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UBC Theses and Dissertations

Alpine melanism in the butterfly Parnassius phoebus F. (Lepidoptera : Papilionidae) Guppy, Crispin Spencer 1984

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ALPINE MELANISM IN THE BUTTERFLY PARNASSIUS PHOEBUS F. (LEPIDOPTERA:PAPILIONIDAE) by CRISPIN SPENCER GUPPY B.Sc.,' U n i v e r s i t y of B r i t i s h C o l u m b i a , 1981 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE i n THE FACULTY OF GRADUATE STUDIES (Department of Z o o l o g y ) We a c c e p t t h i s t h e s i s a s c o n f o r m i n g t o t h e r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA November 1984 (g) C r i s p i n S p e n c e r Guppy, 1984 I n p r e s e n t i n g t h i s t h e s i s i n p a r t i a l f u l f i l m e n t o f t h e r e q u i r e m e n t s f o r an a d v a n c e d d e g r e e a t t h e 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 , I a g r e e t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r r e f e r e n c e and s t u d y . I f u r t h e r a g r e e t h a t p e r m i s s i o n f o r e x t e n s i v e c o p y i n g o f t h i s t h e s i s f o r s c h o l a r l y p u r p o s e s may be g r a n t e d by t h e h e a d o f my d e p a r t m e n t o r by h i s o r h e r r e p r e s e n t a t i v e s . I t i s u n d e r s t o o d t h a t c o p y i n g o r p u b l i c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l g a i n s h a l l n o t be a l l o w e d w i t h o u t my w r i t t e n p e r m i s s i o n . D e p a r t m e n t Of Zoology  The 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 / 1956 Main Mall V a n c o u v e r , Canada V6T 1Y3 D a t e Dec. 19, 1984 DE-6 (3/81) i i ABSTRACT A l p i n e m e l a n i s m , a t e n d e n c y t o become d a r k e r w i t h i n c r e a s e d e l e v a t i o n and l a t i t u d e , o c c u r s i n many s p e c i e s of b u t t e r f l i e s . In P a r n a s s i u s phoebus ( P a p i 1 i o n i d a e ) i t has been assumed t h a t a l p i n e m e l a n i s m i s t h e r e s u l t of a d a p t a t i o n t o c o l d e r e n v i r o n m e n t s a t h i g h e r e l e v a t i o n s and l a t i t u d e s , owing t o t h e a b i l i t y of d a r k e r wings t o t r a n s f e r more s o l a r e n e r g y t o t h e body as h e a t . I examined f i v e i n d e p e n d e n t l y v a r i a b l e components of wing m e l a n i s m f o r agreement w i t h t h i s h y p o t h e s i s , as w e l l as t h e s u b s i d i a r y h y p o t h e s i s t h a t s i z e i s i n v e r s e l y c o r r e l a t e d t o m e l a n i s m b e c a u s e i t a l s o a f f e c t s t h e r m o r e g u l a t i o n . B a s a l p a t c h b l a c k n e s s ( p r o p o r t i o n o f s c a l e s w h i c h a r e b l a c k ) i n b o t h sexes- i s w e l l c o r r e l a t e d w i t h e l e v a t i o n , l a t i t u d e , and a i r t e m p e r a t u r e d u r i n g t h e f l i g h t s e a s o n . In m a l e s , i n c r e a s e d b a s a l p a t c h b l a c k n e s s i s a d v a n t a g e o u s b e c a u s e i t i n c r e a s e s t h e p r o p o r t i o n of t o t a l t i m e s p e n t i n f l i g h t l o o k i n g f o r mates, but i s l o c a l l y d i s a d v a n t a g e o u s b e c a u s e i t i n c r e a s e s t h e p r o b a b i l i t y of d i s p e r s a l . B a s a l p a t c h w i d t h i s o n l y p o o r l y c o r r e l a t e d w i t h e l e v a t i o n , l a t i t u d e , and a i r t e m p e r a t u r e d u r i n g t h e f l i g h t s e a s o n i n m a l e s , and n o t c o r r e l a t e d i n f e m a l e s . I n c r e a s e d p a t c h w i d t h a l s o r e s u l t s i n i n c r e a s e d f l i g h t a c t i v i t y , b u t do e s n o t a f f e c t d i s p e r s a l . B a s a l wing t r a n s p a r e n c y has l i t t l e c o r r e l a t i o n w i t h e l e v a t i o n , and no e f f e c t on a c t i v i t y , a l t h o u g h i t d oes s l i g h t l y a f f e c t body t e m p e r a t u r e . W i t h i n i n d i v i d u a l m o u n t a i n s , d i s t a l wing b l a c k n e s s and t r a n s p a r e n c y a r e c o r r e l a t e d w i t h e l e v a t i o n , b u t show no c o r r e l a t i o n when many m o u n t a i n s a r e c o n s i d e r e d t o g e t h e r . D i s t a l wing c h a r a c t e r s have no e f f e c t on f l i g h t a c t i v i t y o r d i s p e r s a l . S i z e i s w e l l c o r r e l a t e d w i t h e l e v a t i o n and l a t i t u d e , but i s l e s s w e l l c o r r e l a t e d w i t h a i r t e m p e r a t u r e . S m a l l e r m a l e s d i s a p p e a r from t h e p o p u l a t i o n more r a p i d l y t h a n l a r g e m a l e s , a p p a r e n t l y t h r o u g h i n c r e a s e d m o r t a l i t y b e c a u s e s m a l l m a l e s do n o t d i s p e r s e more r a p i d l y . F e m a l e s d i s p e r s e more t h a n m a l e s , w i t h l a r g e r f e m a l e s d i s p e r s i n g more t h a n s m a l l e r f e m a l e s . V a r i a t i o n i n b a s a l p a t c h b l a c k n e s s may have a s i m i l a r a d a p t i v e s i g n i f i c a n c e i n b o t h s e x e s , but a l l t h e o t h e r m e l a n i s m