S i n c e h i s t o r i c a l a c c o u n t s a r e n o t a v a i l a b l e , t h e r o l e o f d i s e a s e o n t h e d i s t r i b u t i o n o f S . s a t u r a t u s c a n n o t b e s u b s t a n t i a t e d . Many f a c t o r s a f f e c t t h e g e o g r a p h i c a l r a n g e a n d d i s t r i b u t i o n o f a s p e c i e s . F o r S . s a t u r a t u s , t h e s e may i n c l u d e d i s p e r s a l b a r r i e r s , g e o l o g i c a l h i s t o r y , c l i m a t e , h a b i t a t r e q u i r e m e n t s , c o m p e t i t i o n a n d d i s e a s e . T h e s e a r e n o t m u t u a l l y e x c l u s i v e ; t h e y may i n t e r a c t t o d e t e r m i n e w h e r e S . s a t u r a t u s o c c u r s . A s s e s s m e n t o f S u r v e y M e t h o d s I n t h i s s e c t i o n , I c o n s i d e r e d t h e p o t e n t i a l v a l u e s a n d l i m i t a t i o n s o f e a c h o f t h e f i v e s u r v e y m e t h o d s u s e d a n d make r e c o m m e n d a t i o n s f o r t h e i r u s e i n f u t u r e r a n g e a n d d i s t r i b u t i o n s u r v e y s . A p p r o a c h e s t h a t c o u l d b e t a k e n t o m a x i m i z e t h e p o t e n t i a l i n f o r m a t i o n w i t h i n t i m e a n d c o s t c o n s t r a i n t s w i l l a l s o be d i s c u s s e d . H i s t o r i c a l r e c o r d s , s u c h a s museum s p e c i m e n s , p r o v i d e t h e b a s i s f o r s t u d i e s o n g e o g r a p h i c a l d i s t r i b u t i o n ( M o n d o l f i 1989 ; B e t t s 1 9 9 0 ) . A l t h o u g h l o c a t i o n s w h e r e S . s a t u r a t u s w e r e c o l l e c t e d w e r e o f t e n n o t s p e c i f i c e n o u g h t o r e v i s i t , t h e r e c o r d s g a v e e n o u g h i n f o r m a t i o n t o c o n s t r u c t a r o u g h o u t l i n e o f t h e r a n g e ( F i g . 3 . 1 ) . O n c e t h i s r o u g h o u t l i n e was e s t a b l i s h e d , t h e s u r v e y t h a t I c o n d u c t e d c o u l d d e t e c t c h a n g e s i n t h e r a n g e . A b u n d a n c e c a n n o t be e s t i m a t e d f r o m museum c o l l e c t i o n s u n l e s s e x t e n s i v e t r a p p i n g was d o n e a n d r e c o r d s o f t r a p p i n g e f f o r t a r e k e p t . 42 T h e u s e o f s u r v e y f o r m s m a i l e d o u t t o v a r i o u s p a r t i e s i s o f q u e s t i o n a b l e v a l u e u n l e s s more c a r e f u l l y t a r g e t t e d . A l t h o u g h o n l y -26% o f t h e 35 f o r m s w e r e r e t u r n e d , t h e s e r e t u r n s s t i l l c o n t r i b u t e d t o 22% o f t h e t o t a l s i g h t i n g s . H o w e v e r , a l l r e s p o n s e s w i t h a c t u a l s i g h t i n g s o f S . s a t u r a t u s came f r o m p e o p l e whom I c o n t a c t e d p e r s o n a l l y a b o u t t h e p r o j e c t a n d e x p l a i n e d t h e v a l u e o f S . s a t u r a t u s s i g h t i n g s . T h e m o s t a p p r o p r i a t e t a r g e t g r o u p s w o u l d i n c l u d e p e o p l e w h e r e p e r s o n a l c o n t a c t becomes e s t a b l i s h e d d u r i n g f i e l d s u r v e y s , a n d p a r k s t a f f who a r e e m p l o y e d i n t h e r a n g e o f t h e a n i m a l . O p p o r t u n i s t i c s i g h t i n g s r e c o r d e d b y m y s e l f a n d two f i e l d a s s i s t a n t s c o m p r i s e d a s u r p r i s i n g 35% o f t h e s i g h t i n g s . T h e d r a w b a c k s o f t h e t e c h n i q u e i s t h a t i t i s n o t s y s t e m a t i c s i n c e s a m p l i n g was b i a s e d t o w a r d s t r a n s p o r t a t i o n c o r r i d o r s a n d c r e d e n c e c a n o n l y b e g i v e n t o S . s a t u r a t u s p r e s e n c e , n o t i t s a b s e n c e . H o w e v e r , t h e t e c h n i q u e c o u l d p r o v i d e v a l u a b l e s t a r t i n g p o i n t s t o b e g i n s y s t e m a t i c s e a r c h e s . T o p r e p a r e f i e l d w b r k e r s f o r o p p o r t u n i s t i c s i g h t i n g s , a l l b i r d a n d mammal a c t i v i t y s e e n d u r i n g t h e c o u r s e o f a w o r k d a y s h o u l d be n o t e d . T h i s m a i n t a i n s t h e o b s e r v a t i o n a n d a w a r e n e s s s k i l l s o f w o r k e r s , p a r t i c u l a r l y when t r a v e l l i n g b e t w e e n l o c a t i o n s . M o s t s i g h t i n g s (43%) w e r e d e t e c t e d b y w a l k i n g / o b s e r v a t i o n s u r v e y s . M o s t t i m e was a l s o d e v o t e d t o t h i s m e t h o d o f s u r v e y . A l t h o u g h s y s t e m a t i c s a m p l i n g was a t t e m p t e d , p a t h w a y s a n d t e r r a i n made some l o c a t i o n s much l e s s a c c e s s i b l e t h a n o t h e r s . S y s t e m a t i c s a m p l i n g , e s p e c i a l l y t o d e t e r m i n e a n y p o s s i b l e a l t i t u d i n a l p r e f e r e n c e s o f S . s a t u r a t u s , s h o u l d b e i n c l u d e d i n f u t u r e w o r k . P e r h a p s a l t i t u d i n a l p r e f e r e n c e was n o t d e t e c t e d i n t h i s s t u d y b e c a u s e o f i n a d e q u a t e s a m p l i n g a t t h e d i f f e r e n t a l t i t u d e s , p a r t i c u l a r l y l o w e r e l e v a t i o n s . B e f o r e s e t t i n g o u t t o do a w a l k i n g / o b s e r v a t i o n s u r v e y , t o p o g r a p h i c a l maps a n d t h e m o s t r e c e n t i n f o r m a t i o n o n a c c e s s r o u t e s s h o u l d be s o u g h t . T h e n a s c h e d u l e s h o u l d b e d r a f t e d , t a k i n g i n t o c o n s i d e r a t i o n t h e p r o x i m i t y o f l o c a t i o n s , t h e r e l a t i v e n e e d t o s a m p l e e a c h l o c a t i o n , a n d t h e t i m e when snow, f l o o d s a n d o t h e r phenomenon a r e l e a s t l i k e l y t o b e a h i n d r a n c e . S a m p l i n g e f f o r t o f d i r e c t o b s e r v a t i o n s u r v e y s c o u l d n o t be a c c u r a t e l y m e a s u r e d u s i n g t i m e s p e n t b e c a u s e o b s e r v a b i l i t y was a f f e c t e d b y w e a t h e r c o n d i t i o n s , v i s u a l o b s t r u c t i o n s ( e g . t r e e s a n d f o g ) , t i m e o f d a y , t i m e o f y e a r , a n d h a b i t u a t i o n t o human a c t i v i t y ( L o c a t i o n s F , A A , A E , A F , AG i n F i g . 3 . 1 a n d A p p e n d i x 4) . B a i l e y (1986) s t a t e s t h a t d i r e c t a n d i n d i r e c t e f f e c t s o f w e a t h e r o n a c t i v i t y a r e more a c u t e f o r s m a l l a n i m a l s t h a n f o r l a r g e a n i m a l s , a n d f o r p o p u l a t i o n s l i v i n g n e a r t h e e d g e s o f t h e i r g e o g r a p h i c a l r a n g e . B o t h p o i n t s a p p l y t o t h i s s u r v e y f o r S . s a t u r a t u s . T h i s g r o u n d s q u i r r e l i s more a c t i v e o n c l e a r d a y s ( B a n f i e l d 1974 , p e r s . o b s e r v . ) a n d d a i l y a c t i v i t y p a t t e r n s a p p e a r e d t o v a r y f r o m s i t e t o s i t e . Where f o o d h a n d o u t s a r e a v a i l a b l e , i n i t i a l a c t i v i t y was l a t e r a n d s q u i r r e l s made t h e i r p r e s e n c e known when p e o p l e w e r e p r e s e n t . D e s p i t e t h e d i f f i c u l t i e s a s s o c i a t e d w i t h s i g h t i n g s , i t i s w o r t h n o t i n g t h a t t h e t i m e t o f i r s t s i g h t i n g a n d v a r i a t i o n i n t i m e s t o f i r s t s i g h t i n g w e r e l e s s f o r l a r g e r p o p u l a t i o n s ( T a b l e 3 . 1 ) . T h e l i k e l i h o o d o f s e e i n g a s q u i r r e l w i t h i n t h e f i r s t h o u r i n c r e a s e s w i t h t h e number o f r e s i d e n t s q u i r r e l s ( T a b l e 3 . 1 ; F i g u r e 3 . 3 ) . S i n c e S . s a t u r a t u s a r e p r o b a b l y l e s s d e t e c t a b l e when t h e p o p u l a t i o n i s a t l o w d e n s i t y , c a u t i o n s h o u l d b e t a k e n when d e t e r m i n i n g i t s a b s e n c e . N e v e r t h e l e s s , t h e p r o b a b i l i t y o f s i g h t i n g S . s a t u r a t u s (78% w i t h i n 60 m i n u t e s ; F i g . 3 . 3 ) g i v e s some c r e d e n c e t o t h e c o n c l u s i o n o f S . s a t u r a t u s ' s a b s e n c e o n v i s i t s o f a n h o u r ' s d u r a t i o n . W i t h a more s o c i a l a n d v o c a l s p e c i e s , d e t e r m i n i n g a b s e n c e may b e more c o n c l u s i v e . A t t e m p t s t o e l i c i t r e s p o n s e b y b l o w i n g a h i g h p i t c h e d w h i s t l e s o m e t i m e s a l e r t e d S . s a t u r a t u s t o i t s u p r i g h t p o s t u r e , b u t v o c a l r e s p o n s e s w e r e o n l y h e a r d f r o m O c h o t o n a p r i n c e p s , M a r m o t a s p p . , a n d S p e r m o p h i l u s c o l u m b i a n u s ( p e r s . o b s e r v . ) . W h i s t l i n g h a s p o t e n t i a l , f o r s u r v e y s o f o t h e r a n i m a l s . T r a p p i n g was a l a b o u r i n t e n s i v e m e t h o d f o r d e t e c t i n g S . s a t u r a t u s a t new l o c a t i o n s . I n f a c t , no new l o c a t i o n s o f i n h a b i t a t i o n w e r e f o u n d by t r a p p i n g . C a p t u r e r a t e s m i g h t h a v e i m p r o v e d i f l o c a t i o n s w e r e p r e b a i t e d ( R i t c h i e a n d S u l l i v a n 1 9 8 9 ) , b u t t h i s i s more t i m e c o n s u m i n g a n d i n e f f e c t i v e when many l o c a t i o n s r e q u i r e s a m p l i n g . T r a p a b i l i t y s u f f e r s f r o m t h e same f a c t o r s a s d o e s o b s e r v a b i l i t y , e x c e p t f o r v i s u a l o b s t r u c t i o n . T h e a d v a n t a g e o f o v e r c o m i n g v i s u a l o b s t r u c t i o n a n d t h e a b i l i t y t o d i s t i n g u i s h i n d i v i d u a l s by m a r k i n g t h e m , make t r a p p i n g a more r e l i a b l e i n d i c a t o r o f r e l a t i v e d e n s i t i e s b e t w e e n h a b i t a t s t h a n do w a l k i n g / o b s e r v a t i o n t r a n s e c t s . I f t i m e a n d l a b o u r a r e a v a i l a b l e , c h a n g e s i n p o p u l a t i o n s i z e , d e m o g r a p h i c c o m p o s i t i o n , a n d h a b i t a t u s e c a n be d e t e r m i n e d f r o m e x t e n s i v e t r a p p i n g r e g i m e s a n d c o l l e c t i o n o f f e c a l p e l l e t s f o r f o o d a n a l y s i s . F o r f u r t h e r d i s c u s s i o n o n s m a l l mammal t r a p p i n g , c o n s u l t C a l l (1986) . I n my s t u d y , t h e m o s t v a l u a b l e i n f o r m a t i o n o b t a i n e d f r o m t r a p p i n g d a t a ( F i g . 3 . 