c h a r a c t e r s a p p e a r t o have a d i f f e r e n t s i g n i f i c a n c e f o r e a c h s e x . I d e v e l o p a new h y p o t h e s i s f o r t h e mechanism o f t h e e f f e c t of wing c o l o r on t h e r m o r e g u l a t i o n i n d o r s a l b a s k i n g b u t t e r f l i e s . I s u g g e s t t h a t most t r a n s f e r of s o l a r r a d i a t i o n t o t h e body by t h e wings o c c u r s w h i l e i n f l i g h t , r a t h e r t h a n w h i l e d o r s a l b a s k i n g on a s u b s t r a t e . T h i s h y p o t h e s i s i s us e d t o e x p l a i n t h e o b s e r v a t i o n t h a t t h e minimum l e v e l s of a i r t e m p e r a t u r e and s o l a r r a d i a t i o n a t w h i c h f l i g h t c a n o c c u r d o e s not depend on wing c o l o r i n P a r n a s s i u s phoebus, i n c o n t r a s t t o p h e n o t y p e d e p e n d e n t f l i g h t t h r e s h o l d s i n l a t e r a l b a s k i n g C o l i a s . i v TABLE OF CONTENTS ABSTRACT i i L I S T OF TABLES v i i L I S T OF FIGURES v i i i ACKNOWLEDGEMENTS X GENERAL INTRODUCTION 1 PHENOTYPIC VARIATION IN PARNASSIUS PHOEBUS 5 INTRODUCTION 5 MATERIALS AND METHODS 9 P o p u l a t i o n s S a m p l e d 9 Q u a n t i f i c a t i o n o f P h e n o t y p e 12 (1) Basal P a t c h W i d t h 12 (2) B a s a l P a t c h C o l o r 12 (3) D i s t a l F r o n t Wing C o l o r 15 (4) W i n g l e n g t h 16 T r a n s f o r m a t i o n o f t h e D a t a 16 D e t e r m i n a t i o n o f Mean A i r T e m p e r a t u r e s 17 C o r r e l a t i o n s w i t h P h e n o t y p e 18 (1) C o r r e l a t i o n s b e t w e e n P o p u l a t i o n s 18 (2) C o r r e l a t i o n s w i t h i n a P o p u l a t i o n 19 I n h e r i t a n c e o f P h e n o t y p e 19 RESULTS 21 P h e n o t y p e C o r r e l a t i o n w i t h E l e v a t i o n and L a t i t u d e ... 21 P h e n o t y p e C o r r e l a t i o n w i t h A i r T e m p e r a t u r e 21 P h e n o t y p e V a r i a t i o n w i t h i n I n d i v i d u a l M o u n t a i n s 28 V P h e n o t y p e C o r r e l a t i o n s between P o p u l a t i o n s 31 P o p u l a t i o n P h e n o t y p e C o r r e l a t i o n s between Sexes 32 P h e n o t y p e C o r r e l a t i o n s w i t h i n a P o p u l a t i o n 34 P h e n o t y p i c V a r i a t i o n o v e r Time 35 I n h e r i t a n c e of P h e n o t y p e . . 36 B a s a l P a t c h W i d t h 44 B a s a l P a t c h B l a c k n e s s . . 44 B a s a l T r a n s p a r e n c y 44 D i s t a l B l a c k n e s s 49 D i s t a l T r a n s p a r e n c y 49 F r o n t Wing L e n g t h 49 DISCUSSION 59 THE ADAPTIVE SIGNIFICANCE OF ALPINE MELANISM 67 INTRODUCTION 67 MATERIALS AND METHODS 71 S t u d y S i t e s 71 M e l a n i s m and Body T e m p e r a t u r e 72 Minimum Body T e m p e r a t u r e R e q u i r e d f o r F l i g h t , . 74 D e t e r m i n a t i o n o f F l i g h t T h r e s h o l d s 75 E f f e c t of P h e n o t y p e and E n v i r o n m e n t on A c t i v i t y ...... 76 P o p u l a t i o n P a r a m e t e r s 77 D i s p e r s a l 77 RESULTS 7 9 B a s a l M e l a n i s m and Body T e m p e r a t u r e 79 Minimum Body T e m p e r a t u r e f o r F l i g h t 79 Minimum F l i g h t T h r e s h o l d 80 M e l a n i s m and A c t i v i t y 94 M e l a n i s m and S u r v i v a l 103 P o p u l a t i o n P a r a m e t e r s 106 D i s p e r s a l 117 DISCUSSION 122 B a s a l M e l a n i s m and Body T e m p e r a t u r e 122 Minimum Body T e m p e r a t u r e f o r F l i g h t 124 Body T e m p e r a t u r e and t h e Minimum F l i g h t T h r e s h o l d ...124 B a s k i n g C o n d i t i o n s and t h e F l i g h t T h r e s h o l d ...125 B a s k i n g and Body T e m p e r a t u r e 126 F l i g h t and Body T e m p e r a t u r e 130 D e t e c t i n g t h e E f f e c t of B a s a l M e l a n i s m on A c t i v i t y ..131 F e e d i n g 132 S u r v i v a l 134 D i s p e r s a l 135 A d a p t i v e S i g n i f i c a n c e of V a r i a t i o n i n M e l a n i s m and S i z e 138 GENERAL DISCUSSION ..141 LITERATURE CITED 147 APPENDIX 1. P o p u l a t i o n s and S u b p o p u l a t i o n s o f P. phoebus ..153 v i i L I S T OF TABLES T a b l e I . L o c a l i t i e s f o r I n h e r i t a n c e D a t a 20 T a b l e I I . Changes i n Phenotype w i t h E l e v a t i o n and L a t i t u d e 22 T a b l e I I I . Changes i n Phenotype w i t h E l e v a t i o n and A i r T e m p e r a t u r e 27 T a b l e IV. P h e n o t y p e D i f f e r e n c e s between M o u n t a i n s 29 T a b l e V. P h e n o t y p e Changes w i t h i n M o u n t a i n s 30 T a b l e V I . I n t e r p o p u l a t i o n C o r r e l a t i o n s between C h a r a c t e r s . 32 T a b l e V I I . I n t e r p o p u l a t i o n C o r r e l a t i o n s between Sexes 33 T a b l e V I I I . C o r r e l a t i o n s between C h a r a c t e r s w i t h i n I n d i v i d u a l s 35 T a b l e IX. Changes i n P h e n o t y p e w i t h Sex and Y e a r ...» 43 T a b l e X. B a s a l M e l a n i s m and Body T e m p e r a t u r e 80 T a b l e X I . M a l e P o p u l a t i o n P a r a m e t e r s 117 T a b l e X I I . Summary of v a r i a t i o n i n wing p h e n o t y p e and i t s c o r r e l a t i o n s w i t h . a c t i v i t y 142 L I S T OF FIGURES F i g u r e 1. Sample p o p u l a t i o n s of P a r n a s s i u s phoebus 10 F i g u r e 2. P h e n o t y p i c measurements 13 F i g u r e 3. Male b a s a l p a t c h b l a c k n e s s as a f u n c t i o n of e l e v a t i o n and a i r t e m p e r a t u r e 23 F i g u r e 4. Female b a s a l p a t c h b l a c k n e s s as a f u n c t i o n of e l e v a t i o n and a i r t e m p e r a t u r e . 25 F i g u r e 5. S e a s o n a l v a r i a t i o n i n male b a s a l p a t c h w i d t h .... 37 F i g u r e 6. S e a s o n a l v a r i a t i o n i n male b a s a l p a t c h b l a c k n e s s 38 F i g u r e 7. S e a s o n a l v a r i a t i o n i n male b a s a l t r a n s p a r e n c y ... 39 F i g u r e 8. S e a s o n a l v a r i a t i o n i n male d i s t a l b l a c k n e s s 40 F i g u r e 9. S e a s o n a l v a r i a t i o n i n male d i s t a l t r a n s p a r e n c y .. 41 F i g u r e 10. S e a s o n a l v a r i a t i o n i n male f r o n t wing l e n g t h ... 42 F i g u r e 11. I n h e r i t a n c e of b a s a l p a t c h w i d t h 45 F i g u r e 12. I n h e r i t a n c e o f b a s a l p a t c h b l a c k n e s s 47 F i g u r e 13. I n h e r i t a n c e o f b a s a l t r a n s p a r e n c y 50 F i g u r e 14. I n h e r i t a n c e o f d i s t a l b l a c k n e s s 52 F i g u r e 15. I n h e r i t a n c e o f d i s t a l t r a n s p a r e n c y 54 F i g u r e 16. I n h e r i t a n c e o f f r o n t wing l e n g t h 56 F i g u r e 17. Ma l e minimum f l i g h t t h r e s h o l d 82 F i g u r e 18. Female minimum f l i g h t t h r e s h o l d 84 F i g u r e 19. P r e d i c t e d minimum f l i g h t t h r e s h o l d s 86 F i g u r e 20. E f f e c t of b a s a l p a t c h b l a c k n e s s on f l i g h t t h r e s h o l d 88 F i g u r e 21. E f f e c t o f b a s a l p a t c h w i d t h on f l i g h t t h r e s h o l d 90 F i g u r e 22. Combined e f f e c t of b a s a l p a t c h w i d t h and b l a c k n e s s on f l i g h t t h r e s h o l d 92 F i g u r e 23. P r o p o r t i o n o f t i m e m a l e s spend i n f l i g h t 95 F i g u r e 24. P r o p o r t i o n of t i m e s p e n t f e e d i n g 98 F i g u r e 25. P r o p o r t i o n of t i m e s p e n t b a s k i n g 100 F i g u r e 26. F r e q u e n c y d i s t r i b u t i o n o f r e s i d e n c e t i m e 104 F i g u r e 27. C u m u l a t i v e f r e q u e n c y d i s t r i b u t i o n of r e s i d e n c e t i m e 107 F i g u r e 28. B a s a l p a t c h b l a c k n e s s and r e s i d e n c e t i m e 109 F i g u r e 29. B a s a l p a t c h w i d t h and r e s i d e n c e t i m e 111 F i g u r e 30. B a s a l p a t c h w i d t h and b l a c k n e s s and r e s i d e n c e t ime 113 F i g u r e 31. F r o n t wing l e n g t h and r e s i d e n c e t i m e 115 F i g u r e 32. D i s p e r s a l ,..118 X ACKNOWLEGEMENTS T h i s r e s e a r c h would not have been p o s s i b l e w i t h o u t t h e s u p p o r t o f my g r a d u a t e s u p e r v i s o r , D r . J u d i t h M y e r s , t h r o u g h o u t my r e s e a r c h p r o g r a m . I thank my f e l l o w o f f i c e i n m a t e s , D r . K a t h y W i l l i a m s and R o b e r t P o w e l l , f o r t h e i r s u p p o r t and a d v i c e . W i t h o u t t h e i r f r i e n d l y e n couragement my y e a r s a s a g r a d u a t e s t u d e n t would have been much l e s s r e w a r d i n g . S p e c i a l a p p r e c i a t i o n goes t o Mr. D a v i d L. T h r e a t f u l who l e d me t o many of t h e P a r n a s s i u s phoebus c o l o n i e s used, i n t h i s s t u d y , and a s s i s t e d i n s a m p l i n g some of them. I a l s o w i s h t o thank Mr. A.B. C a m p b e l l , R e g i o n a l D i r e c t o r o f t h e P a r k s and O u t d o o r R e c r e a t i o n D i v i s i o n (Thompson-Okanagan R e g i o n ) of t h e B.C. M i n i s t r y of L a n d s , P a r k s , and H o u s i n g . He g e n e r o u s l y gave p e r m i s s i o n • t o sample P a r n a s s i u s c o l o n i e s i n S i l v e r S t a r P r o v i n c i a l R e c r e a t i o n A r e a and 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 , and a l w a y s r e p l i e d p r o m p t l y and c o u r t e o u s l y t o my i n q u i r i e s . I am g r a t e f u l t o t h e s t a f f , p a s t and p r e s e n t , of t h e B i o s c i e n c e s D a t a C e n t e r a t U.B.C. F o r t h e i r p a t i e n c e and t h e i n s t r u c t i o n w h i c h was n e c e s s a r y f o r t h e e x t e n s i v e d a t a a n a l y s i s u s e d i n t h i s s t u d y . S h i r l e y G r a y s t o n e , Susan E r t i s , and Dave Z i t t i n were e s p e c i a l l y h e l p f u l . 