2 ) i s t h e p r e s e n c e o f y o u n g who s u r v i v e d t h r o u g h t h e i r f i r s t w i n t e r , t h e l i f e s t a g e w i t h h i g h e s t m o r t a l i t y f o r m o s t r o d e n t s ( C a u g h l e y 1 9 7 7 ) . A c o m b i n a t i o n o f d i f f e r e n t s u r v e y m e t h o d s i s p r o b a b l y t h e m o s t e f f e c t i v e means o f d e t e r m i n i n g t h e r a n g e a n d d i s t r i b u t i o n o f a s p e c i e s s u c h a s S . s a t u r a t u s . T h e c o n s t r a i n t s a n d p o t e n t i a l o f t h e t e c h n i q u e s may c o m p l e m e n t e a c h o t h e r s o t h a t s e v e r a l s o u r c e s o f i n f o r m a t i o n a r e e x p l o i t a b l e . N e v e r t h e l e s s , t h e p r o b l e m o f c o n c l u d i n g t h a t a r a r e a n i m a l i s n o t t h e r e s h o u l d b e r e c o g n i z e d ( H a i l a a n d J a r v i n e n 1983 ; M c A r d l e 1 9 9 0 ) . CHAPTER 4: G E N E R A L H A B I T A T COMPONENTS A S S O C I A T E D WITH SPERMOPHILUS SATURATUS 46 INTRODUCTION S p e r m o p h i l u s s a t u r a t u s a r e f o u n d i n c o n i f e r o u s f o r e s t s , o p e n meadows ( B r o a d b o o k s 1958; T r o m b u l a k 1 9 8 7 ) , k r u m h o l t z a n d r o c k d o m i n a t e d h a b i t a t ( R e i c h e l 1 9 8 6 ) . W i t h i n t h e s e h a b i t a t s , t h e y m u s t meet t h e i r b a s i c f o o d a n d s h e l t e r r e q u i r e m e n t s . T h e d i e t o f S . s a t u r a t u s i n c l u d e s A c h i l l e a , D r a b a . E p i l o b i u m , E r i q e r o n . E r y t h r o n i u m , F r a q a r i a , G a u l t h e r i a . L u p i n u s . O x y t r o p i s . P e n s t e m o n , P h a c e l i a . P h l o x . P i n u s . R h i z o p o q o n . S e n e c i o , S i l e n e , S o r b u s . T a r a x a c u m . T r i f o l i u m . V a c c i n i u m . V i c i a . b a r k , c o r m s , f u n g i a n d h y p o g e o u s f u n g i ( D a l q u e s t 1948 ; B r o a d b o o k s 1958 ; T r o m b u l a k 1988 ; C o r k a n d K e n a g y 1 9 8 9 ; p e r s . o b s e r v . ) . S . s a t u r a t u s c o n s t r u c t b u r r o w s f o r s h e l t e r a n d p r o t e c t i o n f r o m p r e d a t o r s . P e r c h e s f o r v i g i l a n c e ( M a c H u t c h o n a n d H a r e s t a d 1990) c o u l d a l s o b e e s s e n t i a l c o m p o n e n t s o f S . s a t u r a t u s h a b i t a t . S . s a t u r a t u s s p e n d m o s t o f t h e i r t i m e a b o v e - g r o u n d s i t t i n g ( T r o m b u l a k 1 9 8 7 ) ; some o f t h i s t i m e i s p r o b a b l y s p e n t i n v i g i l a n c e b e h a v i o u r . When c o v e r r e d u c e s v i s u a l d e t e c t i o n o f p r e y , p r e d a t o r s n e e d t o e x p e n d more e f f o r t s e a r c h i n g f o r g r o u n d d w e l l i n g p r e y ( J a n e s 1 9 8 5 ) . P o t e n t i a l p r e d a t o r s o f S . s a t u r a t u s i n c l u d e t h e g r e a t - h o r n e d o w l (Bubo v i r q i n i a n u s ) , goshawk ( A c c i p i t e r g e n t i l i s ) , , r e d - t a i l e d hawk ( B u t e o i a m a i c e n s i s ) , m e r l i n ( F a l c o c o l u m b a r i u s ) , l o n g - e a r e d o w l ( A s i o o t u s ) , c o y o t e ( C a n i s l a t r a n s ) , r e d f o x ( V u l p e s v u l p e s ) , l o n g t a i l e d w e a s e l ( M u s t e l a f r e n a t a ) , m a r t e n ( M a r t e s a m e r i c a n a ) a n d b o b c a t ( L y n x r u f u s ) ( T r o m b u l a k 1 9 8 8 ) . By e x a m i n i n g s p e c i e s ' h a b i t a t u s e , w i l d l i f e m a n a g e r s c a n b e t t e r a s s e s s a r e a s w h i c h t h e s p e c i e s c a n p o t e n t i a l l y i n h a b i t . I n t h i s c h a p t e r , I p r e s e n t a p r e l i m i n a r y d e s c r i p t i o n o f t h e h a b i t a t o f S . s a t u r a t u s b y m e a s u r i n g t h e p e r c e n t c o v e r o f p l a n t a n d u n v e g e t a t e d c a t e g o r i e s a t l o c a t i o n s i n h a b i t e d b y S . s a t u r a t u s . METHODS N i n e l o c a t i o n s i n h a b i t e d b y S . s a t u r a t u s w e r e c h o s e n . H a b i t a t s w e r e a s s e s s e d b e t w e e n 24 J u n e a n d 23 J u l y 1 9 9 0 , b e g i n n i n g w i t h t h e l o c a t i o n s o f e a r l i e s t p l a n t p h e n o l o g y . T h e n i n e l o c a t i o n s w e r e d i f f e r e n t i a t e d i n t o ' f i v e w h e r e f o o d was n o t s u p p l e m e n t e d ( C a t h e d r a l P a r k S l o p e , C a t h e d r a l L a k e s P r o v i n c i a l P a r k 4 9 ° 0 5 ' N 1 2 0 ° 1 2 ' W ; I r o n M o u n t a i n 5 0 ° 0 3 ' N 1 2 0 ° 4 6 ' W ; L a d y s l i p p e r L a k e , C a t h e d r a l L a k e s P r o v i n c i a l P a r k 4 9 ° 0 3 ' N 1 2 0 ° 1 1 ' W ; M o u n t O u t r a m 4 9 ° 1 7 ' N 1 2 1 ° 1 0 ' W ; T h y n n e M o u n t a i n 4 9 ° 4 2 ' N 1 2 0 ° 5 5 ' W ) a n d f o u r w h e r e f o o d was s u p p l e m e n t e d b y v i s i t o r h a n d o u t s a n d r e f u s e ( B r o o k m e r e Dump 4 9 ° 4 9 ' N 1 2 0 ° 5 2 ' W ; C o l d s p r i n g C a m p g r o u n d , M a n n i n g P r o v i n c i a l P a r k 4 9 ° 0 5 ' N 1 2 0 ° 4 8 ' W ; C a s c a d e L o o k o u t , M a n n i n g 48 P r o v i n c i a l P a r k 4 9 ° 0 5 ' N 1 2 0 ° 4 8 ' W ; Q u i n i s c o e C a m p g r o u n d , C a t h e d r a l L a k e s P r o v i n c i a l P a r k 4 9 ° 0 4 ' N 1 2 0 ° 1 2 ' W ) . A t e a c h l o c a t i o n , b e t w e e n two a n d f i v e a c t i v e b u r r o w s w e r e c h o s e n f o r t h e s a m p l i n g o f a l o c a t i o n ' s h a b i t a t c h a r a c t e r i s t i c s . T h e b u r r o w s c h o s e n w e r e r o u g h l y s p a c e d o u t a c r o s s e a c h l o c a t i o n . A t e a c h b u r r o w , a 5x5 m p l o t was c e n t r e d o v e r t h e b u r r o w o p e n i n g . F r o m t h e b u r r o w , a r a n d o m d i r e c t i o n was t a k e n t o e s t a b l i s h f o u r m o r e 5x5 m p l o t s , p l a c e d c o n c e n t r i c a l l y a t d i s t a n c e s i n c r e a s i n g i n 10 m i n c r e m e n t s ( F i g . 4 . 1 ) . T h e f u r t h e s t d i s t a n c e s a m p l e d f r o m a b u r r o w was t h u s 40 m a n d i t was a s s u m e d t h a t , o n a v e r a g e , S . s a t u r a t u s u s e d t h e h a b i t a t a t l e a s t w i t h i n a 45 m r a d i u s o f i t s b u r r o w . T h i s i s r e a s o n a b l e b e c a u s e S . s a t u r a t u s t r a v e l f u r t h e r t h a n t h a t d u r i n g t h e i r d a i l y a c t i v i t y ( p e r s . o b s e r v . ) . I n e a c h p l o t , t h e p e r c e n t c o v e r ( i f > 1%) o f t h e d i f f e r e n t p l a n t s p e c i e s a n d u n v e g e t a t e d c a t e g o r i e s ( e g . r o c k , g r a v e l ) w e r e v i s u a l l y e s t i m a t e d u s i n g t h e a v e r a g e v a l u e o f two o b s e r v e r s , e a c h a d i f f e r e n t p o i n t s o f v i e w . T h e s e d a t a w e r e r e c o r d e d a t t h r e e d i f f e r e n t h e i g h t c l a s s e s : 1) h e r b l a y e r (< 30 cm) ; 2) s h r u b l a y e r (> 30 cm a n d < 150 c m ) ; a n d 3) t r e e l a y e r (> 150 cm). P l a n t s p e c i e s w e r e c l a s s i f i e d b y t a x o n o m i c g e n e r a . T h e d a t a w e r e s u m m a r i z e d a n d s i m p l i f i e d b y g r o u p i n g p e r c e n t c o v e r o f p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s i n t o p e r c e i v e d f u n c t i o n a l c o m p o n e n t s : f o o d s p e c i e s , h y p o g e o u s f u n g i a s s o c i a t e s , b u r r o w s o r b u r r o w p o t e n t i a l , p e r c h p o t e n t i a l , p r o t e c t i v e c o v e r o r t o t a l p l a n t c o v e r . T a b l e 4 . 1 l i s t s t h e g e n e r a o r u n v e g e t a t e d c a t e g o r i e s i n c l u d e d i n e a c h f u n c t i o n a l c o m p o n e n t , e x c e p t t o t a l 49 3 F i g . 4 . 1 . A s c h e m a t i c d i a g r a m s h o w i n g t h e p l a c e m e n t o f a s e t o f f i v e p l o t s u s e d f o r s a m p l i n g h a b i t a t . T h e "+" i n d i c a t e s t h e l o c a t i o n o f a S . s a t u r a t u s b u r r o w . 50 T a b l e 4 . 1 . T h e g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s i n c l u d e d i n t h e f u n c t i o n a l c o m p o n e n t s o f S . s a t u r a t u s h a b i t a t . A l i s t o f c a t e g o r i e s t h a t w e r e m e a s u r e d a r e i n A p p e n d i x 7 . F u n c t i o n a l c o m p o n e n t C a t e g o r i e s i n c l u d e d R e l e v a n t L a y e r s F o o d s p e c i e s A s t e r a c e a e : C r u c i f e r a e E r i c a c e a e : F a b a c e a e : A c h i l l e a  E r i g e r o n  S e n e c i o  T a r a x a c u m  G a u i t h e r i a  D r a b a  V a c c i n i u m  L u p i n u s  O x y t r o p i s  T r i f o l i u m  V i c i a H y d r o p h y l l a c e a e : P h a c e l i a L i l i a c e a e : E r y t h r o n i u m  E p i l o b i u m  P i n u s  A a r o p y r o n  Bromus C a l a m a q r o s t i s F e s t u c a H o r d e u m P h l e u m P o a T r i s e t u m P o l e m o n i a c e a e : P h l o x R o s a c e a e : F r a g a r i a S c r o p h u l a r i a c a e : P e n s t e m o n H e r b , S h r u b O n a g r a c e a e P i n a c e a e : P o a c e a e : B u r r o w o r b u r r o w p o t e n t i a l G r a v e l / d e b r i s / s o i l H e r b H y p o g e o u s f u n g i a s s o c i a t e s C o n i f e r s : P i n a c e a e : A b i e s  P i c e a P i n u s S h r u b , T r e e P s e u d o t s u q a  T s u q a P e r c h e s B e d r o c k G r a v e l / d e b r i s / s o i l S t u m p s / d e a d f a l l L a r g e r o c k s (> 8 cm) S h r u b p l a n t c o v e r w h i c h i n c l u d e s a l l p l a n t g e n e r a i n e a c h l a y e r ( A p p e n d i x 7 ) . T o t a l p l a n t c o v e r i n t h e s h r u b a n d t r e e l a y e r s r e p r e s e n t e d p r o t e c t i v e c o v e r . T h e h y p o g e o u s f u n g i , R h i z o p o q o n s p . , common i n t h e d i e t o f S . s a t u r a t u s ( C o r k a n d K e n a g y 1 9 8 9 ) , i s a s s o c i a t e d w i t h c o n i f e r o u s f o r e s t s ( S m i t h a n d Z e l l e r 1 9 6 6 ) . C o n i f e r s w i t h g r e a t e r c o v e r w e r e a s s u m e d t o h a v e a more e x t e n s i v e r o o t s y s t e m a n d t h e r e f o r e r e p r e s e n t a t i v e o f a g r e a t e r a b u n d a n c e o f h y p o g e o u s f u n g i . T h e mean p e r c e n t c o v e r a n d s t a n d a r d d e v i a t i o n f o r e a c h g e n u s a n d u n v e g e t a t e d c a t e g o r y was c a l c u l a t e d f o r e a c h l o c a t i o n . T h e p e r c e n t c o v e r o f t h e p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s c o m p r i s i n g e a c h f u n c t i o n a l c o m p o n e n t was summed f o r e a c h p l o t a n d u s e d t o c a l c u l a t e t h e mean p e r c e n t c o v e r a n d s t a n d a r d d e v i a t i o n o f e a c h f u n c t i o n a l c o m p o n e n t a t e a c h l o c a t i o n . W i t h i n e a c h l o c a t i o n , e a c h s e t o f f i v e p l o t s was a s s o c i a t e d w i t h a p a r t i c u l a r b u r r o w a n d w o u l d b e t t e r r e p r e s e n t h a b i t a t u s e b y t h e s q u i r r e l s t h a n p l o t s p l a c e d r a n d o m l y . F o r t h e p u r p o s e o f t h i s t h e s i s , no s t a t i s t i c a l b u t o n l y d e s c r i p t i v e c o m p a r i s o n s w e r e i m p l e m e n t e d . R E S U L T S T h e mean p e r c e n t c o v e r a n d s t a n d a r d d e v i a t i o n s f o r a l l p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s i n e a c h l a y e r f o r e a c h l o c a t i o n a p p e a r i n A p p e n d i x 7 . T h e mean p e r c e n t c o v e r a n d T a b l e 4 . 2 . P e r c e n t c o v e r o f f u n c t i o n a l c o m p o n e n t s . V a l u e s h a v e b e e n r o u n d e d o f f t o t h e n e a r e s t i n t e g e r . Herb Layer Shrub Layer Tree Layer No. of Plant Food spp. Burrow Protective Food spp. Conifers Perches Protective Conifers plots cover potential cover cover Supplemented food sites Brookmere Dump 25 mean 43 11 54 29 11 1 1 19 18 sd 23 16 24 24 18 4 3 18 18 Coldspring Campground 10 mean 56 39 39 64 10 5 0 54 54 sd 20 18 19 21 14 11 0 27 27 Cascade Lookout 15 mean 40 30 32 35 3 8 7 28 27 sd 26 19 28 33 5 12 16 28 27 Quiniscoe Campground 25 mean 31 40 53 17 0 14 10 41 41 sd 21 14 25 17 1 15 14 26 26 Natural food sites Cathedral Park Slope 10 mean 63 40 16 33 0 2 22 27 27 sd 31 24 12 28 1 5 33 27 27 Iron Mountain 25 mean 60 19 20 33 8 1 5 28 27 sd 21 14 13 23 10 2 10 28 29 Ladyslipper Lake 10 mean 79 35 13 23 1 9 14 0 0 sd 9 6 8 21 2 14 7 0 0 Mount Outram 25 mean 71 18 12 4 1 2 0 9 8 sd 25 12 7 6 4 4 0 19 19 Thynne Mountain 25 mean 52 24 37 28 2 9 4 10 10 sd 21 17 24 23 4 11 9 17 17 s t a n d a r d d e v i a t i o n s o f t h e f u n c t i o n a l c o m p o n e n t s f o r e a c h l o c a t i o n a r e s u m m a r i z e d i n T a b l e 4 . 2 . D I S C U S S I O N W h i l e I d e s c r i b e d e a c h p e r c e i v e d f u n c t i o n a l c o m p o n e n t s e p a r a t e l y , t h e y may n o t b e i n d e p e n d e n t o f one a n o t h e r . Where one c o m p o n e n t i s s c a r c e a n o t h e r c o m p o n e n t may c o m p e n s a t e f o r i t . C o n i f e r f o r e s t h a b i t a t h a s l e s s p e r c e n t c o v e r o f h e r b a c e o u s f o o d t h a n o p e n a r e a s , b u t p r o v i d e s more h y p o g e o u s f u n g i a n d b a r k f o r f o o d t h a n o p e n a r e a s . Where k e y f o o d s p e c i e s a r e r a r e , g r e a t e r p r o t e c t i v e c o v e r may f a c i l i t a t e t r a v e l f o r f o o d o v e r a l a r g e r a r e a . C o n v e r s e l y , w h e r e k e y f o o d s p e c i e s a r e common, S . s a t u r a t u s may a g g r e g a t e d e s p i t e l i t t l e p r o t e c t i v e c o v e r ( e . g . s e e A n d e r s o n 1 9 8 5 ) . O f t h e l o c a t i o n s s a m p l e d , o n l y M o u n t O u t r a m h a d l i t t l e p r o t e c t i v e c o v e r . T h i s l a c k o f p r o t e c t i v e c o v e r s u g g e s t s t h a t e i t h e r more f o o d was a v a i l a b l e a t t h i s s i t e t h a n m e a s u r e d , few c o m p e t i t o r s f o r f o o d r e s o u r c e s w e r e p r e s e n t , o r my l i s t o f f o o d s p e c i e s was i n c o m p l e t e . S i m i l a r l y , r e f u s e a n d h a n d -o u t s a t f o o d s u p p l e m e n t e d l o c a t i o n s p r o v i d e d a s u p p l e m e n t a l a n d e a s i l y a c c e s s i b l e f o o d s o u r c e t h a t c o u l d o u t w e i g h t h e p r e d a t i o n r i s k r e s u l t i n g f r o m l a c k o f p r o t e c t i v e c o v e r . T h e l a c k o f c o n s i s t e n c y i n t h e c o m p o s i t i o n o f f u n c t i o n a l c o m p o n e n t s a t v a r i o u s l o c a t i o n s may r e f l e c t a g r e a t e r f l e x i b i l i t y i n t h e d i e t o f S . s a t u r a t u s . T h e h y p o g e o u s f u n g i a n d p l a n t g e n e r a i n g e s t e d b y S . s a t u r a t u s a l s o a p p e a r i n f o o d i t e m s l i s t e d f o r i t s s i b l i n g s p e c i e s , S . l a t e r a l i s ( T e v i s 1954 , M c K e e v e r 1964 , C a r l t o n 1 9 6 6 ) . S . l a t e r a l i s i s known t o h a v e a w i d e r a n g e o f f o o d s i n i t s d i e t ; t h e same i s p r o b a b l y t r u e f o r S . s a t u r a t u s , i t s s i b l i n g s p e c i e s . T r o m b u l a k (1987) f o u n d t h a t f o r e s t d w e l l i n g a n d meadow d w e l l i n g S . s a t u r a t u s h a d d i f f e r e n t d i e t s r e f l e c t i n g t h e a v a i l a b i l i t y o f d i f f e r e n t f o o d s . T o b e t t e r a s s e s s t h e f o o d a v a i l a b i l i t y a t a l o c a t i o n , more i n f o r m a t i o n on t h e v a r i e t y o f f o o d s p e c i e s r e q u i r e d t o s a t i s f y S . s a t u r a t u s o f d i f f e r e n t age a n d s e x g r o u p s o v e r d i f f e r e n t s e a s o n s i s n e e d e d . Some i n f o r m a t i o n on t h e n u t r i t i o n a l v a l u e o f h y p o g e o u s f u n g i a l r e a d y e x i s t s ( C o r k a n d K e n a g y 1 9 8 9 ) . I n t h i s h a b i t a t a s s e s s m e n t , t h e f u n c t i o n a l c o m p o n e n t s w e r e m e a s u r e d a n d c o m p a r e d among l o c a t i o n s w i t h o u t t a k i n g i n t o a c c o u n t t h e d i f f e r e n t d e n s i t i e s o f s q u i r r e l s i n e a c h l o c a t i o n . I f t h e number o f s q u i r r e l s u t i l i z i n g a l o c a t i o n i s k n o w n , i t may be e a s i e r t o d e t e r m i n e w h e t h e r a p a r t i c u l a r f u n c t i o n a l c o m p o n e n t i s a l i m i t i n g f a c t o r ( e . g . s e e C a r l 1971 a n d Y o u n g 1 9 9 0 , r e : b u r r o w i n g s i t e s l i m i t i n g p o p u l a t i o n s i z e s o f some S p e r m o p h i l u s s p p . ) . F u r t h e r s t u d i e s s h o u l d a l s o i n c l u d e c o m p a r i s o n s o f o c c u p i e d l o c a t i o n s w i t h a p p a r e n t l y u n o c c u p i e d a d j a c e n t l o c a t i o n s . T h e d a i l y a c t i v i t y p a t t e r n o f i n d i v i d u a l S . s a t u r a t u s a t o c c u p i e d l o c a t i o n s s h o u l d be f o l l o w e d t o d e t e r m i n e w h i c h h a b i t a t c h a r a c t e r i s t i c s i n t h e s e l o c a t i o n s p e r m i t S . s a t u r a t u s i n h a b i t a t i o n a n d w h i c h o f t h e s e h a b i t a t c h a r a c t e r i s t i c s a r e a b s e n t i n a d j a c e n t u n o c c u p i e d s i t e s . D e s c r i p t i o n s o f S . s a t u r a t u s h a b i t a t t h a t r e p r e s e n t S . s a t u r a t u s a c r o s s i t s r a n g e s h o u l d b e o b t a i n e d b y s u r v e y i n g h a b i t a t a c r o s s i t s r a n g e . Some v e r t e b r a t e s p e c i e s ( e g . b i r d s ) may b e l e s s s e l e c t i v e o f v e g e t a t i o n a l c h a r a c t e r i s t i c s a t t h e n o r t h e r n e d g e a s o p p o s e d t o t h e s o u t h e r n p o r t i o n o f t h e i r r a n g e ( E e v a e t a l . 1 9 8 9 ) . H a b i t a t d e s c r i p t i o n s s h o u l d i n c l u d e t h e r o l e o f c o m p e t i t o r s f o r t h e same r e s o u r c e s a n d p r e d a t o r r e l i a n c e o n S . s a t u r a t u s a s p r e y s i n c e t h e y c o u l d c h a n g e t h e s u i t a b i l i t y o f h a b i t a t f o r t h e g r o u n d s q u i r r e l . A l t h o u g h m e a s u r e m e n t s o f h a b i t a t c h a r a c t e r i s t i c s i n t h i s s t u d y p r o v i d e a g e n e r a l d e s c r i p t i o n o f S . s a t u r a t u s h a b i t a t i n B r i t i s h C o l u m b i a , t h e r e s e a r c h s u g g e s t e d a b o v e may f a c i l i t a t e more d e f i n i t i v e a s s e s s m e n t s o f p o t e n t i a l h a b i t a t t h a n p r e s e n t l y p o s s i b l e . 56 CHAPTER 5: G E N E R A L D I S C U S S I O N AND RECOMMENDATIONS F o r t h e C o m m i t t e e o n t h e S t a t u s o f E n d a n g e r e d W i l d l i f e i n C a n a d a ( C O S E W I C ) , t h e t e c h n i c a l d e f i n i t i o n o f s p e c i e s i n c l u d e s "any s p e c i e s , s u b s p e c i e s , o r g e o g r a p h i c a l l y s e p a r a t e d p o p u l a t i o n " . S . s a t u r a t u s f i t s t h i s d e f i n i t i o n b e c a u s e i t s g e n e t i c c o m p o s i t i o n a n d m o r p h o l o g y a r e s u f f i c i e n t l y d i f f e r e n t f r o m i t s c l o s e s t r e l a t i v e , S . l a t e r a l i s ( C h a p t e r 2) , a n d i t s g e o g r a p h i c r a n g e i s d i s j u n c t f r o m S . l a t e r a l i s . t h e r e b y p r e v e n t i n g i n t e r b r e e d i n g b e t w e e n t h e two s p e c i e s . COSEWIC d e f i n e s a " v u l n e r a b l e s p e c i e s " a s "any i n d i g e n o u s s p e c i e s o f f a u n a o r f l o r a t h a t i s p a r t i c u l a r l y a t r i s k b e c a u s e o f l o w o r d e c l i n i n g n u m b e r s , o c c u r r e n c e a t t h e f r i n g e o f i t s r a n g e o r i n r e s t r i c t e d a r e a s , o r f o r some o t h e r r e a s o n , b u t i s n o t a t h r e a t e n e d s p e c i e s " (Munro 1 9 9 0 ) . C l e a r l y , b y t h i s d e f i n i t i o n , S . s a t u r a t u s i n B r i t i s h C o l u m b i a f i t s i n t o t h e c a t e g o r y o f " v u l n e r a b l e s p e c i e s " b e c a u s e t h e y a r e a t t h e n o r t h e r n " f r i n g e " o f t h e i r r a n g e . A l t h o u g h t h e r a n g e o f S . s a t u r a t u s i s f u r t h e r w e s t a n d n o r t h t h a n p r e v i o u s l y c i t e d , my d a t a s u g g e s t t h a t t h i s s q u i r r e l i s n o t common t h r o u g h o u t i t s r a n g e i n B r i t i s h C o l u m b i a ( C h a p t e r 3 ) . S . s a t u r a t u s w e r e n o t f o u n d a s f a r e a s t a s t h e y w e r e p r e v i o u s l y r e p o r t e d , p o s s i b l y d u e t o h a b i t a t a l t e r a t i o n a n d s u b s e q u e n t d i s p l a c e m e n t b y c o m p e t i n g s p e c i e s ( C h a p t e r 3) . T h e l o w g e n e t i c v a r i a t i o n f o u n d i n S . s a t u r a t u s may b e a r e s u l t o f f o u n d e r e f f e c t s o r s t r o n g s e l e c t i o n i n p e r i p h e r a l 57 a r e a s , s i n c e m o s t o f t h e S . s a t u r a t u s s a m p l e d w e r e f r o m t h e n o r t h e r n e x t e n t o f i t s r a n g e ( C h a p t e r 2) . T h i s l o w v a r i a t i o n r e l a t i v e t o S . l a t e r a l i s may r e s t r i c t S . s a t u r a t u s t o a s m a l l e r r a n g e t h a n S . l a t e r a l i s . P e r h a p s t h e l o w v a r i a t i o n e x p l a i n s why S . s a t u r a t u s d o e s n o t r a n g e a s f a r n o r t h a s d o e s S . l a t e r a l i s ( F i g . 1 . 1 ) . When S . l a t e r a l i s f r o m d i f f e r e n t a l t i t u d e s w e r e e x p o s e d t o t h e same l a b c o n d i t i o n s , t h e y s t i l l h a d p h y s i o l o g i c a l d i f f e r e n c e s i n t h e i r c i r c a n n u a l c y c l e ( B l a k e 1 9 7 2 ) , s u g g e s t i n g t h a t t h e r e i s g e o g r a p h i c v a r i a t i o n i n g e n o t y p e . B l a k e ' s (1972) r e s u l t s a r e s u p p o r t e d b y t h e r e l a t i v e l y h i g h g e n e t i c v a r i a t i o n f o u n d i n S . l a t e r a l i s i n t h i s s t u d y a n d c o n t r a s t s w i t h t h e r e l a t i v e l y l o w g e n e t i c v a r i a t i o n i n S . s a t u r a t u s . ( P o l y m o r p h i s m f o r e n z y m e s G P I , LDH a n d E S T w e r e f o u n d i n S . s a t u r a t u s f r o m L a k e W e n a t c h e e , W a s h i n g t o n S t a t e ; K e n a g y u n p u b l . d a t a . ) D e s p i t e i t s l o w g e n e t i c v a r i a t i o n , S . s a t u r a t u s c a n s t i l l be f o u n d i n a v a r i e t y o f d i f f e r e n t h a b i t a t s ( C h a p t e r 4) . A c c o r d i n g t o R e i c h e l ( 1 9 8 6 ) , i n t h e a l p i n e , b o t h S . s a t u r a t u s a n d S . l a t e r a l i s u s e r o c k d o m i n a t e d a n d k r u m h o l t z h a b i t a t , b u t o n l y S . l a t e r a l i s u s e f e l l f i e l d a n d g r a s s l a n d h a b i t a t . H o w e v e r , i n my s t u d y a r e a , S . s a t u r a t u s w e r e a l s o p r e s e n t i n f e l l f i e l d h a b i t a t . A s w e l l , t h e v a r i e t y o f f o o d s i n g e s t e d b y S . s a t u r a t u s . i n c l u d i n g h e r b s , f u n g i a n d v i s i t o r h a n d o u t s , d e m o n s t r a t e s f l e x i b i l i t y i n t h e s p e c i e s ' d i e t ( C h a p t e r 4 ) . T h e n e e d t o e n h a n c e h a b i t a t s f o r S . s a t u r a t u s c a n n o t b e d e t e r m i n e d a t p r e s e n t , b u t more r e s e a r c h o n t h e i r b i o l o g y i s r ecommended ( T a b l e 5 . 