1 GENERAL INTRODUCTION Most b u t t e r f l i e s have complex wing c o l o r p a t t e r n s formed from many component c h a r a c t e r s , e a c h of w h i c h may v a r y c o n t i n u o u s l y or d i s c r e t e l y . I f t h e same s e l e c t i o n p r e s s u r e i s a c t i n g on t h e e n t i r e wing p a t t e r n , v a r i a t i o n i n e a c h component of t h e p h e n o t y p e w i l l t e n d t o c o v a r y between p o p u l a t i o n s . However, L e p i d o p t e r a have many s e l e c t i o n p r e s s u r e s a c t i n g on them, and a l s o have a l a r g e number of chromosomes ( F o r d , 1937). I t i s t h e r e f o r e l i k e l y t h a t a l t h o u g h some of t h e v a r i a t i o n i n wing p a t t e r n components may be c o r r e l a t e d , much of i t w i l l n ot show any c o r r e l a t i o n , a s i n E u p h y d r y a s e d i t h a ( L e p i d o p t e r a : N y m p h a l i d a e ) (Mason e t a_l. , 1968) and Heteronympha merope ( L e p i d o p t e r a : S a t y r i n a e ) ( P e a r s e and M u r r a y , 1981, 1982). The o v e r a l l wing p h e n o t y p e w i l l be t h e r e s u l t of many s e l e c t i v e p r e s s u r e s a c t i n g t o g e t h e r ( D o u g l a s , 1978). To u n d e r s t a n d t h e a d a p t i v e s i g n i f i c a n c e of p h e n o t y p i c v a r i a t i o n i n wing c o l o r a t i o n , t h e s e l e c t i o n p r e s s u r e s a c t i n g on b o t h i n d i v i d u a l components and t h e o v e r a l l wing p a t t e r n must be d e t e r m i n e d . S i n c e a c h a r a c t e r may be i n f l u e n c e d by s e v e r a l d i f f e r e n t s o u r c e s o f s e l e c t i o n , o r t h e a d a p t i v e s i g n i f i c a n c e may change under d i f f e r e n t c o n d i t i o n s , t h e ' o p t i m a l ' p h e n o t y p e w i l l be a compromise w h i c h r e s u l t s i n t h e g r e a t e s t o v e r a l l f i t n e s s . The r e l a t i v e s t r e n g t h of e a c h s e l e c t i o n p r e s s u r e w i l l p r o b a b l y v a r y b o t h t e m p o r a l l y and s p a t i a l l y , and t h e r e f o r e t h e p h e n o t y p e components w i l l a l s o v a r y between p o p u l a t i o n s and o v e r t i m e . S e l e c t i o n p r e s s u r e s may v a r y w i t h i n one g e n e r a t i o n , c a u s i n g h i g h 2 v a r i a b i l i t y w i t h i n p o p u l a t i o n s . When g e o g r a p h i c p a t t e r n s of v a r i a t i o n o c c u r , i t i s f r e q u e n t l y p o s s i b l e t o d e v e l o p h y p o t h e s e s as t o the a d a p t i v e s i g n i f i c a n c e of t h e v a r i a t i o n (Mayr, 1956). One p a r t i c u l a r l y n o t i c e a b l e p a t t e r n of v a r i a t i o n i n i n s e c t s i s t h e t e n d e n c y f o r a s p e c i e s t o be d a r k e r a t h i g h e r e l e v a t i o n s and l a t i t u d e s (Benham e t a l . , 1974; D o u g l a s and G r u l a , 1978; Downes, 1964,1965; Freeman, 1958; H o v a n i t z , 1941,1950; K e t t l e w e l l , 1973; M a n i , 1962,1968; R o l a n d 1978,1982; Watt, 1968), w h i c h i s t e rmed ' a l p i n e m e l a n i s m ' . S i n c e a i r t e m p e r a t u r e d e c r e a s e s w i t h i n c r e a s e d e l e v a t i o n and l a t i t u d e , a l p i n e m e l a n i s m has g e n e r a l l y been i n t e r p r e t e d as b e i n g r e l a t e d t o t h e r m o r e g u l a t i o n . The b u t t e r f l y genus C o l i a s e x h i b i t s a l p i n e m e l a n i s m ( R o l a n d , 1978; Watt, 1968), and v a r i a t i o n i n m e l a n i s m i n t h i s genus has been shown t o a f f e c t body t e m p e r a t u r e , f l i g h t a c t i v i t y , and o v i p o s i t i o n r a t e ( D o u g l a s , 1978; K i n g s o l v e r , 1983; R o l a n d , 1982; Watt, 1969). Thus a l p i n e m e l a n i s m i n C o l i a s can more p r e c i s e l y be t e r med ' t h e r m a l a l p i n e m e l a n i s m ' , t o s t r e s s t h e p o s s i b i l i t y t h a t o t h e r mechanisms c o u l d a l s o r e s u l t i n a g e n e r a l i n c r e a s e i n m e l a n i s m w i t h i n c r e a s e d e l e v a t i o n and l a t i t u d e . The b a s k i n g p o s t u r e of a b u t t e r f l y d e t e r m i n e s w h i c h s u r f a c e of t h e wings a f f e c t s body t e m p e r a t u r e t h r o u g h t h e a b s o r p t i o n o f s o l a r r a d i a t i o n . The f o u r most f r e q u e n t l y u s e d c a t a g o r i e s o f b a s k i n g p o s t u r e ( C l e n c h , 1966) a r e l a t e r a l b a s k i n g ( v e n t r a l wing s u r f a c e p e r p e n d i c u l a r t o s o l a r r a d i a t i o n ) , d o r s a l b a s k i n g ( d o r s a l wing s u r f a c e p e r p e n d i c u l a r t o s o l a r r a d i a t i o n ) , v e r t i c a l 3 b a s k i n g (wing s u r f a c e s c l o s e d d o r s a l l y and o r i e n t e d p a r a l l e l t o s o l a r r a d i a t i o n ) , and body b a s k i n g (wing s u r f a c e s p a r a l l e l t o s o l a r r a d i a t i o n but open s u f f i c i e n t l y t o a l l o w t h e body dorsum t o be i l l u m i n a t e d ) . The c o l o r o f t h e wing s u r f a c e s p e r p e n d i c u l a r t o s o l a r r a d i a t i o n w i l l have a much g r e a t e r e f f e c t on body t e m p e r a t u r e t h a n v a r i a t i o n i n c o l o r of t h e o t h e r wing s u r f a c e s . T h e r e f o r e b a s k i n g p o s t u r e i s one f a c t o r w h i c h w i l l d e t e r m i n e whether v a r i a t i o n i n a wing p a t t e r n component a f f e c t s body t e m p e r a t u r e . C o l i a s