1 ) s o t h a t p o t e n t i a l t h r e a t s t o t h e i r e x i s t e n c e may b e a v o i d e d . S . s a t u r a t u s d o e s n o t h a v e a h i g h 58 r i s k o f i m m e d i a t e e x t i n c t i o n b e c a u s e o f i t s a b i l i t y t o t h r i v e i n a v a r i e t y o f d i f f e r e n t h a b i t a t s a n d t h e a d e q u a t e s i z e o f i t s p r e s e n t r a n g e i n s o u t h w e s t e r n B r i t i s h C o l u m b i a a n d W a s h i n g t o n S t a t e . H o w e v e r , d u e t o i t s l o w d e n s i t y , l o w g e n e t i c v a r i a t i o n a n d p o s s i b l e d i s p l a c e m e n t b y i n c o m p a t i b l e s p e c i e s whose d i s p e r s a l i s f a c i l i t a t e d b y h a b i t a t c h a n g e s , S . s a t u r a t u s s h o u l d b e g i v e n t h e c o n s e r v a t i o n s t a t u s o f " v u l n e r a b l e s p e c i e s " . 59 T a b l e 5 . 1 . R e c o m m e n d a t i o n s f o r t h e c o n s e r v a t i o n o f S p e r m o p h i l u s  s a t u r a t u s . 1) R e s e a r c h p r i o r i t i e s i d e n t i f i e d f r o m p r e s e n t s t u d y a . Does h a b i t a t a l t e r a t i o n p r o m o t e e x p l o i t a t i v e c o m p e t i t i o n w h e r e b y S . s a t u r a t u s i s d i s p l a c e d b y S . c o l u m b i a n u s ? T h i s c a n b e t e s t e d by m e a s u r i n g h a b i t a t u s e b y S . s a t u r a t u s a n d S . c o l u m b i a n u s l i v i n g p r o x i m a t e t o e a c h o t h e r , t h e n m e a s u r i n g h a b i t a t u s e b y S . s a t u r a t u s a f t e r S . c o l u m b i a n u s h a s b e e n r e m o v e d . I f S . s a t u r a t u s e x p a n d s i t s h a b i t a t u s e i n t h e a b s e n c e o f S . c o l u m b i a n u s . S . c o l u m b i a n u s was l i k e l y d i s p l a c i n g S . s a t u r a t u s . b . M e a s u r e t h e g e n e t i c v a r i a t i o n o f S . s a t u r a t u s i n W a s h i n g t o n S t a t e a n d c o m p a r e i t w i t h t h a t o f t h e S . s a t u r a t u s i n B r i t i s h C o l u m b i a . I s t h e l o w g e n e t i c v a r i a t i o n o f S . s a t u r a t u s i n B . C . d u e t o f o u n d e r e f f e c t s o r some a l t e r n a t i v e f a c t o r ? c . D e t e r m i n e t h e d i s p e r s a l a n d c o l o n i z i n g a b i l i t y o f S . s a t u r a t u s . d . D e t e r m i n e t h e f o o d a n d h a b i t a t r e q u i r e m e n t s o f S . s a t u r a t u s a c r o s s i t s r a n g e . See C h a p t e r 4 D i s c u s s i o n . How w e l l do p a r k s a n d e c o l o g i c a l r e s e r v e s f u n c t i o n a s w i l d l i f e r e s e r v e s f o r S . s a t u r a t u s ? 2) G e n e r a l r e s e a r c h r e q u i r e d a . P o p u l a t i o n s o f S . s a t u r a t u s n e e d t o b e m o n i t o r e d p e r i o d i c a l l y . P o p u l a t i o n t r e n d s a r e u n k n o w n . I n f o r m a t i o n i s l a c k i n g on t h e age s t r u c t u r e , s e x r a t i o , b i r t h r a t e s a n d age s p e c i f i c m o r t a l i t y o f S . s a t u r a t u s . b . U n d e r s t a n d i n g t h e i n t e r a c t i o n s o f s p e c i e s , p r o v i d e s i n s i g h t oh t h e s t a b i l i t y a n d r e s i l i e n c e o f a c o m m u n i t y ( H o l l i n g 1 9 7 3 ; Pimm 1 9 8 6 ) . T h e r e l a t i o n s h i p o f S . s a t u r a t u s t o o t h e r s p e c i e s i n t h e c o m m u n i t y m u s t be e x a m i n e d . 60 L I T E R A T U R E C I T E D A n d e r s o n , P . K . 1 9 8 5 . F o r a g i n g r a n g e i n m i c e a n d v o l e s : t h e r o l e o f r i s k . C a n . J . Z o o l . 64: 2 6 4 5 - 2 6 5 3 . A n d r u s i a k , L . A . a n d A . S . H a r e s t a d . 1 9 8 7 . F e e d i n g b e h a v i o u r a n d d i s t a n c e f r o m b u r r o w s o f C o l u m b i a n g r o u n d s q u i r r e l s . C a n . J . Z o o l . 6 7 : 3 8 1 - 3 8 4 . A u f f r a y , J . C . , E . T c h e r n o v , F . Bonhomme, G . H e t h , S . S i m s o n a n d E . Newo. 199 0 . P r e s e n c e a n d e c o l o g i c a l d i s t r i b u t i o n o f Mus  s p r e t o i d e s a n d Mus m u s c u l u s d o m e s t i c u s i n I s r a e l : C i r c u m -M e d i t e r r a n e a n v i c a r i a n c e i n t h e G e n u s M u s . Z . S a e u g e t i e r k De 5 5 ( 1 ) : 1 - 1 0 . A v i s e , J . 1 9 7 6 . G e n e t i c d i f f e r e n t i a t i o n d u r i n g s p e c i a t i o n . I n M o l e c u l a r E v o l u t i o n . E d i t e d b y F . J . A y a l a . S i n a u e r A s s o c . I n c . , S u n d e r l a n d , M a s s a c h u s e t t s , p p . 1 0 6 - 1 2 2 . A y a l a , F . J . a n d J . A . K i g e r . 1984 . M o d e r n G e n e t i c s . 2 n d e d . B e n j a m i n / C u m m i n g s P u b l i s h i n g C o . I n c . O n t a r i o . B a i l e y J . A . 1 9 8 6 . . W e a t h e r a n d c l i m a t e . I n I n v e n t o r y a n d m o n i t o r i n g o f w i l d l i f e h a b i t a t . E d i t e d b y A . Y . C o o p e r i d e r , J . B o y d a n d H . R . S t u a r t . U . S . D . I . B u r . L a n d M a n a g e . S e r v i c e C e n t e r , C o l o r a d o , p p . 7 1 1 - 1 2 5 . B a n f i e l d A . W. F . 1974 . T h e Mammals o f C a n a d a . U n i v e r s i t y o f T o r o n t o P r e s s , T o r o n t o . B a r r o w c l o u g h , G . F . a n d R . J . G u t i e r r e z . 1 9 9 0 . G e n e t i c v a r i a t i o n a n d d i f f e r e n t i a t i o n i n t h e s p o t t e d o w l ( S t r i x o c c i d e n t a l i s m . A u k 1 0 7 : 7 3 7 - 7 4 4 . B a r t o n , N . H . a n d B . C h a r l e s w o r t h . 1984 . G e n e t i c r e v o l u t i o n s , f o u n d e r e f f e c t s , a n d s p e c i a t i o n . A n n . R e v . E c o l . S y s t . 15 : 1 3 3 - 1 6 4 . B e g o n , M . a n d M . M o r t i m e r . 1 9 8 1 . P o p u l a t i o n e c o l o g y : A u n i f i e d s t u d y o f a n i m a l s a n d p l a n t s . S i n a u e r A s s o c . I n c . , M a s s a c h u s e t t s . B e s t g e n , K . R . a n d P . L . P r o p s t . 1 9 8 9 . D i s t r i b u t i o n , s t a t u s a n d n o t e s o n t h e e c o l o g y o f G i l a r o b u s t a ( C y p r i n i d a e ) . S o u t h w e s t . N a t . 3 4 ( 3 ) : 4 0 2 - 4 1 2 . B e t t s , B . J . 1 9 9 0 . G e o g r a p h i c d i s t r i b u t i o n a n d h a b i t a t p r e f e r e n c e s o f W a s h i n g t o n g r o u n d s q u i r r e l s ( S p e r m o p h i l u s  w a s h i n q t o n i ) . N o r t h w e s t . N a t . 7 1 ( 2 ) : 2 7 - 3 7 . 61 B i c k m a n , J . W. 1 9 8 3 . S i b l i n g s p e c i e s . I n G e n e t i c s a n d c o n s e r v a t i o n . E d i t e d b y C . M . S c h o n e w a l d - C o x , S . M . C h a m b e r s , B . M a c B r y d e , W. L . T h o m a s . B e n j a m i n / Cummings P u b l i s h i n g C o . I n c . , O n t a r i o , p p . 9 6 - 1 0 6 . B l a k e , B . H . 1 9 7 2 . T h e a n n u a l c y c l e a n d f a t s t o r a g e i n two p o p u l a t i o n s o f g o l d e n m a n t l e d g r o u n d s q u i r r e l s . J . Mammal . 5 3 : 1 5 7 - 1 6 7 . B r o a d b o o k s , H . E . 1 9 5 8 . L i f e h i s t o r y a n d e c o l o g y o f t h e c h i p m u n k , E u t a m i a s amoenus , i n e a s t e r n W a s h i n g t o n . M i s c . P u b l . M u s . Z o o l . , U n i v . M i c h i g a n 103: 1 - 4 2 . B r o n s o n , M . T . 1 9 7 9 . A l t i t u d i n a l v a r i a t i o n i n t h e l i f e h i s t o r y o f t h e g o l d e n m a n t l e d g r o u n d s q u i r r e l S p e r m o p h i l u s l a t e r a l i s . E c o l o g y 60 : 2 7 2 - 2 7 9 . Brown J . H . 1 9 7 1 . Mammals on m o u n t a i n t o p s : n o n e q u i l i b r i u m i n s u l a r b i o g e o g r a p h y . A m e r . N a t . 105: 4 6 7 - 4 7 8 . B u r t , W. H . a n d R . P . G r o s s e n h e i d e r . 197 6 . 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A p r e l i m i n a r y a c c o u n t o f t h e N o r t h A m e r i c a n s p e c i e s o f R h i z o p o q o n . Mem. New Y o r k B o t . G a r d e n s 1 4 ( 2 ) : 1 - 1 7 8 . S o u l e , M . E . 1 9 8 0 . T h r e s h o l d s f o r s u r v i v a l : m a i n t a i n i n g f i t n e s s a n d e v o l u t i o n a r y p o t e n t i a l . I n C o n s e r v a t i o n B i o l o g y : a n E v o l u t i o n a r y - E c o l o g i c a l P e r s p e c t i v e . E d i t e d b y M . E . S o u l e a n d B . W i l c o x . S i n a u e r A s s o c i a t e s I n c . , M a s s a c h u s e t t s , p p . 1 5 1 - 1 6 9 . 67 S t a r k , H . E . , B . W. H u d s o n , B . P i t t m a n . 1 9 6 6 . P l a g u e e p i d e m i o l o g y . P u b l i c H e a l t h S e r v i c e . G e o r g i a . T o m i c h , P . Q . 1 9 8 2 . G r o u n d s q u i r r e l s . I n W i l d mammals o f N o r t h / A m e r i c a . E d i t e d b y J . A . Chapman a n d G . A . F e l d h a m e r . J o h n H o p k i n s U n i v . P r e s s , B a l t i m o r e . 1 1 4 7 p p . T e v i s J r . , L . 1 9 5 3 . S t o m a c h c o n t e n t s o f c h i p m u n k s a n d m a n t l e d s q u i r r e l s i n n o r t h e a s t e r n C a l i f o r n i a . J . Mammal . 34: 3 1 6 -3 2 4 . T r o m b u l a k , S . C . 1 9 8 7 . L i f e h i s t o r y o f t h e C a s c a d e m a n t l e d g r o u n d s q u i r r e l ( S p e r m o p h i l u s s a t u r a t u s ) . J . Mammal . 68 : 5 4 4 - 5 5 4 . T r o m b u l a k , S . C . 1 9 8 8 . S p e r m o p h i l u s s a t u r a t u s . M a m m a l i a n S p e c i e s . N o . 322 pp 1 - 4 . V a n D e V e e r , D . a n d C . P i e r c e , e d s . 1 9 8 6 . P e o p l e , p e n g u i n s , a n d p l a s t i c t r e e s : b a s i c i s s u e s i n e n v i r o n m e n t a l e t h i c s . W a d w o r t h P u b l i s h i n g C o . , C a l i f o r n i a . W a r l s b e r g , G . E . 1 9 9 0 . C o n v e r g e n c e o f s o l a r h e a t g a i n i n two s q u i r r e l s p e c i e s w i t h c o n t r a s t i n g c o a t c o l o r s . P h y s i o l . Z o o l . 6 3 : 1 0 2 5 - 1 0 4 2 . Y o u n g P . J . 1 9 9 0 . S t r u c t u r e , l o c a t i o n a n d a v a i l a b i l i t y o f h i b e r n a c u l a o f C o l u m b i a n g r o u n d s q u i r r e l s ( S p e r m o p h i l u s c o l u m b i a n u s ) . Am. M i d i . N a t . 123: 3 5 7 - 3 6 4 . 68 A p p e n d i x 1. Museum s p e c i m e n s e x a m i n e d a n d m e a s u r e d f o r c r a n i a l c h a r a c t e r i s t i c s . Museums: C o n n e r Museum, W a s h i n g t o n S t a t e U n i v e r s i t y (WSU); T h e B u r k e Museum, U n i v e r s i t y o f W a s h i n g t o n (UWA) ; S l a t e r Museum, U n i v e r s i t y o f P u g e t S o u n d (UPS),; Cowan V e r t e b r a t e Museum, U n i v e r s i t y o f B r i t i s h C o l u m b i a ( U B C ) ; N a t i o n a l Museums o f C a n a d a ( N M C ) ; R o y a l B r i t i s h C o l u m b i a Museum ( R B C M ) . " * " d e n o t e s s k u l l s n o t m e a s u r e d , b u t u s e d i n r a n g e map. Species Sex Location Museum ID# Date Ss f Yakima Co., WA WSU 6474 1964 Ss f Skamania Co., WA WSU 83146 -Ss f Chelan Co., WA WSU 83144 1980 Ss f Chelan Co., WA WSU 83143 1980 Ss f Okanogan Co., WA WSU 76309 -Ss f Okanogan Co., WA WSU 53168 -Ss f Okanogan Co., WA WSU 76310 -• Ss f Lewis Co., WA WSU 77501 - . Ss f Pierce Co., WA WSU 119 -Ss m Yakima Co., WA WSU 6485 1964 Ss m Chelan Co., WA WSU 81379 1980 Ss m Okanogan Co., WA WSU 8394 -Ss m Kittitas Co., WA WSU 48176 -Ss m Okanogan Co., WA WSU 8046 1979 Ss m Stevens Co., WA WSU 82140 1981 Ss m Okanogan Co., WA WSU 82135 1943 Ss m Yakima Co., WA WSU 117 ' 1917 Ss m Okanogan Co., WA WSU 47162 1947 Ss m Pierce Co., WA WSU 121 -Ss m Chelan Co., WA WSU 83145 1980 Ss m Chelan Co., WA WSU 81381 1980 Ss m Yakima Co., WA WSU 6294 -Ss f Kittitas Co., WA UWA 15004 1936 Ss f Lewis Co., WA UWA 35506 1987 Ss f Skamania Co., WA UWA 3400 1963 Ss f Chelan Co., WA UWA 32988 1981 Ss f Yakima Co, WA UWA 1522 1966 Ss f Yakima Co., WA UWA 495 1941 Ss f Yakima Co., WA UWA 35507 1987 Ss f Yakima Co, WA UWA 1027 1965 Ss m Skamania Co., WA UWA 3401 1963 Ss m Yakima Co, WA UWA 1020 1965 Ss m Kittitas Co., WA UWA 16457 1958 Ss m Chelan Co., WA UWA 33089 1981 Ss m Skamania Co., WA UWA 4412 1966 Ss m Pierce Co.,WA UWA 31137 1976 Ss m Yakima Co, WA UWA 3877 , 1965 Ss m Pierce Co., WA UWA 31558 1976 Ss m Yakima Co., WA UWA 2575 1961 Ss m Skamania Co., WA UWA 3878 1965 A p p e n d i x 1. C o n t i n u e d . Species Sex Location Museum ID# Date Ss m Chelan Co., WA UWA 16394 1958 Ss m Skamania Co., WA UWA 3851 1965 Ss m Skamania Co., WA UWA 1037 1965 Ss m Lewis Co., WA UWA 35505 1987 Ss m Chelan Co., WA UWA 33008 . 1981 Ss f Yakima Co., WA UPS 9886 1962 Ss f Pierce Co., WA UPS 19944 1974 Ss f Pierce Co., WA UPS 26422 1979 Ss f Kittitas Co, WA UPS 2140 1949 Ss f Pierce Co., WA UPS 19952 1974 Ss f Pierce Co., WA UPS 19946 1974 Ss f Pierce Co., WA UPS 19942 1974 Ss f Pierce Co., WA UPS 26421 1979 Ss f Pierce Co., WA UPS 19951 1974 Ss f Chelan Co., WA UPS 1172 1947 Ss f Chelan Co., WA UPS 1173 1947 Ss f Kittitas Co., WA UPS 7966 1948 Ss f Pierce Co., WA UPS 19948 1974 Ss f Pierce Co., WA UPS 19943 1974 Ss m Kittitas Co, WA UPS 4037 1953 Ss m Chelan Co., WA UPS 1524 1948 Ss m Klickitat Co., WA UPS 1991 1949 Ss m Pierce Co., WA UPS 557 1934 Ss m Chelan Co., WA UPS 1171 1947 Ss m Kittitas Co., WA UPS 7967 1948 Ss m Pierce Co., WA UPS 19949 1974 Ss m Pierce Co., WA UPS 19953 1974 Ss f Manning Prov. Park, BC , UBC 2968 1949 Ss f Princeton, BC • UBC 3685 1943. Ss f Manning Prov. Park, BC UBC 2969 1949 Ss f Princeton, BC UBC 3683 1943 Ss f 17 miles up Ashnola R., BC UBC 5629 1950 Ss f Kittitas Co., WA UBC 667 1937 Ss f Ashnola R., BC UBC 3556 1951 Ss m Ashnola Forks, BC UBC 3554 1951 Ss m Ashnola Forks, BC UBC 3555 1951 Ss m Gibson Pass, Manning Prov. Park, BC UBC 4298 1952 Ss m Princeton, BC UBC 3684 1943 Ss m Blue R.(Northnortheast of Princeton), BC UBC 5630 1947 Ss m Manning Prov. Park, BC UBC 2966 1949 70 A p p e n d i x 1. C o n t i n u e d . Species Sex Location Museum ID# Date Ss m 12 miles East of Princeton, BC UBC 5631 1952 Ss f Second Summit, Skagit Ft., BC NMC 1207 1905 * Ss f Similkameen R., BC NMC 8966 1928 * Ss f Second Summit, Skagit Ft., BC NMC 1198 1905 * Ss f Ashnola Ck, BC NMC 28847 1950 * Ss f Similkameen R., BC NMC 8967 1928 * Ss f Hope-Princeton Summit Pass, BC NMC 7858 1927 * Ss f Hope-Princeton Summit Pass, BC NMC 7856 1927 * Ss f Stirling Ck (Northwest of Hedley), BC NMC 8884 1928 * Ss f near Princeton, BC NMC 28848 1950 * Ss f Stirling Ck (Northwest of Hedley), BC NMC 8883 1928 * Ss f Stirling Ck (Northwest of Hedley), BC NMC 8908 1928 * Ss f Second Summit, Skagit R., BC NMC 1199 1905 * Ss f Stirling Ck (Northwest of Hedley), BC NMC 8857 1928 * Ss m Second Summit, Skagit R., BC NMC 1205 1905 * Ss m Whipsaw Ck, BC NMC 1163 1905 * Ss m Skagit, BC NMC 1165 1905 X Ss m Second Summit, Skagit R., BC NMC 1202 1905 * Ss m Hope-Princeton Summit Pass, BC NMC 7857 1927 * Ss m Stirling Ck (Northwest of Hedley), BC NMC 8889 1928 * Ss - Manning Prov. Park, BC RBCM 5175 1945 Ss - Mt Gordon, Manning Prov. Pk, BC RBCM 5485 1949 Ss - Tulameen, BC RBCM 225 1917 Ss - Princeton, BC RBCM 4956 1939 Ss - Princeton, BC RBCM 4955 1939 Ss - Manning Prov. Park, BC RBCM 5174 1945 Sit f Okanogan Co., WA WSU 82282 -Sit f Idaho Co., ID WSU 78445 -Sit f Pend Oreille Co., WA WSU 47144 1947 Sit m Pend Oreille Co., WA WSU 83151 1981 Sit m Pend Oreille Co., WA WSU 81392 1980 Sit m Ravalli Co., WA WSU 83517 1983 Sit m Pend Oreille Co., WA WSU 82281 1981 Sit f Missoula Co., MT UWA 4245 1975 Sit f Missoula Co., MT UWA 4072 1974 Sit f Missoula Co., MT UWA 4058 1974 Sit f Missoula Co., MT UWA 4077 1974 Sit m Missoula Co., MT UWA 4061 1974 Sit m Missoula Co., MT UWA 4073 1974 Sit m Missoula Co., MT UWA 4084 1974 A p p e n d i x 1. C o n t i n u e d . Species Sex Location Museum ID# Date Sit m Missoula Co., MT UWA 4060 1974 Sit m Missoula Co., MT UWA 4244 1975 Sit m Missoula Co., MT UWA 4083 1974 Sit m Missoula Co., MT UWA 5993 1974 Sit f Pend Oreille Co., WA UPS 4193 1953 Sit m Pend Oreille Co., WA UPS 3751 1952 Sit m Bonner Co., ID UPS 9907 1964 Sit m Sanders Co., MT UPS 9374 1962 Sit m 22 miles West of Edgewood, BC UPS 5652 1955 Sit f Medicine L, AB UBC 1570 1945 Sit f 20 miles West of Banff, AB UBC 748 1943 Sit f Spray R., Banff, AB UBC 750 1943 Sit f Sunwapta Pass, Jasper, AB UBC 1567 1945 Sit f Coleman, AB UBC 8796 -Sit f South of Waterton Park, AB UBC 1569 1945 Sit f Maynard Pass, Jasper, AB UBC 1471 1945 Sit m Pine R., BC UBC 5625 1929 Sit m Akamina Pass, BC UBC 3553 1951 Sit m Monashee Pass, BC UBC 5627 1937 Sit m Well's Gray Park, BC UBC 3208 1950 Sit m Waterton Park, AB UBC 1565 1945 Sit m Jasper, AB UBC 2181 1946 Sit m 20 miles W of Banff, AB UBC 747 1943 Sit m Creston, BC UBC 447 1941 Sit m Toby Ck, BC UBC 1568 1945 Sit m Jasper, AB UBC 2180 1946 Sit m Bow L, Banff, AB UBC 1571 1945 Sit m Thompson Pass, AB UBC 1614 1945 Slco f Garfield Co., WA WSU 77690 1977 Slco f Garfield Co., WA WSU 46156 -Slco f Garfield Co., WA WSU 54191 1954 Slco f Garfield Co., WA WSU 78347 1977 Slco f Asotin Co., WA WSU 53237 1953 Slco m Asotin Co., WA WSU 4274 1942 Slco m Garfield Co., WA WSU 55420 -Slco m Columbia Co., WA WSU 46244 -Slco m Garfield Co., WA WSU 5836 -Slco m Wallowa Co., OR WSU 82113 1981 Slco m Columbia Co., WA WSU 46226 1946 Slco m Garfield Co., WA WSU 7855 1977 A p p e n d i x 1. C o n t i n u e d . Species Sex Location Museum ID# Date Slco m Garfield Co., WA WSU 7854 1977 Slco m Garfield Co., WA WSU 48338 1948 Slco f Wallowa Co., OR UWA 884 1943 Slco f Asotin-Garfield Co., WA UWA 905 1943 Slco f Wallowa Co., OR UWA 921 1943 Slco f Asotin-Garfield Co., WA UWA 912 1943 Slco f Wallowa Co., OR UWA 882 1943 Slco m Wallowa Co., OR UWA 883 1943 Slco m Asotin-Garfield Co., WA UWA 12803 1943 Slco m Wallowa Co., OR UWA 881 1943 Slco f Asotin Co., WA UPS 738 1939 Slco m Asotin Co., WA UPS 740 1939 Slch f Grant Co., OR WSU 82121 1981 Slch f Hood R. Co., OR WSU 82112 1980 Slch f Hood R. Co., OR WSU 82109 1980 Slch f Deschutes Co., OR WSU 80741 1980 Slch f Hood R. Co., OR WSU" 82108 1980 Slch f Grant Co., OR WSU 82119 1981 Slch f Plush Lk Co., OR WSU 57187 1957 Slch f Hood R. Co., OR WSU 81307 -Slch f Hood R. Co., OR WSU 82111 1980 Slch f Hood R. Co., OR WSU 82106 1980 Slch f Hood R. Co., OR WSU 82107 1980 Slch f Baker Co., OR WSU 82114 1981 Slch f Deschutes Co., OR WSU 80738 1980 Slch f Hood R. Co., OR WSU 82105 1980 Slch f Deschutes Co.," OR WSU 80740 1980 Slch f Lake Co., OR WSU 57189 1957 Slch m, Plush L Co., OR WSU 57188 1957 Slch m Grant Co., OR WSU 82120 1981 Slch m Deschutes Co., OR WSU 80739 1.980 Slch f Wasco Co., OR UWA 20074 1959 Slch m Wasco Co., OR UWA 20048 1958 Slch f Jefferson Co., OR UPS 8662 1956 Slch f Lake Co., OR UPS 8020 1951 Slch f Lake Co., OR UPS 2728 1951 Slch f Linn Co., OR UPS 8654 1952 Slch f Nevada Co., CA UPS 9371 1960 Slch f Linn Co., OR UPS 8658 1952 Slch f Lake Co., OR UPS 8022 1952 73 A p p e n d i x 1. C o n t i n u e d . Species Sex Location Museum ID# Date Slch f Lake Co., OR UPS 1976 1937 Slch f Jefferson Co., OR UPS 8007 1952 Slch f Jefferson Co., OR UPS 8008 1952 Slch f Lake Co., OR UPS 8014 1948 Slch f Klamath Co., OR UPS 8012 1945 Slch f Lake Co., OR UPS 2733 1951 Slch f Jefferson Co., OR UPS 8005 1952 Slch f Harney Co., OR UPS 1888 1937 Slch f Jefferson Co., OR UPS 8659 1956 Slch f Jefferson Co., OR UPS 8664 1956 Slch f Lake Co., OR UPS 2730 1951 Slch f Linn Co., OR UPS 8656 1956 Slch f Linn Co., OR UPS 11923 -Slch f Jefferson Co., OR UPS 8666 1956 Slch m Lake Co., OR UPS 8016 1950 Slch m Siskiyou Co., CA UPS 24364 1971 Slch m Jefferson Co., OR UPS 8003 1952 Slch m Klamath Co., OR UPS 13523 -Slch m Linn Co., OR UPS 7999 1952 Slch m Klamath Co., OR UPS 13521 Slch m Klamath Co., OR UPS 24362 Slch m Lake Co., OR UPS 8018 1950 Slch m Lake Co., OR UPS 2731 1951 74 A p p e n d i x 2 . T h e s u r v e y f o r m u s e d t o g a t h e r i n f o r m a t i o n o n S . s a t u r a t u s s i g h t i n g s f r o m t h e p u b l i c . Information on the Sighting of the Cascades Mantled Ground Squirrel  (Spermophilus Saturatus) 1) Date 2) Time 3) Location 4) Weather 5) Habitat type (eg. alpine meadow, rocky slope, f i r forest) 6) Number of individuals seen 7) Estimated distance from observer 8) Where was the squirrel(s) and what was i t doing? (eg. on ground feeding on leaves) 9) Apparent reaction of squirrel to observer 10) Observer act iv i ty when he/she came upon the squirrel 11) Observer's name address (optional) 12) Additional comments on back Identification: larger than chipmunk; no stripes on face (distinguishes i t from chipmunks); poorly defined black stripes on sides (distinguishes i t from the golden mantled ground squirrel) PLEASE return form to: Maria Leung Department of Animal Science PHOTOGRAPH OF SPERMOPHILUS SATURATUS HERE 75 A p p e n d i x 3 . O r g a n i z a t i o n s a n d p e r s o n s t o whom s u r v e y f o r m s w e r e d i s t r i b u t e d . " * " i n d i c a t e s a r e s p o n d e n t . 1 . A l p i n e C l u b o f C a n a d a , V a n c o u v e r , B . C . 2 . B . C . M o u n t a i n e e r i n g C l u b , V a n c o u v e r , B . C . * 3 . B a l l a m , I . , B u r n a b y , B . C . * 4 . C a n n i n g s , R . , V a n c o u v e r , B . C . * 5 . C a t h e d r a l L a k e s P r o v i n c i a l P a r k S t a f f , B . C . 6 . C e n t r a l O k a n a g a n N a t u r a l i s t C l u b , K e l o w n a , B . C . * 7 . C h i l l i w a c k F i e l d N a t u r a l i s t s , V e d d e r C r o s s i n g , B . C . * 8 . C h i l l i w a c k O u t d o o r C l u b , V e d d e r C r o s s i n g , B . C . * 9 . D a v i d s o n , P . , P r i n c e t o n , B . C . * 1 0 . F o l k a r d , N . , V a n c o u v e r , B . C . 1 1 . F r a s e r V a l l e y H i k e r s , M i s s i o n , B . C . 1 2 . K l i s t e r O u t d o o r K l u b , V a n c o u v e r , B . C . * 1 3 . L o g a n , I . , T u l a m e e n , B . C . * 1 4 . M a n n i n g P r o v i n c i a l P a r k S t a f f , B . C . 1 5 . N . V . R . C . O u t d o o r C l u b , N o r t h V a n c o u v e r , B . C . 1 6 . N o r t h O k a n a g a n N a t u r a l i s t s , V e r n o n , B . C . 1 7 . N o r t h S h o r e H i k e r s , V a n c o u v e r , B . C . 1 8 . O l i v e r - O s o y o o s N a t u r a l i s t s , O s o y o o s , B . C . 1 9 . O t h e r i n t e r e s t e d i n d i v i d u a l s . 2 0 . O u t s e t t e r s C l u b , V a n c o u v e r , B . C . 2 1 . R i c h m o n d O u t d o o r s C l u b , R i c h m o n d , B . C . 2 2 . S o u t h O k a n a g a n N a t u r a l i s t s , P e n t i c t o n , B . C . 2 3 . V a n c o u v e r N a t u r a l H i s t o r y S o c i e t y , V a n c o u v e r , B . C . 2 4 . V a n c o u v e r O u t d o o r C l u b , V a n c o u v e r , B . C . 2 5 . V a l l e y O u t d o o r A s s o c i a t i o n , W h i t e R o c k , B . C . 2 6 . V a r s i t y O u t d o o r C l u b , V a n c o u v e r , B . C . 2 7 . W e s t e r n C a n a d a W i l d e r n e s s C o m m i t t e e , V a n c o u v e r , B . C . 2 8 . Y - O u t d o o r C l u b , V a n c o u v e r , B . C . 