have a v e r y s i m p l e wing p a t t e r n , p a r t i c u l a r i l y on t h e v e n t r a l ( b a s k i n g ) s u r f a c e . A l p i n e m e l a n i s m o c c u r s i n o t h e r b u t t e r f l y s p e c i e s ( D o u g l a s and G r u l a , 1978; Downes, 1965; Freeman, 1958; H o v a n i t z , 1941; K e t t l e w e l l , 1973; M a n i , 1962,1968), many of w h i c h have a more complex wing p a t t e r n on t h e b a s k i n g s u r f a c e t h a n C o l i a s . In d o r s a l b a s k i n g b u t t e r f l i e s most of t h e d i s t a l wing a r e a i s t o o f a r from t h e body t o a f f e c t body t e m p e r a t u r e when t h e b u t t e r f l y i s s u s p e n d e d i n m i d - a i r ( D o u g l a s , 1978), and t h e r e may n o t be any e f f e c t o f t h e wings on t h e b u t t e r f l y ' s body t e m p e r a t u r e when i t i s d o r s a l b a s k i n g on a s u b s t r a t e ( H e i n r i c h , 1972; Kammer and B r a c c h i , 1972). N e v e r t h e l e s s , a l p i n e m e l a n i s m i s e v i d e n t i n b o t h t h e b a s a l and d i s t a l wing a r e a s of some d o r s a l b a s k i n g b u t t e r f l i e s , s u g g e s t i n g t h a t t h e r e may be o t h e r , n o n - t h e r m o r e g u l a t o r y , r e a s o n s f o r v a r i a t i o n i n m e l a n i s m . My s t u d y i s an a t t e m p t t o d e t e r m i n e : ( l ) t h e p a t t e r n s of v a r i a t i o n i n m e l a n i s m i n a d o r s a l b a s k i n g b u t t e r f l y , and whether t h e y f i t t h e p a t t e r n of a l p i n e m e l a n i s m , ( 2 ) w h e t h e r t h e 4 v a r i a t i o n i s due t o g e n e t i c or e n v i r o n m e n t a l d i f f e r e n c e s , , ( 3 ) w h e t h e r t h e r m a l e x p l a n a t i o n s a r e s u f f i c i e n t t o a c c o u n t f o r a l p i n e m e l a n i s m i n a d o r s a l b a s k i n g b u t t e r f l y , ( 4 ) w h e t h e r t h o s e c h a r a c t e r s w h i c h f o l l o w t h e p a t t e r n of t h e r m a l a l p i n e m e l a n i s m , have a m e a s u r a b l e t h e r m o r e g u l a t o r y a d a p t i v e s i g n i f i c a n c e , and ( 5 ) w h e t h e r t h o s e c h a r a c t e r s w h i c h do n o t f o l l o w t h e p a t t e r n of t h e r m a l a l p i n e m e l a n ism, a l s o do n o t have a m e a s u r a b l e t h e r m o r e g u l a t o r y a d a p t i v e s i g n i f i c a n c e . 5 PHENOTYPIC VARIATION IN PARNASSIUS PHOEBUS INTRODUCTION The b u t t e r f l y P a r n a s s i u s phoebus F a b r i c i u s ( P a p i l i o n i d a e ) i s a v i d l y s o u g h t by c o l l e c t o r s , p r i m a r i l y owing t o i t s immense v a r i a b i l i t y i n m e l a n i s m ( F e r r i s , 1976; S h e p a r d and S h e p a r d , i n Howe 1975). D e s p i t e t h i s p o p u l a r i t y , knowledge o f t h e s p e c i e s ' b i o l o g y i s l i m i t e d t o a s t u d y of i t s p o p u l a t i o n b i o l o g y i n C o l o r a d o by S c o t t ( 1 9 7 3 a ) , a r a t h e r l i m i t e d s t u d y of s p o t v a r i a t i o n and p o p u l a t i o n d y n a m i c s of t h r e e p o p u l a t i o n s i n C o l o r a d o by C l i f f o r d ( 1 9 8 1 ) , and a few s h o r t comments i n o t h e r s o u r c e s . D e s c r i p t i o n s of v a r i a t i o n i n m e l a n i s m i n P. phoebus ( F e r r i s , 1976; S h e p a r d and S h e p a r d , i n Howe 1975) i n d i c a t e t h a t i t p r o b a b l y f o l l o w s t h e p a t t e r n o f a l p i n e m e l a n i s m , a s do most o t h e r s p e c i e s o f P a r n a s s i u s (Mani, 1962,1968). O n l y e l e v a t i o n a l g r a d i e n t s i n m e l a n i s m , and not l a t i t u d i n a l , have been n o t e d f o r t h i s genus. In s o u t h e r n B r i t i s h C o l u m b i a P. phoebus has been f o u n d f r o m as low as 61Om t o as h i g h a s 2400m, and i n C o l o r a d o i t o c c u r s up t o a t l e a s t 4000m e l e v a t i o n . The s p e c i e s i s f o u n d f r o m n o r t h e r n M e x i c o t o n o r t h e r n A l a s k a i n N o r t h A m e r i c a , and a c r o s s much o f E u r a s i a . T h i s s p e c i e s has a f a r g r e a t e r a l t i t u d i n a l and l a t i t u d i n a l r ange t h a n any s i n g l e s p e c i e s of t h e o n l y o t h e r genus ( C o l i a s spp.) f o r which a l p i n e m e l a n i s m has been q u a n t i f i e d ( R o l a n d , 1978). Such a wide d i s t r i b u t i o n , c o m b i n e d w i t h a p p a r e n t l y low r a t e s o f d i s p e r s a l ( S c o t t , 1973a) make P. phoebus an a l m o s t i d e a l c h o i c e i n w h i c h t o q u a n t i f y and 6 i n v e s t i g a t e t h e a d a p t i v e s i g n i f i c a n c e of a l p i n e m e l a n i s m . A g e n e r a l d e s c r i p t i o n of p h e n o t y p i c v a r i a t i o n i n P a r n a s s i u s  p hoebus i s g i v e n by F e r r i s ( 1 9 7 6 ) : " T h r o u g h o u t t h e Rocky M t s . , l o w - e l e v a t i o n phoebus t e n d t o be much l a r g e r and more b r i g h t l y marked t h a n s p e c i m e n s t a k e n a t h i g h e l e v a t i o n s . W i t h i n c r e a s i n g a l t i t u d e , t h e f e m a l e s become s m a l l e r and d a r k e r , w i t h t h e g r o u n d c o l o r of t h e w i n g s t e n d i n g t o w a r d s c h a r c o a l g r e y . On t h e o t h e r hand, t h e m a l e s become s m a l l e r and more l i g h t l y marked. T h i s change i n c o l o r and m a c u l a t i o n i s e a s i l y o b s e r v e d i n t h e c o n t i n u o u s c l i n e f r o m t h e L a r a m i e P l a i n s ( c a . 7,000') t h r o u g h t h e T r a n s i t i o n Zone F o o t h i l l s ( c a . 8,000-9,000') and i n t o t h e Snowy Range Mts., A l b a n y Co., Wyoming. In t h i s r e g i o n , t h e r e i s a c o n t i n u o u s band of Sedum l a n c e o l a t u m t h e h o s t p l a n t , and t h e r e a r e no b a r r i e r s t h a t p r o d u c e i s o l a t e d c o l o n i e s . " W i t h o u t g i v i n g f u r t h e r e v i d e n c e , he c o n c l u d e d t h a t : "The r e a s o n f o r t h e d a r k c o l o r i n t h e f e m a l e s r e l a t e s t o d e v e l o p m e n t a t r e d u c e d t e m p e r a t u r e ( h i g h a l t i t u d e s ) r e s u l t s [ s i c ] i n t h e p r o d u c t i o n of more m e l a n i n and hence d a r k e r t o n e s . T h i s phenomenon i s sometimes c a l l e d t h e r m a l a d a p t a t i o n . " However t h e a d a p t i v e s i g n i f i c a n c e o f v a r i a t i o n i n m e l a n i s m can not be t h e same f o r m a l e s and f e m a l e s i f t h e y do n o t f o l l o w t h e same p a t t e r n of v a r i a t i o n . F u r t h e r m o r e , p r e l i m i n a r y o b s e r v a t i o n s s u g g e s t e d t h a t f e m a l e s become d a r k e r w i t h i n c r e a s e d e l e v a t i o n o n l y on some m o u n t a i n s , w h i c h c a s t f u r t h e r d o u b t on a t h e r m a l c a u s e of m e l a n i s m v a r i a t i o n i n t h i s s p e c i e s . Many b u t t e r f l i e s a l s o t e n d t o become s m a l l e r w i t h i n c r e a s e d e l e v a t i o n and l a t i t u d e ( H o v a n i t z , 1950), w h i c h I t e r m " a l p i n e d w a r f i s m " . F e r r i s ( 1 9 7 6 ) , S h e p a r d and S h e p a r d ( i n Howe, 1975) and o t h e r s a s s o c i a t e i n c r e a s e d f e m a l e m e l a n i s m of P. phoebus w i t h a l p i n e d w a r f i s m . I f a l p i n e d w a r f i s m i s i n d e e d s t r o n g l y 7 c o r r e l a t e d w i t h a l p i n e m e l a n i s m , t h e y may be a c t e d on by t h e same e n v i r o n m e n t a l f a c t o r . F o r example, D o u g l a s (1978) s u g g e s t e d t h a t b o t h i n c r e a s e d b a s a l wing m e l a n i s m and d e c r e a s e d body s i z e ( f o r a l a r g e b u t t e r f l y ) w i l l be t h e r m a l l y a d v a n t a g e o u s . S i n c e t h e r e p o r t e d p a t t e r n s of v a r i a t i o n i n m e l a n i s m and s i z e i n P. phoebus have been q u a l i t a t i v e ( w i t h t h e e x c e p t i o n of C l i f f o r d , 1981, who u s e d r a n k e d v a l u e s ) , I q u a n t i f y some o f t h e v a r i a t i o n p r e s e n t w i t h i n and between p o p u l a t i o n s o v e r a range o f a l t i t u d e s and l a t i t u d e s . The a s s u m p t i o n b e h i n d p r e v i o u s s t u d i e s of a l p i n e m e l a n i s m has been t h a t s i n c e a i r t e m p e r a t u r e d e c r e a s e s w i t h e l e v a t i o n , t h e o b s e r v e d c o r r e l a t i o n between m e l a n i s m and e l e v a t i o n i s r e a l l y an i n v e r s e c o r r e l a t i o n between m e l a n i s m and a i r t e m p e r a t u r e . However, s i n c e t h e a i r t e m p e r a t u r e d u r i n g t h e f l i g h t s e a s o n of a b u t t e r f l y i s o n l y r o u g h l y c o r r e l a t e d w i t h e l e v a t i o n and l a t i t u d e when c o n s i d e r i n g a l a r g e g e o g r a p h i c a r e a , t h e c o r r e l a t i o n o f m e l a n i s m w i t h a i r t e m p e r a t u r e d u r i n g t h e f l i g h t s e a s o n s h o u l d be b e t t e r t h a n t h e c o r r e l a t i o n w i t h e l e v a t i o n and l a t i t u d e . T h i s p r e d i c t i o n has n o t p r e v i o u s l y been t e s t e d . In s t u d i e s o f t h e a d a p t i v e s i g n i f i c a n c e of m e l a n i s m i n C o l i a s , i t was assumed t h a t d i f f e r e n c e s i n m e l a n i s m a r e g e n e t i c a l l y b a s e d , and t h e r e f o r e n a t u r a l s e l e c t i o n c a n o c c u r . No e v i d e n c e has been p u b l i s h e d t o t e s t t h i s a s s u m p t i o n however, and a l l s t a t e m e n t s as t o t h e g e n e t i c b a s i s o f v a r i a t i o n i n m e l a n i s m a r e b a s e d on u n p u b l i s h e d d a t a ( R o l a n d , 1982 [pers.comm. f r o m W.B. W a t t ] ; Watt, 1968 [ u n p u b l i s h e d d a t a ] ; Hoffmann, 1978 8 [ u n p u b l i s h e d d a t a ] ) . Remington (1954) d i s c u s s e d t h e u n q u a n t i f i e d o b s e r v a t i o n of G e r o u l d (1923 and i_n l i t t . ) t h a t t h e o f f s p r i n g o f an a b n o r m a l l y d a r k f e m a l e C o l i a s were a l s o a b n o r m a l l y d a r k . Remington s u g g e s t e d t h a t a s i n g l e dominant a l l e