76 A p p e n d i x 4 . L o c a t i o n s w h e r e S . s a t u r a t u s w e r e s i g h t e d i n B r i t i s h C o l u m b i a . E a c h s i g h t i n g ' s r e s p e c t i v e a l t i t u d e , m e t h o d o f s u r v e y b y w h i c h t h e a n i m a l was d e t e c t e d , a n d b i o g e o c l i m a t i c z o n e a r e g i v e n ( k e y f o l l o w s A p p e n d i x 5 ) . M e t h o d s o f s u r v e y : F = s u r v e y f o r m ; R = r o a d / o p p o r t u n i s t i c s i g h t i n g ; 0 = o b s e r v a t i o n w i t h t i m e t o f i r s t s i g h t i n g i n p a r e n t h e s e s . L o c a t i o n / c o o r d i n a t e s A l t i t u d e S u r v e y B i o g e o c l i -m e t h o d m a t i c z o n e A . I r o n M t n 5 0 ° 0 3 ' N 1 2 0 ° 4 6 'W 1650 B . G i l l i s L a k e c a t t l e g u a r d 4 9 ° 4 4 ' N 1 2 0 ° 5 5 'W 950 C . W o f G i l l i s L a k e ' 4 9 ° 5 4 ' N 1 2 1 ° 0 2 ' W 800 D. G i l l i s M t n 4 9 ° 5 4 ' N 1 2 0 ° 5 9 ' W 1550 E . W o f B r o o k m e r e 4 9 ° 5 0 ' N 1 2 0 ° 5 4 ' W 1000 F . B r o o k m e r e G a r b a g e Dump 4 9 ° 4 9 ' N 1 2 0 ° 5 2 ' W 1000 G . N o f T h y n n e M t n 4 9 ° 4 6 ' N 1 2 0 ° 5 5 ' W 1450 H . T h y n n e M t n 4 9 ° 4 2 ' N 1 2 0 ° 5 5 ' W 1950 I . S o f C o q u i h a l a L a k e s 4 9 ° 3 8 ' N 1 2 1 ° 0 0 ' W 1100 J l . 4 km SW o f C o q u i h a l a L a k e s 4 9 ° 3 6 ' N 1 2 1 ° 0 4 ' W 1000 J 2 . 8km SW o f C o q u i h a l a L a k e s 4 9 ° 3 5 ' N 1 2 1 ° 0 6 ' W 1000 K . N o f G r a s s h o p p e r M t n 4 9 ° 3 5 « N 1 2 0 ° 5 4 " W 1250 L . W o f N e e d l e P e a k ' 4 9 ° 3 4 ' N 1 2 1 ° 0 7 ' W 1500 M . NW o f C o a l m o n t 4 9 ° 3 2 ' N 1 2 0 ° 4 3 ' W 800 N . r o a d t o L o d e s t o n e L a k e 4 9 ° 3 0 ' N 1 2 0 ° 4 6 ' W ?1350 0 . N o f T u l a m e e n R i v e r 4 9 ° 2 9 ' N 1 2 0 ° 5 9 ' W 1000 P . B l a c k b u r n R o a d 4 9 ° 2 9 ' N 1 2 0 ° 4 4 ' W 1250 Q . b e t w e e n C o a l m o n t a n d P r i n c e t o n 4 9 ° 3 0 ' N 1 2 0 ° 3 9 " W 800 R . NWW o f P r i n c e t o n 4 9 ° 2 8 ' N 1 2 0 ° 3 7 ' W 700 0 ( 6 0 ) R R 0 ( 3 1 ) R 0 ( 5 ) R 0 ( 7 2 ) R R R F R F R 0 ( 3 1 ) F R MSxk E S S F d c p I D F d k I D F d k E S S F d c E S S F d c p ESSFwm MSdm, ESSFwm I D F x h MSdm MS I D F d k I D F d k A p p e n d i x 4 . C o n t i n u e d . L o c a t i o n / c o o r d i n a t e s S . T . U . V . W. X . Y . Z . A A . A B . A C . A D . A E . A F . A G . A H . A l . NE o f Mt S n i d e r 4 9 ° 2 6 N 1 2 1 ° 0 7 ' W T r e a s u r e M t n 4 9 ° 2 5 ' N 1 2 1 ° 0 3 ' W M t O u t r a m 4 9 ° 1 7 ' N 1 2 1 ° 1 0 , W A l t i t u d e 1200 S u r v e y B i o g e o c l i -m e t h o d m a t i c z o n e 1700 1 8 0 0 -2100 3km N o f D r y L a k e 4 9 ° 1 7 ' N 1 2 1 ° 0 2 ' W W e n d o f M a n n i n g P a r k 4 9 0 1 3 ! N 1 2 1 ° 0 1 ' W K i c k i n g H o r s e C a m p g r o u n d , M a n n i n g P a r k 4 9 ° 1 1 ' N 1 2 0 ° 4 9 ' W G a r i s o n L a k e 4 9 ° 1 3 ' N 1 2 0 ° 4 1 ' W E o f C o p p e r C r e e k 4 9 ° 1 5 ' N 1 2 0 ° 4 6 ' W 8 km E o f M a n n i n g P a r k 4 9 ° 1 1 ' N 1 2 0 ° 3 3 ' W F i r s t B r o t h e r M t n 4 9 ° 0 5 ' N 1 2 0 ° 4 6 ' W W i l l i a m ' s P e a k 4 9 ° 0 7 ' N 1 2 1 ° 3 0 ' W S t r a w b e r r y F l a t s , M a n n i n g P a r k 4 9 ° 0 4 ' N 1 2 0 ° 5 3 ' W C o l d s p r i n g C a m p g r o u n d , M a n n i n g P a r k 4 9 ° 0 5 ' N 1 2 0 o 4 8 ' W " C a s c a d e L o o k o u t , M a n n i n g P a r k 4 9 ° 0 4 ' N 1 2 0 ° 4 6 ' W R e s o r t L o d g e , C a t h e d r a l P a r k 4 9 ° 0 4 ' N 1 2 0 ° 1 2 ' W N o f L a d y s l i p p e r L a k e , C a t h e d r a l P a r k 4 9 ° 0 3 ' N 1 2 0 ° 1 1 ' W L a k e v i e w M t n 4 9 ° 0 2 ' N 1 2 0 ° 0 9 ' W 2500 1200 1000 1900 1300 1000 1050 2250 1900 1400 1250 1750 2100 2250 0 ( 5 4 ) 0 ( 4 9 ) 0 ( 3 1 ) F F F F R R 0 ( 5 3 ) 0 ( 4 5 ) 0 ( 3 5 ) 0 ( 5 ) 0 ( 1 ) , F 0 ( 1 ) 0 ( 4 4 ) 0 ( 3 1 ) A E p , Mb A E p A E p MS dm I D F d k I D F d k A E p Mb E f I F u E e E S S F x c A T E x c p A T E x c p A p p e n d i x 5 . L o c a t i o n s s u r v e y e d w h e r e no S . s a t u r a t u s was s i g h t e d . T h e r e s p e c t i v e a l t i t u d i n a l r a n g e s , n u m b e r o f h o u r s s p e n t o b s e r v i n g , a n d b i o g e o c l i m a t i c z o n e s a r e i n c l u d e d ( k e y f o l l o w s t h i s t a b l e ) . L o c a t i o n / c o o r d i n a t e s A l t i t u d e O b s e r v a t i o n B i o g e o c l i -r a n g e h o u r s m a t i c z o n e 1. Mt T o d d 5 0 ° 5 5 ' N 1 1 9 ° 5 6 ! W 2100 5 — 2 . C o r n w a l l H i l l s 5 0 ° 4 2 ' N 1 2 1 ° 2 8 ' W 2000 2 — 3 . Mt S a v o n a 5 1 ° 4 2 ' N 1 2 0 ° 4 9 * W 1500 3 — 4 . G r e e n s t o n e M t n 5 0 ° 3 7 ' N 1 2 0 ° 3 8 ' W 1800 3 — 5 . C h u w e l s M t n 5 0 ° 3 2 ' N 1 2 0 ° 4 ' W 1900 4 — 6. T u k t a k a m i n M t n 5 0 ° 2 8 ' N 1 1 9 ° 3 4 , W 1800 4 MSdm 7 . N o f D e v i l ' s L a k e 5 0 ° 2 5 ' N 1 2 1 ° 5 3 ' W . , 1700 - 2 1 0 0 10 — 8 . B o t a n i e M t n 5 0 ° 2 0 ' N 1 2 1 ° 3 6 ' W 1400 - 1 9 0 0 6 9 . S i l v e r s t a r M t n 5 0 ° 2 2 ' N 1 1 9 ° 0 3 I W 1600 - 1 8 5 0 6 E S S F d c 1 0 . Swakum M t n 5 0 ° 1 8 ' N 1 2 0 ° 4 2 ' W 1700 3 MSxk 1 1 . P r o m o n t o r y H i l l s 5 0 ° 1 2 ' N 1 2 0 ° 5 8 ' W 1 0 0 0 , 4 MSxk 1700 12 . Mt H a m i l t o n 5 0 ° 0 8 ' N 1 2 0 ° 2 1 ' W 1500 2 I D F d k 13 . T e r r a c e M t n 5 0 ° 0 6 ' N 1 1 9 ° 3 7 ' W 1600 - 1 9 0 0 6 E S S F x c 1 4 . E u r e k a M t n 5 0 ° 0 5 ' N 1 1 8 0 2 3 ' W 1900 2 — 1 5 . S t o y o m a M t n 4 9 ° 5 9 , N 1 2 1 ° 1 3 ' W 2200 5 -1 6 . K a n e V a l l e y T r a i l 4 9 ° 4 9 ' N 1 2 0 ° 4 4 ' W 1350 10 I D F d k 1 7 . C a r r o t M t n 4 9 ° 4 5 ' N n g ^ ' w 1500 6 MSdm 18 . E o f B o s t o n B a r 1100 1 E f , AMp 4 9 ° 5 4 ' N 1 2 1 ° 2 3 ' W 1 9 . W o f G i l l i s L a k e 4 9 ° 5 5 ' N 1 2 1 ° 0 5 ' W 1300 3 — 2 0 . B l u e y L a k e 4 9 ° 5 2 ' N 1 2 0 ° 3 5 ' W 1100 2 I D F d k 2 1 . S E o f B o s t o n B a r 4 9 ° 3 9 ' N 121o22 'W 800 2 Hb 22 . U z t i m u s Rd 4 9 ° 4 7 ' N 1 2 1 ° 2 0 ' W . 500 1 H e , Hb A p p e n d i x 5 . C o n t i n u e d . L o c a t i o n / c o o r d i n a t e s A l t i t u d e O b s e r v a t i o n B i o g e o c l i -r a n g e h o u r s m a t i c z o n e 23 . 8km E o f B r o o k m e r e 4 9 ° 4 9 ' N 1 2 0 ° 4 0 ' W 1100 1 — 24 . S i w a s h L a k e Rd 4 9 ° 4 3 ' N 1 2 0 ° 2 1 ' W 1250 3 I D F d k 2 5 . Mt P i k e 4 9 ° 4 3 ' N 1 2 0 ° 4 0 ' W 1400 8 MS dm 2 6 . S E o f B o s t o n B a r 4 9 ° 4 0 ' N 1 2 1 0 2 4 ' W 500 2 Hb 2 7 . M i s s e z u l a M t n 4 9 ° 4 0 ' N 1 2 0 ° 3 2 ' W 1500 6 E S S F d c , MSdm 2 8 . C h a i n L a k e 4 9 ° 4 4 ' N 1 2 0 ° 1 9 ' W 1350 2 MS dm 2 9 . R a m p a r t L a k e 4 9 ° 3 9 ' N 1 2 0 ° 3 0 ' W 1400 2 . I D F d k 3 0 . N o f S p u z z u m C r e e k 4 9 ° 3 8 ' N 1 2 1 ° 3 4 ' W 1000 4 M b , AMp 3 1 . D o u g l a s L a k e T r a i l 4 9 ° 3 8 ' N 1 2 1 ° 0 8 ' W 1300 4 -32 . I s i n t o k M t n 4 9 ° 3 4 ' N 1 1 9 ° 5 8 ' W 1800 5 MSdm 33 . E o f N e e d l e P e a k 4 9 ° 3 4 ' N 1 2 1 ° 0 6 ' W 1500 5 H b , Mb 3 4 . E m a n c i p a t i o n M t n 4 9 ° 3 0 , N 1 2 1 ° 1 6 ' W 1100 5 A M p , Mb 3 5 . L o d e s t o n e M t n 4 9 ° 2 8 ' N 1 2 1 ° 5 0 ' W 1850 6 E S S F d c p 3 6 . Dog M t n 4 9 ° 2 4 ' N 1 2 1 ° 3 2 » W 1000 2 M a , Hb 37 . O g l i v i e M t n 4 9 ° 2 5 ' N 1 2 1 ° 2 3 ' W 750 14 A M p , Mb - 1 0 0 0 3 8 . S t e m w i n d e r Rd 4 9 ° 2 5 ' N 1 2 0 ° 0 9 ' W 1400 4 I D F d k 3 9 . A p e x M t n 4 9 ° 2 8 ' N 1 1 9 ° 5 5 ' W 2200 7 E S S F x c p 4 0 . A g a t e M t n 4 9 ° 2 3 ' N 1 2 0 ° 2 4 ' W 1600 4 MSdm 4 1 . S o f E u r e k a C r e e k 4 9 ° 2 0 I N 1 2 1 ° 2 8 ' W 500 - 9 0 0 10 H b , Mb 42 . W h i s t l e C r e e k Rd 4 9 ° 1 8 ' N 1 2 0 ° 0 6 ' W 1600 6 I D F d k , MSxk 43 . NW o f I R 1 0 4 9 ° 1 6 ' N 1 2 0 ° 0 9 ' W 2000 6 I D F d k , MSxk 44 . O r o f i n o P e a k 4 9 ° 1 5 , N l i g ^ l ' W 1500 3 I D F d k 4 5 . Y o l a C r e e k 4 9 ° 1 1 ' N 1 2 1 ° 2 6 ' W 1000 3 H b , Mb 4 6 . C r a t e r M t n 2000 7 A T E x c p , 4 9 ° 1 1 ' N 1 2 0 ° 0 5 ' W - 2 3 0 0 E S S F X C A p p e n d i x 5 . C o n t i n u e d . L o c a t i o n / c o o r d i n a t e s 4 7 . S i l v e r h o p e C r e e k 4 9 ° 0 7 ' N 1 2 1 ° 2 2 ' W 4 8 . Shawatum M t n 4 9 ° 0 7 ' N 1 2 1 ° 0 5 ' W 4 9 . Mt K o b a u 4 9 ° 0 7 ' N 1 1 9 ° 4 0 ' W 5 0 . W o f D u r u i s s e a u Ck 4 9 ° 0 6 ' N 1 2 0 ° 2 3 ' W 5 1 . S o f Nepopekum M t n 4 9 ° 0 3 ' N 1 2 1 ° 0 0 ' W 5 2 . C h u r c h M t n 4 9 ° 0 2 ' N 1 2 1 ° 5 1 ' W 5 3 . S k y l i n e T r a i l , M a n n i n g P a r k 4 9 ° 0 2 ' N 1 2 0 ° 5 5 ' W 5 4 . W o f Mt K r u g e r 4 9 ° 0 2 ' N 1 1 9 ° 3 2 ' W 5 5 . . F r o s t y M t n 4 9 ° 0 1 ' N 1 2 0 ° 5 0 ' W A l t i t u d e O b s e r v a t i o n B i o g e o c l i -r a n g e h o u r s m a t i c z o r 1000 6 H b , Mb 650 12 E f 1800 13 M S x k , I D F d k 1300 8 E S S F x c - 1 7 0 0 600 16 E f - 1 5 0 0 1000 4 Mb - 1 2 5 0 1300 16 E f , A E p - 1 7 0 0 1000 6 I D F x h 1300 19 A E p - 2 4 0 0 81 K e y t o B i o g e o c l i m a t i c Z o n e s ( b a s e d on K r a j i n a 1959) A . V a n c o u v e r F o r e s t R e g i o n AMp - u n d i f f e r e n t i a t e d a l p i n e t u n d r a a n d u n d i f f e r e n t i a t e d p a r k l a n d m o u n t a i n h e m l o c k A E p - u n d i f f e r e n t i a t e d a l p i n e t u n d r a a n d u n d i f f e r e n t i a t e d p a r k l a n d E n g l e m a n n s p r u c e - s u b a l p i n e f i r Ma - m a r i t i m e f o r e s t e d m o u n t a i n h e m l o c k Mb - s u b m a r i t i m e f o r e s t e d m o u n t a i n h e m l o c k E e - d r y s o u t h e r n f o r e s t e d E n g l e m a n n s p r u c e - s u b a l p i n e f i r E f - s u b c o n t i n e n t a l f o r e s t e d E n g l e m a n n s p r u c e - s u b a l p i n e f i r I F u - u n d i f f e r e n t i a t e d i n t e r i o r d o u g l a s f i r Hb5 - s o u t h e r n s u b m a r i t i m e c o a s t a l w e s t e r n h e m l o c k H c l - d r i e r m a r i t i m e c o a s t a l w e s t e r n h e m l o c k B . K a m l o o p s F o r e s t R e g i o n I D F x h - v e r y d r y h o t i n t e r i o r d o u g l a s f i r I D F d k - d r y c o o l i n t e r i o r d o u g l a s f i r MSxk - v e r y d r y c o o l m o n t a n e s p r u c e MSdm - d r y m i l d m o n t a n e s p r u c e MS - u n d i f f e r e n t i a t e d m o n t a n e s p r u c e E S S F x c p - v e r y d r y c o l d E n g l e m a n n s p r u c e - s u b a l p i n e f i r E S S F d c p - d r y c o l d E n g l e m a n n s p r u c e - s u b a l p i n e f i r ESSFwm - w e t m i l d E n g l e m a n n s p r u c e - s u b a l p i n e f i r A T E x c p - v e r y d r y c o l d a l p i n e t u n d r a E n g l e m a n n s p r u c e -s u b a l p i n e f i r p a r k l a n d 82 A p p e n d i x 6 - T r a p p i n g e f f o r t , l o c a t i o n s , a n d m a m m a l i a n s p e c i e s c a p t u r e d . " * " i n d i c a t e s u s e o f S h e r m a n f o l d i n g t r a p s . L o c a t i o n D a t e 1989 B l a c k b u r n R d May 2 6 -27 Y o l a C r e e k Rd J u n e 7 S u t t e r C r e e k J u n e 2 0 T h y n n e M t n J u n e 2 3 -24 T r e a s u r e M t n J u l y 6 J u l y 8 J u l y 12 -13 A u g 3-4 A u g 2 3 - 2 4 K o b a u M t n T h y n n e M t n T h y n n e M t n T h y n n e M t n T h y n n e M t n T h y n n e M t n 1990 C a s c a d e L o o k o u t , M a n n i n g P a r k May 29 - J u n e 1 B r o o k m e r e Dump J u n e 5-8 Number o f S p e c i e s c a u g h t t r a p p i n g d a y s 226 E u t a m i a s amoenus T a m i a s c i u r u s h u d s o n i c u s I r o n M t n J u n e 1 4 - 1 7 25 30 n i l E u t a m i a s amoenus  T a m i a s c i u r u s h u d s o n i c u s 100 E u t a m i a s amoenus S p e r m o p h i l u s s a t u r a t u s 60 E u t a m i a s amoenus E u t a m i a s t o w n s e n d i i 25 E u t a m i a s amoenus 200 E u t a m i a s amoenus S p e r m o p h i l u s c o l u m b i a n u s  S p e r m o p h i l u s s a t u r a t u s 120 E u t a m i a s amoenus S p e r m o p h i l u s s a t u r a t u s 120 +22* E u t a m i a s amoenus C l e t h r i o n o m y s q a p p e r i  P e r o m y s c u s m a n i c u l a t u s  S p e r m o p h i l u s s a t u r a t u s S e p t . 