l e was i n v o l v e d i n d e t e r m i n i n g t h e a b n o r m a l d a r k n e s s , however a p p a r e n t l y t h e f e m a l e was an a b e r r a t i o n , r a t h e r t h a n p a r t of t h e n o r m a l v a r i a t i o n i n m e l a n i s m . No e v i d e n c e as t o t h e h e r i t a b i l i t y of v a r i a t i o n i n m e l a n i s m i n P. phoebus has been p r e s e n t e d , a l t h o u g h F e r r i s (1976) b e l i e v e d t h a t i t i s d e t e r m i n e d by d e v e l o p m e n t a l t e m p e r a t u r e s , and J.A. S c o t t (rn l i t t . ) b e l i e v e s t h a t i t i s g e n e t i c a l l y b a s e d . A c c o r d i n g l y , I w i l l a l s o a t t e m p t t o d e m o n s t r a t e whether v a r i a t i o n i n m e l a n i s m and s i z e i n P. phoebus i s due t o g e n e t i c or e n v i r o n m e n t a l d i f f e r e n c e s , o r b o t h . The p u r p o s e of t h i s i n i t i a l e x p l o r a t o r y c h a p t e r i s t o examine v a r i a t i o n i n m e l a n i s m i n P a r n a s s i u s p h o e b u s , under t h e g e n e r a l h y p o t h e s i s t h a t t h e v a r i a t i o n i s t h e r e s u l t of a d a p t a t i o n t o d i f f e r e n t a i r t e m p e r a t u r e s . S i x p h e n o t y p i c c h a r a c t e r s a r e e xamined: t h r e e as components o f o v e r a l l b a s a l wing m e l a n i s m , two as components of d i s t a l w ing m e l a n i s m , and one as a measure of s i z e . A l l c h a r a c t e r s a r e s e p a r a t e l y e v a l u a t e d as t o ( l ) w h e t h e r t h e i r v a r i a t i o n i s c o r r e l a t e d w i t h e l e v a t i o n , ( 2 ) w h e t h e r t h e y a r e b e t t e r c o r r e l a t e d w i t h a i r t e m p e r a t u r e d u r i n g t h e f l i g h t s e a s o n than w i t h e l e v a t i o n , and ( 3 ) w h e t h e r t h e c h a r a c t e r s a r e d e v e l o p m e n t a l l y , g e n e t i c a l l y , and e v o l u t i o n a r i l y i n d e p e n d e n t . I do not a t t e m p t t o a s s e s s t h e i m p o r t a n c e o f any o t h e r e n v i r o n m e n t a l f a c t o r . 9 MATERIALS AND METHODS P o p u l a t i o n s Sampled One h u n d r e d and f i v e p o p u l a t i o n s and s u b p o p u l a t i o n s were sampled t h r o u g h o u t N o r t h A m e r i c a ( F i g u r e 1), a l t h o u g h most o f the s a m p l i n g was done i n 1982 and 1983 i n t h e Okanagan V a l l e y 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 and a d j a c e n t a r e a s . The samples o u t s i d e t h e Thompson and Okanagan V a l l e y s of B.C. were o b t a i n e d by m y s e l f and o t h e r s p r i o r t o 1982. O n l y a r b i t r a r y samples of t h e p o p u l a t i o n s were u s e d . Number of samples p e r p o p u l a t i o n , and number of i n d i v i d u a l s p e r sample, was h i g h l y v a r i a b l e ( A p p e n d i x 1 ) , w i t h a t o t a l of 2480 s p e c i m e n s examined. One p o p u l a t i o n ( V e r n o n H i l l , 1100-1200m) was sampl e d t h r o u g h o u t t h e 1983 f l i g h t s e a s o n . L o c a t i o n and e l e v a t i o n were f i x e d u s i n g 1:50,000 t o p o g r a p h i c a l maps w i t h c o n t o u r i n t e r v a l s of 50 or 100 f e e t . When a p o p u l a t i o n o c c u r r e d o v e r an e l e v a t i o n r a n g e g r e a t e r t h a n 150 m e t e r s , i t was s p l i t i n t o two or more s u b p o p u l a t i o n s a l o n g l i n e s o f low p o p u l a t i o n d e n s i t y . S i m i l a r i l y i f a p o p u l a t i o n o c c u r r e d o v e r more t h a n 0.5 km h o r i z o n t a l l y , o r was b r o k e n by bands of f o r e s t , i t was s p l i t i n t o two o r more s u b p o p u l a t i o n s . A ' p o p u l a t i o n ' was t h e r e f o r e a r b i t r a r i l y d e f i n e d by a r e a s o f a p p a r e n t l y r e d u c e d gene f l o w . P r e c i s e l o c a l i t i e s , e l e v a t i o n s , c o l l e c t i o n d a t e s , e s t i m a t e d f l i g h t s e a s o n s , and sample s i z e s a r e g i v e n i n A p p e n d i x 1. 10 F i g u r e 1. Sample p o p u l a t i o n s of P a r n a s s i u s phoebus. The p o i n t s i n s o u t h e r n B r i t i s h C o l u m b i a may r e p r e s e n t more t h a n one p o p u l a t i o n . See A p p e n d i x 1 f o r l o c a l i t y d e t a i l s . 11 4 2 Q u a n t i f i c a t i o n of P h e n o t y p e Two a r e a s of t h e d o r s a l wing s u r f a c e s were c o n s i d e r e d i n d e t a i l : t h e b a s a l a r e a of t h e h i n d wings and t h e d i s t a l a r e a of t h e f r o n t w i n g s . W i t h i n e a c h a r e a , o v e r a l l m e l a n i s m depends on t h e p r o p o r t i o n o f b l a c k t o w h i t e s c a l e s , and t h e p r o p o r t i o n of s c a l e d t o u n s e a l e d a r e a . The v e n t r a l s u r f a c e c o l o r i s m o s t l y a b l u r r e d v e r s i o n of d o r s a l c o l o r e x c e p t f o r some r e d s c a l i n g a t th e base of t h e h i n d w i n g s . W i n g l e n g t h measurements were made w i t h a m e t r i c r u l e r , a l l o t h e r measurements were made u s i n g a Bausch and