9 - 1 0 120 S e p t . 2 3 - 2 4 120 E u t a m i a s amoenus  S p e r m o p h i l u s s a t u r a t u s E u t a m i a s amoenus E u t a m i a s amoenus 80 S p e r m o p h i l u s s a t u r a t u s T a m i a s c i u r u s h u d s o n i c u s 80 160 E u t a m i a s amoenus  S p e r m o p h i l u s s a t u r a t u s E u t a m i a s amoenus  M a r m o t a f l a v i v e n t r i s T h y n n e M t n J u l y 4 -8 160 E u t a m i a s amoenus  S p e r m o p h i l u s s a t u r a t u s A p p e n d i x 7 A . The p e r c e n t c o v e r o f p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s : HERB L A Y E R . L o c a t i o n s : B r o o k m e r e Dump ( B D ) , C o l d s p r i n g C a m p g r o u n d ( C C ) , C a s c a d e L o o k o u t ( C L ) , C a t h e d r a l P a r k s l o p e ( C P ) , I r o n M o u n t a i n ( I M ) , L a d y s l i p p e r L a k e ( L L ) , M o u n t O u t r a m (MO), Q u i n i s c o e C a m p g r o u n d ( Q C ) , T h y n n e M o u n t a i n ( T M ) . T h e number o f p l o t s s a m p l e d a t e a c h l o c a t i o n a r e : B D ( 2 5 ) , C C ( 1 0 ) , C L ( 1 5 ) , C P ( 1 0 ) , I M ( 2 5 ) , L L ( 1 0 ) , M O ( 2 5 ) , T M ( 2 5 ) . Apiaceae Apocynaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Location Lomatium Apocynum Achillia Antennaria Arnica Aster Cirsium Erigc BD mean 0.00 2.92 0.38 0.29 2.28 0.00 . 2.31 0.00 sd 0.00 10.43 1.34 1.43 6.56 0.00 6.28 0.00 CC mean 0.43 0.00 1.03 0.48 3.56 0.80 0.00 0.00 sd 1.35 0.00 3.27 1.51 2.78 2.53 0.00 0.00 CL mean 0.00 0.00 0.00 0.37 1.93 0.00 0.00 0.00 sd 0.00 0.00 0.00 1.42 5.47 0.00 0.00 0.00 CP mean 0.00 0.00 3.05 0.80 5.13 0.00 0.43 3.15 sd 0.00 0.00 2.20 1.75 5.96 0.00 1.37 3.53 IM mean 0.00 0.00 2.39 0.42 5.84 0.00 0.00 1.33 sd 0.00 0.00 2.87 1.49 7.51 0.00 0.00 4.07 LL mean 0.00 0.00 0.29 2.62 0.00 0.00 0.00 6.20 sd 0.00 0.00 0.93 2.28 0.00 0.00 0.00 2.44 MO mean 0.00 0.00 0.63 4.26 2.45 0.00 0.00 0.49 sd 0.00 0.00 1.59 4.48 5.41 0.00 0.00 1.02 QC mean 0.00 0.00 0.53 0.00 0.00 0.00 0.00 0.11 sd 0.00 0.00 1.50 0.00 0.00 0.00 0.00 0.53 TM mean 0.00 0.00 1.13 4.18 0.87 0.09 0.00 1.74 sd 0.00 0.00 1.68 5.39 2.10 0.44 0.00 3.16 CO A p p e n d i x 7 A . HERB L A Y E R ( c o n t i n u e d ) . Asteraceae Asteraceae Asteraceae Asteraceae Berberidaceae Boraginaceae Caprifoliaceae Cary< Location Hieracium Senecio Solidago Taraxacum Berberis Unidentified Linnaea Aren< BD mean 0.00 0.00 0.00 0.00 0.32 0.27 0.00 0.00 sd 0.00 0.00 0.00 0.00 1.58 1.33 0.00 0.00 CC mean 0.53 1.98 0.00 0.00 0.00 0.00 1.26 0.00 sd 1.69 2.18 0.00 0.00 0.00 0.00 3.14 0.00 CL mean 2.15 0.74 0.39 0.00 0.00 0.00 0.00 0.37 sd 2.89 2.12 1.50 0.00 0.00 0.00 0.00 1.45 CP mean 0.83 0.00 0.53 0.00 0.00 0.00 0.00 0.51 sd 1.74 0.00 1.69 0.00 0.00 0.00 0.00 1.60 IM mean 0.00 0.43 0.00 1.28 0.21 0.00 0.00 1.09 sd 0.00 1.55 0.00 3.45 1.03 0.00 0.00 2.48 LL mean 0.00 1.08 0.00 0.00 0.00 0.00 0.00 3.91 sd 0.00 1.15 . 0.00 0.00 0.00 0.00 0.00 1.57 MO mean 2.24 0.34 0.00 0.10 0.00 0.00 0.00 4.14 sd 3.86 0.97 0.00 0.52 0.00 0.00 0.00 5.06 QC mean 0.00 0.09 0.51 0.00 0.00 0.00 0.00 0.12 sd 0.00 0.44 1.47 0.00 0.00 0.00 0.00 0.58 TM mean 0.00 1.85 0.00 0.00 0.00 0.00 0.00 1.92 sd 0.00 2.93 0.00 0.00 0.00 0.00 0.00 3.22 co A p p e n d i x 7 A . HERB L A Y E R ( c o n t i n u e d ) . Caryophyllaceae Celastraceae Cornaceae Crassulaceae Cruciferae Cruciferae Cyperaceae Empetrace< Location Stellarina Pachistima Cornus Sedum Arabis Draba Carex Empetrum BD mean 0.00 1.66 0.00 000 0.00 0.00 11.56 0.00 sd 0.00 2.29 0.00 000 0.00 0.00 14.36 0.00 CC mean 0.00 0.00 2.95 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 6.70 0.00 0.00 o.oo 0.00 0.00 CL mean 0.47 0.57 0.00 0.24 0.52 0.00 0.34 0.00 sd 1.24 2.23 0.00 0.95 2.02 0.00 1.32 0.00 CP mean 0.00 0.00 0.00 0.00 0.29 0.00 5.11 0.00 sd 0.00 0.00 0.00 0.00 0.92 0.00 4.20 0.00 IM mean 0.00 3.86 0.00 1.28 0.13 0.00 17.71 1.00 sd 0.00 6.67 0.00 1.98 0.66 0.00 16.11 5.00 LL mean 0.00 0.00 0.00 1.46 0.00 0.21 4.52 0.54 sd 0.00 0.00 0.00 1.30 0.00 0.66 5.53 1.70 MO mean 0.00 0.00 0.00 0.00 1.20 0.00 6.31 0.10 sd 0.00 0.00 0.00 0.00 2.56 0.00 9.08 0.49 QC mean 0.00 0.00 0.00 0.00 0.00 0.00 1.88 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 6.89 0.00 TM mean 0.00 0.10 0.00 1.05 0.12 0.00 1.69 0.00 sd 0.00 0.50 0.00 1.74 0.62 0.00 3.48 0.00 A p p e n d i x 7 A . HERB LAYER ( c o n t i n u e d ) . Ericaceae Ericaceae Ericaceae Ericaceae Ericaceae Fabaceae Fabaceae Fabaceae Faba Location Arctostaphvlos Cassiope Gaultheria Phyllodoce Vaccinium Lupinus Oxytropis Trifolium Vicia BD mean 5.67 0.00 0.00 0.00 0.00 0.09 0.00 0.00 0.00 sd 14.08 0.00 0.00 0.00 0.00 0.45 0.00 0.00 0.00 CC mean 0.00 0.00 0.00 0.00 19.45 0.42 0.00 0.71 0.00 sd 0.00 0.00 0.00 0.00 13.03 1.32 0.00 2.26 0.00 CL mean 0.00 0.00 0.00 0.00 17.44 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 17.38 0.00 0.00 0.00 0.00 CP mean 0.00 0.00 0.00 0.00 20.93 2.22 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 23.06 3.66 0.00 0.00 0.00 1M mean 3.48 0.00 0.00 0.00 7.71 1.31 0.00 0.00 0.67 sd 8.08 0.00 0.00 0.00 12.52 3.66 0.00 0.00 2.46 LL mean 7.83 0.72 0.00 0.42 3.28 11.49 2.62 0.00 0.00 sd 9.46 1.56 0.00 1.34 4.27 7.58 1.50 0.00 0.00 MO mean 0.00 2.76 0.00 6.12 5.47 3.29 0.00 0.00 0.00 sd 0.00 5.31 0.00 9.82 6.65 4.41 0.00 0.00 0.00 QC mean 0.49 0.00 0.00 2.80 18.73 0.13 0.00 0.00 0.00 sd 2.45 0.00 0.00 5.95 15.39 0.66 0.00 0.00 0.00 TM mean 0.00 0.00 0.19 0.66 3.04 2.81 0.00 0.00 0.00 sd 0.00 0.00 0.93 3.30 12.43 4.84 0.00 0.00 0.00 co A p p e n d i x 7 A . HERB L A Y E R ( c o n t i n u e d ) Location Gentianacea Hydrophyliacea Juncaceae Juncaceae Lilliaceae Gentianella Phacelia Juncus Luzula Clintonia BD mean 0.00 0.14 0.00 0.00 0.00 sd 0.00 0.68 0.00 0.00 0.00 CC mean 0.00 0.00 . 0.00 0.00 1.20 sd 0.00 0.00 0.00 0.00 3.79 CL mean 0.00 1.39 0.00 0.00 0.00 sd 0.00 2.94 0.00 0.00 0.00 CP mean 0.00 0.00 0.51 0.00 0.00 sd 0.00 0.00 1.60 0.00 0.00 IM mean 0.14 0.20 0.00 0.00 0.00 sd 0.70 1.00 0.00 0.00 0.00 LL mean 0.00 0.00 0.00 0.22 0.00 sd 0.00 0.00 0.00 0.68 0.00 MO mean 0.00 0.00 4.41 3.93 0.00 sd 0.00 0.00 4.43 4.86 0.00 QC mean 0.00 0.00 0.00 0.18 0.00 sd 0.00 0.00 0.00 0.91 0.00 TM mean 0.00 0.55 1.15 7.78 0.00 sd 0.00 1.57 4.06 10.67 0.00 Lilliaceae Erythronium 0.00 0.00 0.00 0.00 2.03 2.70 0.00 0.00 0.00 0.00 0.00 0.00 2.27 3.17 0.00 0.00 0.61 1.44 Lilliaceae Lilium 0.00 0.00 0.74 1.62 0.19 0.75 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Lilliaceae Smilacina 0.00 0.00 0.28 0.89 0.00 0.00 0.00 6.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 A p p e n d i x 7 A . HERB L A Y E R ( c o n t i n u e d ) . Orchidaceae Onagraceae Pinaceae Pinac Location Goodyera Epilobium Abies Picee BD mean 0.00 0.40 0.00 0.00 sd 0.00 1.17 0.00 0.00 CC mean 0.81 0.00 1.19 0.00 sd • 2.56 0.00 3.76 0.00 CL mean - 0.00 0.65 0.26 0.23 sd 0.00 1.78 1.00 0.91 CP mean 0.00 1.72 0.00 0.00 sd 0.00 3.94 0.00 0.00 IM mean 0.00 0.14 0.00 0.00 sd 0.00 0.68 0.00 0.00 LL mean 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 MO mean 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 QC mean 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 TM mean 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 Pinaceae Poaceae Poaceae Poaceae Poaceae Pinus Agropyron Bromus Calamagrostis Festuca 0.00 0.16 8.13 0.93 0.00 0.00 0.81 15.02 4.63 0.00 2.38 0.00 0.00 0.00 0.00 7.53 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 3.07 0.00 0.00 0.00 0.00 4.05 0.19 0.00 0.00 0.00 0.00 0.96 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.83 0.95 0.00 0.00 0.00 2.17 1.24 0.00 0.00 0.00 0.00 0.80 0.00 0.00 0.00 0.00 1.52 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0,00 0.00 0.00 0.00 1.21 0.00 0.00 0.00 0.00 2.41 03 CO A p p e n d i x 7 A . HERB LAYER ( c o n t i n u e d ) . Poaceae Poaceae Poaceae Poaceae Poaceae Polemoniaceae Polemoniaceae Polyc Location Hordeum Phleum Poa Trisetum Unidentified Phlox Polemonium Eriog BD mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 CC mean 0.00 0.00 0.00 0.00 11.70 1.28 0.00 0.00 sd 0.00 0.00 0.00 0.00 12.50 4.04 0.00 0.00 CL mean . 0.61 0.00 0.00 0.00 3.91 1.63 0.00 0.00 sd 1.64 0.00 0.00 0.00 6.01 2.43 0.00 0.00 CP mean 0.00 0.00 0.76 0.00 0.45 0.42 0.29 0.00 sd 0.00 0.00 2.40 0.00 1.42 , 1.33 0.92 0.00 IM mean 0.00 0.00 0.45 0.00 0.00 0.00 2.05 0.14 sd 0.00 0.00 1.71 0.00 0.00 0.00 4.95 0.69 LL mean 0.00 0.00 0.67 0.81 0.00 0.00 2.27 0.00 sd 0.00 0.00 1.45 1.33 0.00 0.00 1.47 0.00 MO mean 0.00 0.00 0.53 0.00 0.00 3.25 0.00 0.00 sd 0.00 0.00 1.09 0.00 0.00 3.93 0.00 0.00 QC mean 0.00 0.00 0.00 0.00 3.34 0.00 0.29 0.00 sd 0.00 0.00 0.00 0.00 6.36 0.00 1.03 0.00 TM mean 0.00 0.16 1.75 0.26 0.00 5.10 0.36 0.35 sd 0.00 0.82 3.62 0.90 0.00 4.92 1.28 1.22 A p p e n d i x 7A . HERB L A Y E R ( c o n t i n u e d ) . Polygonaceae Polypodiaceae Portulaceae Location Rumex Unidentified Claytonia BD mean 0.80 0.00 0.00 sd 3.16 0.00 0.00 CC mean 0.00 0.00 0.00 sd 0.00 o.oo- 0.00 CL mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 CP mean 0.00 0.72 0.00 sd 0.00 1.54 0.00 IM mean 0.17 0.33 0.00 sd 0.83 1.16 0.00 LL mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 MO mean 0.00 0.00 1.33 sd 0.00 0.00 3.54 QC mean 0.00 0.11 0.00 sd 0.00 0.53 0.00 TM mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 Portulaceae Pyrolaceae Ranunculaceae Ranunculaceae Montia Pyrola Anemone Ranunculus o.bo 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.53 0.00 0.00 0.00 1.69 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.51 1.14 0.00 0.00 1.60 1.87 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 3.30 0.00 0.00 0.00 2.32 0.00 0.00 0.10 1.22 0.00 0.00 0.48 2.09 0.00 0.00 0.00 0.09 0.00 0.00 0.00 0.44 0.00 0.28 0.00 0.15 0.00 0.98 0.00 0.76 0.00 A p p e n d i x 7 A . HERB LAYER ( c o n t i n u e d ) . Ranunculaceae Ranunculaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosa Location Thalictrum Trbllius Amelanchier Dryas Fragaria Luetkea Potentilla Rubus Sibbi BD mean 0.00 0.51 0.59 0.00 0.30 0.00 0.96 0.00 0.00 sd 0.00 2.56 2.13 0.00 1.05 0.00 2.68 0.00 0.00 CC mean 0.00 0.00 0.00 0.00 0.00 0.00 0.54 0.28 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 1.71 0.89 0.00 CL mean 0.00 0.00 0.52 0.00 0.41 0.00 0.00 0.00 0.00 sd 0.00 0.00 1.38 0.00 1.59 0.00 0.00 0.00 0.00 CP mean 0.00 0.00 0.00 0.00 4.52 0.00 1.38 0.00 0.00 sd 0.00 0.00 0.00 0.00 4.23 0.00 2.37 0.00 0.00 IM mean 0.00 0.00 1.22 0.15 1.51 0.00 0.24 0.00 0.00 sd 0.00 0.00 2.64 0.77 2.32 0.00 0.84 0.00 0.00 LL mean 0.00 0.00 0.00 10.23 0.22 0.00 4.24 0.00 0.00 sd 0.00 0.00 0.00 4.28 0.68 0.00 1.83 0.00 0.00 MO mean 0.11 0.00 0.00 0.00 0.00 5.26 0.70 0.00 0.45 sd 0.54 0.00 0.00 0.00 0.00 5.91 2.64 0.00 1.41 QC mean 0.00 0.16 0.00 0.00 0.58 0.00 0.37 0.00 0.00 sd 0.00 0.79 0.00 0.00 1.66 0.00 1.03 0.00 0.00 TM mean 0.00 0.00 0.00 0.00 0.00 3.14 0.88 0.00 0.27 sd 0.00 0.00 0.00 0.00 0.00 7.35 3.51 0.00 0.97 A p p e n d i x 7 A . HERB L A Y E R ( c o n t i n u e d ) Saxifragaceae Saxifragaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scro| Location Heuchera Saxifraga Castilleja Collinisia Pedicularis Pens BD mean 0.00 0.00 0.00 0.00 0.00 0.71 sd 0.00 0.00 0.00 0.00 0.00 2.73 CC mean 0.00 0.00 . 0.00 0.46 0.00 0.00 sd 0.00 0.00 0.00 1.45 0.00 0.00 CL mean 0.00 0.00 0.00 0.00 1.25 1.08 sd 0.00 0.00 0.00 0.00 2.76 2.36 CP mean 0.51 0.43 0.00 0.00 0.00 0.00 sd 1.60 1.36 0.00 0.00 0.00 0.00 IM mean 0.21 0.28 0.13 0.00 0.00 1.49 sd 1.04 0.99 0.66 0.00 0.00 2.20 LL mean 0.24 2.11 0.00 0.00 0.00 5.06 sd 0.76 1.58 0.00 0.00 0.00 5.07 MO mean 0.00 0.69 1.21 0.00 2.21 0.65 sd 0.00 1.63 2.56 0.00 4.48 1.36 QC mean 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 TM mean 0.14 0.34 0.00 0.00 1.55 3.85 sd 0.68 1.23 0.00 0.00 2.96 5.68 to A p p e n d i x 7 A . HERB LAYER ( c o n t i n u e d ) . Scrophulariaceae Scrophulariaceae Valerianaceae Violai Location Verbascum Veronica Valeriana Viola BD mean 2.10 0.00 0.00 0.00 sd 3.27 0.00 0.00 0.00 CC mean 0.00 0.00 0.00 1.10 sd 0.00 0.00 0.00 1.91 CL mean 0.00 0.00 0.00 0.57 sd 0.00 0.00 0.00 1.54 CP mean 0.00 0.40 2.97 0.00 sd 0.00 1.25 9.39 0.00 IM mean 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 LL mean 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 MO mean 0.00 o.bo 2.70 0.26 sd 0.00 0.00 6.53 1.31 QC mean 0.00 0.00 0.40 0.00 sd 0.00 0.00 1.99 0.00 TM mean 0.00 0.00 0.00 0.18 sd 0.00 0.00 0.00 0.91 Unvegetated Bedrock Unvegetated Gravel/debris/soil 0.42 53.85 1.46 23.60 0.00 38.58 0.00 19.23 1.58 31.63 4.91 28.38 0.00 15.65 0.00 12.29 19.32 20.13 17.44 13.09 0.00 13.19 0.00 . 7.59 0.66 11.53 2.83 7.31 0.00 52.72 0.00 25.28 7.29 37.40 9.60 23.54 A p p e n d i x 7A. HERB LAYER ( c o n t i n u e d ) . Unvegetated Unvegetated Location Stumps/deadfall Garbage Large BD mean 0.34 0.91 1.00 sd 1.70 3.13 2.64 CC mean 0.46 0.00 3.89 sd 1.45 0.00 6.74 CL mean 1.24 0.00 18.61 sd 3.54 0.00 33.56 CP mean 0.00 0.00 21.59 sd 0.00 0.00 30.31 IM mean 1.03 0.00 0.00 sd 2.46 0.00 0.00 LL mean 0.00 0.00 7.47 sd 0.00 0.00 9.00 MO mean 0.27 0.00 8.09 sd 1.34 0.00 10.61 QC mean 0.96 0.00 2.66 sd 3.33 0.00 5.50 TM mean 2.71 0.00 0.20 sd 7.59 0.00 1.00 Unvegetated Snow Unvegetated Tarmac Unvegetated Sign/building Unvegetated Water 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.98 0.00 0.00 0.00 2.06 0.00 0.00 6.67 0.00 0.00 0.00 25.82 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 8.67 0.00 0.00 0.00 24.94 0.00 0.00 0.00 0.00 0.00 0,76 12.00 0.00 0.00 3.81 33.17 0.88 0.00 0.00 0.00 4.41 0.00 0.00 0.00 4^  A p p e n d i x 7 B . T h e p e r c e n t c o v e r o f p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s : SHRUB L A Y E R . L o c a t i o n s : B r o o k m e r e Dump ( B D ) , C o l d s p r i n g Campground (CC) , C a s c a d e L o o k o u t ( C L ) , C a t h e d r a l P a r k S l o p e ( C P ) , I r o n M o u n t a i n ( I M ) , L a d y s l i p p e r L a k e ( L L ) , M o u n t O u t r a m ( M O ) , Q u i n i s c o e C a m p g r o u n d ( Q C ) , T h y n n e M o u n t a i n ( T M ) . Apocynaceae Asteraceae Asteraceae Location Apocynum Achillia Anaphalis BD mean 0.80 0.72 0.00 sd 4.00 2.64 0.00 CC mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 CL mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 QC mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 CP mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 IM mean 0.00 0.00 0.16 sd 0.00 0.00 0.80 LL mean 0.00 • 0.00 0.00 sd 0.00 0.00 0.00 MO mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 TM mean 0.00 0.00 0.00 sd 0.00 0.00 0.00 Asteraceae Asteraceae Asteraceae Asteraceae Betul Aitemesia Aster Cirsium Solidaqo Alnus 0.16 0.00 0.52 0.60 0.00 0.80 0.00 1.61 3.00 0.00 0.00 5.40 0.00 0.00 0.00 0.00 6.75 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.16 0.00 0.00 0.00 0.00 0.80 0,00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 VO (Jl A p p e n d i x 7 B . SHRUB LAYER ( c o n t i n u e d ) . Location Caprifbliaceae Capriloliaceae Celastraceae Cruciferae Cupressaceae Cyperaceae Elaeagnaceae Ericaceae Lonicera Linnaea Pachistima Arabis Juniperus Carex Sheppherdia Menziesia BD mean 0.00 0.00 0.00 0.12 0.00 0.16 0.00 0.00 sd 0.00 0.00 0.00 0.60 0.00 0.80 0.00 0.00 CC mean 8.90 2.00 25.40 0.00 0.00 0.00 0.00 0.00 sd 7.58 6.32 8.86 0.00 0.00 0.00 0.00 0.00 CL mean 2.93 0.27 11.73 0.00 0.47 0.00 0.00 0.00 sd 5.79 1.03 14.77 0.00 1.81 0.00 0.00 0.00 QC mean 0.00 0.00 0.00 0.00 1.00 0.00 0.00 0.64 sd 0.00 0.00 0.00 0.00 3.54 0.00 0.00 2.29 CP mean 0.00 0.00 0.00 0.00 14.60 0.00 0.00 9.50 sd 0.00 0.00 0.00 0.00 18.08 0.00 0.00 15.71 IM mean 0.00 0.00 4.52 0.00 12.44 0.00 0.16 0.00 sd 0.00 0.00 7.19 0.00 12.00 0.00 0.80 0.00 LL mean 0.00 0.00 0.00 0.00 14.10 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 8.95 0.00 0.00 0.00 MO mean 0.00 0.00 0.00 0.00 0.00 0.00 - 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 TM mean 0.00 0.00 0.80 0.00 1.76 0.00 0.00 0.00 sd 0.00 0.00 4.00 0.00 5.17 0.00 0.00 0.00 A p p e n d i x 7 B . SHRUB L A Y E R ( c o n t i n u e d ) . Ericaceae Ericaceae Ericaceae Fabaceae Fabaceae Fabaceae Grossulariaceae Lilliace Location Rhododendron Phvllodoce Vaccinium Lupinus Trifolium Vicia Ribes Veratri BD mean 0.00 0.00 0.00 2.40 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 3.7,7 0.00 0.00 0.00 0.00 CC mean 0.00 0.00 0.00 0.60 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 1.90 0.00 0.00 0.00 0.00 CL mean 6.80 0.00 0.00 0.00 0.00 0.00 0.27 0.00 sd 14.82 0.00 0.00 0.00 0.00 0.00 1.03 0.00 QC mean 0.00 0.00 0.00 ,0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 CP mean 0.00 0.00 0.00 0.00 0.40 0.00 0.00 3.50 sd 0.00 0.00 0.00 0.00 1.26 0.00 0.00 11.07 IM mean 0.00 0.00 1.60 3.84 0.00 1.24 2.84 0.00 sd 0.00 0.00 5.17 7.24 0.00 6.20 3.97 0.00 LL mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 MO mean 0.00 0.00 0.00 0.00 0.00 0.00 0.48 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 1.66 0.00 TM mean 0.00 4.16 0.00 1.32 0.00 0.00 0.00 0.00 sd 0.00 13.65 0.00 2.84 0.00 0.00 0.00 0.00 A p p e n d i x 7 B . SHRUB LAYER ( c o n t i n u e d ) . Location BD mean sd CC mean sd CL mean sd QC mean sd CP mean sd IM mean sd LL mean sd MO mean sd TM mean sd Onagraceae Pinaceae Epilobium Abies 0.00 0.88 0.00 4.00 0.00 0.40 0.00 1.26 0.00 6.13 0.00 9.76 0.00 12.48 0.00 14.80 0.00 1.50 0.00 4.74 0.40 0.00 1.41 0.00 0.00 7.40 0.00 12.75 0.00 1.80 0.00 3.32 0.00 8.12 0.00 10.67 Pinaceae Pinaceae Picea Pinus 0.00 0.40 0.00 1.15 1.20 0.00 2.70 0.00 0.27 0.67 1.03 1.80 1.64 0.16 4.33 0.80 0.70 0.00 2.21 0.00 0.00 0.48 0.00 1.33 0.00 1.10 0.00 2.42 0.00 0.28 0.00 1.40 0.32 0.56 1.11 1.58 Pinaceae Poaceae Poaceae Poaceae Poac Pseudotsuga Agropyron Bromus Calamagrostis Poa 0.00 1.60 5.12 0.24 0.16 0.00 3.00 16.74 1.20 0.80 3.10 0.00 0.00 0.00 0.00 8.49 0.00 0.00 0.00 0.00 1.27 0.00 0.00 0.00 0.00 2.79 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.16 0.16 0.00 0.00 0.16 0.80 0.80 0.00 0.00 0.80 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 CO A p p e n d i x 7B. SHRUB L A Y E R ( c o n t i n u e d ) . Poaceae Poaceae Polygonaceae Ranunculaceae Rosaceae Rosaceae Rosaceae Rosaceae Rosa Location Stjpa Unidentified Rumex Anemone Amelanchier Potentilla Rosa Rubus Spire BD mean 0.40 0.00 0.40 0.00 7.84 2.08 2.24 0.00 0.00 sd 2.00 0.00 2.00 0.00 10.26 6.47 4.69 0.00 0.00 CC mean 0.00 4.50 0.00 0.00 10.90 0.00 0.00 0.00 0.00 sd 0.00 11.17 0.00 0.00 9.99 0.00 0.00 0.00 0.00 CL mean 0.00 0.00 0.00 0.00 1.73 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 3.13 0.00 0.00 0.00 0.00 QC mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 CP mean 0.00 0.00 0.00 0.00 0.00 1.80 0.00 0.70 0.00 sd 0.00 0.00 0.00 0.00 0.00 5.69 0.00 2.21 0.00 IM mean 0.00 0.00 0.00 0.00 3.00 0.16 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 4.40 0.80 0.00 0.00 0.00 LL mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 MO mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.24 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.20 TM mean 0.00 000 0.00 1.32 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 000 5.06 0.00 0.00 0.00 0.00 0.00 U3 A p p e n d i x 7 B . SHRUB L A Y E R ( c o n t i n u e d ) . Rosaceae Salicaceae Salicaceae Scrophulariaceae Scrophulariaceae Scrophulariaceae Scroj Location Sorbus Populus Salix Caslilleja Pedicularis Penstemon Verb. BD mean 0.00 1.24 0.00 0.00 0.16 0.16 0.24 sd 0.00 4.81 0.00 0.00 0.80 0.80 0.88 CC mean 1.30 0.00 0.00 0.00 0.00 0.00 0.00 sd 4.11 0.00 0.00 0.00 0.00 0.00 0.00 CL mean 1.40 0.00 1.07 0.00 0.00 0.00 0.00 sd 3.50 0.00 2.25 0.00 0.00 0.00 0.00 QC mean 0.00 0.00 0.76 0.00 0.00 0.00 0.00 sd 0.00 0.00 3.80 0.00 0.00 0.00 0.00 CP mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 IM mean 0.00 0.16 1.08 0.48, 0.16 0.00 0.00 sd 0.00 0.80 2.94 1.33 0.80 0.00 0.00 LL mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 MO mean 1.04 0.00 0.00 0.00 0.00 0.00 0.00 sd 3.88 0.00 0.00 0.00 0.00 0.00 0.00 TM mean 0.00 0.00 0.00 0.00 0.72 0.00 0.00 sd 0.00 0.00 0.00 0.00 2.23 0.00 0.00 o o A p p e n d i x 7B. SHRUB L A Y E R ( c o n t i n u e d ) Valerianaceae Violaceae Location Valeriana Viola BD mean 0.00 0.00 sd 0.00 0.00 CC mean 0.00 0.00 sd 0.00 0.00 CL mean 0.00 0.00 sd 0.00 0.00 QC mean 0.60 0.00 sd 3.00 0.00 CP mean 0.00 0.00 sd 0.00 0.00 IM mean 0.00 0.00 sd 0.00 0.00 LL mean 0.00 0.00 sd 0.00 0.00 MO mean 0.00 0.00 sd 0.00 0.00 TM mean 7.08 1.40 sd . 12.23 3.96 Unvegetate Unvegetated Bedrock Gravel/debris/soil Unvegetated Stumps/deadfall Unvegetated Large rocks 0.80 0.00 0.08 0.00 3.12 ' 0.00 0.40 .0.00 0.00 0.00 - 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 3.80 3.40 0.00 0.00 6.90 11.61 0.00 0.00 1.40 8.68 0.00 0.00 5.02 12.19 0.00 0.00 0.00 22.00 0.00 0.00 0.00 33.35 1.56 1.80 1.92 0.00 5.82 6.60 5.12 0.00 0.00 0.00 0.60 13.00 0.00 .0.00 1.90 8.14 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 2.84 1.28 0.00 0.00 6.03 3.55 A p p e n d i x 7 C . T h e p e r c e n t c o v e r o f p l a n t g e n e r a a n d u n v e g e t a t e d c a t e g o r i e s : T R E E L A Y E R . L o c a t i o n s : B r o o k m e r e Dump ( B D ) , C o l d s p r i n g C a m p g r o u n d ( C C ) , C a s c a d e L o o k o u t ( C L ) , C a t h e d r a l P a r k S l o p e ( C P ) , I r o n M o u n t a i n ( I M ) , L a d y s l i p p e r L a k e ( L L ) , M o u n t O u t r a m ( M O ) , Q u i n i s c o e C a m p g r o u n d (QC) , T h y n n e M o u n t a i n ( T M ) . Betulaceae Pinaceae Pinaceae Pinaceae Pinaceae Pinaceae Rosaceae Salicaceae Salicaceae Location Alnus Abies Picea Pinus Pseudotsuqa Tsuga Sorbus Populus Salix BD mean 0.00 2.40 0.00 15.40 0.32 0.00 0.00 1.36 0.00 sd 0.00 6.06 0.00 16.39 1.60 0.00 0.00 4.42 0.00 CC mean 0.00 9.40 . 6.00 25:80 9.50 3.50 0.00 0.00 0.00 sd 0.00 9.90 15.78 22.38 17.64 11.07 0.00 0.00 0.00 CL mean 0.00 13.60 0.00 3.67 10.07 0.00 0.33 0.00 0.00 sd 0.00 13.60 0.00 3.67 10.07 o:oo 0.33 0.00 0.00 QC mean 0.00 14.52 20.44 5.04 1.00 0.00 0.00 0.00 0.00 sd 0.00 17.56 18.13 10.31 5.00 0.00 0.00 0.00 0.00 CP mean 0.00 7.50 5.20 14.30 0.00 0.00 0.00 0.00 0.00 sd 0.00 7.50 5.20 14.30 0.00 0.00 0.00 0.00 0.00 IM mean 0.68 2.20 0.64 23.60 0.60 0.00 0.00 0.16 0.72 sd 3.40 6.78 1.89 28.95 3.00 0.00 0.00 0.80 3.60 LL mean 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 MO mean 0.00 7.68 0.00 0.40 0.00 0.00 0.48 0.00 0.00 sd 0.00 18.72 0.00 2.00 0.00 0.00 2.40 0.00 0.00 TM mean 0.00 9.04 0.64 0.00 0.00 0.00 0.00 0.00 0.00 sd 0.00 16.25 2.29 0.00 0.00 0.00 0.00 0.00 0.00 

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