Lomb "Zoom" d i s s e c t i n g m i c r o s c o p e w i t h an o p t i c a l m i c r o m e t e r i n one e y e p i e c e and an o p t i c a l g r i d i n t h e o t h e r e y e p i e c e . Measurement '2b' i s a p p r o x i m a t e l y e q u i v a l e n t t o t h e measurement u s e d by R o l a n d (1982) as an i n d e x o f m e l a n i s m . F o r e a c h sex t h e f o l l o w i n g s i x measures of p h e n o t y p e were made: (1) B a s a l P a t c h W i d t h The i n d e x u s e d was t h e p r o p o r t i o n of t h e c e n t e r l i n e o f t h e h i n d wing d i s c a l c e l l w h i c h i s c o v e r e d by t h e p r e d o m i n a n t l y b l a c k a r e a (measured u s i n g an o p t i c a l m i c r o m e t e r , F i g u r e 2 b ) . (2) B a s a l P a t c h C o l o r The d e g r e e of m e l a n i s m w i t h i n t h e b a s a l p a t c h i s d e t e r m i n e d by t h e r e l a t i v e a r e a s c o v e r e d by b l a c k s c a l e s , w h i t e s c a l e s , and b a r e wing membrane. O n l y b a r e wing membrane w h i c h was c o n g e n i t a l , n ot t h a t due t o wear, was c o n s i d e r e d . In t h e b a s a l 1 3 F i g u r e 2. P h e n o t y p i c measurements. The p o s i t i o n o f t h e measurements of p h e n o t y p e u s e d i n t h i s s t u d y . ( a ) D i s t a l wing b l a c k n e s s ; ( b ) B a s a l p a t c h w i d t h ; ( c ) B a s a l p a t c h b l a c k n e s s and t r a n s p a r e n c y . 14 15 a r e a v e r y l i t t l e s c a l e wear o c c u r r e d , even when t h e o u t e r wing a r e a s ' were h e a v i l y worn. An o p t i c a l m i c r o m e t e r was l a i d down t h e c e n t e r l i n e of t h e h i n d wing c e l l , w i t h t h e b a s a l end l i n e d up w i t h v e i n Sc+R, and t h e d i s t a l end about 10% of t h e p a t c h w i d t h i n from t h e o u t e r edge o f t h e p a t c h ( F i g u r e 2 c ) . The 202 p o i n t s d e f i n e d by t h e t i p s o f t h e m i c r o m e t e r c r o s s - b a r s were c l a s s i f i e d as b e i n g o v e r b l a c k , w h i t e , or no s c a l e s . Number of b l a c k + number of w h i t e + number no s c a l e s = 202. ( 2 a ) B a s a l P a t c h T r a n s p a r e n c y - The p r o p o r t i o n of t h e m i c r o m e t e r p o i n t s w h i c h l a y o v e r b a r e wing membrane. ( 2 b ) B a s a l P a t c h B l a c k n e s s - The number of t h e m i c r o m e t e r p o i n t s w h i c h l a y o v e r b l a c k s c a l e s , as a p r o p o r t i o n of a l l p o i n t s o v e r s c a l e s . ( 3 ) D i s t a l F r o n t Wing C o l o r The d i s t a l p a r t of the- f r o n t wing between v e i n s M 3 and C u 2 was u s e d as an i n d e x o f t h e d e g r e e of d i s t a l m e l a n i s m . F o r measurement 3a, an o p t i c a l g r i d was l a i d a c r o s s t h e wing w i t h t h e 2 o u t e r c o r n e r s m e e t i n g t h e wing m a r g i n a t t h e ends o f v e i n s M 3 and C u 2 ( F i g u r e 2 a ) . F o r measurement 3b, t h e m i c r o m e t e r was a l i g n e d p a r a l l e l and a n t e r i o r t o v e i n Cu,, w h i l e t h e s p e c i m e n was a t t h e p o s i t i o n f o r 3a, and t h e m a g n i f i c a t i o n d o u b l e d . M i n o r a d j u s t m e n t s were t h e n made t o p o s i t i o n t h e m i c r o m e t e r a b o u t 2 m i c r o m e t e r w i d t h s a n t e r i o r t o t h e v e i n w h i l e a v o i d i n g any minor s c a l e e r o s i o n , and t h e number of p o i n t s o v e r b a r e wing membrane c o u n t e d . M a j o r s c a l e e r o s i o n was f r e q u e n t l y p r e s e n t , i n w h i c h c a s e t h i s measurement was not made. ;16 ( 3 a ) D i s t a l B l a c k n e s s - The p r o p o r t i o n of t h e 100 q u a d r a t s i n t h e o p t i c a l g r i d i n w h i c h >25% ( m a l e s ) o r >50% ( f e m a l e s ) of t h e s c a l e s were b l a c k . ( 3 b ) D i s t a l T r a n s p a r e n c y - The p r o p o r t i o n o f p o i n t s on t h e o p t i c a l m i c r o m e t e r w h i c h l a y o v e r b a r e wing membrane. ( 4 ) W i n g l e n q t h The l e n g t h of t h e f r o n t wing from t h e t h o r a c i c a t t a c h m e n t p o i n t t o t h e apex of t h e wing. S i n c e b u t t e r f l y w i n g l e n g t h i s w e l l c o r r e l a t e d w i t h t h o r a c i c d i a m e t e r ( D o u g l a s , 1978), t h i s i s a l s o a measure of body s i z e . T r a n s f o r m a t i o n of t h e D a t a B e f o r e any s t a t i s t i c a l a n a l y s i s was done, a l l p h e n o t y p i c measurements, e x c e p t f r o n t wing l e n g t h , were a r c s i n e t r a n s f o r m e d t o n o r m a l i z e t h e d i s t r i b u t i o n s . The v a l u e s were c o n v e r t e d t o p e r c e n t a g e s f o r p r e s e n t a t i o n where p o s s i b l e . The a r c s i n e t r a n s f o r m a t i o n u s e d was t h a t recommended by Zar ( 1 9 7 4 ) : Y = s q r t ( z + . 5 ) a r c s i n ( s q r t ( x + . 3 7 5 ) / ( z + . 7 5 ) ) where x=the raw datum, z=the maximum p o s s i b l e v a l u e f o r x, and Y=the t r a n s f o r m e d p r o p o r t i o n . 17 D e t e r m i n a t i o n of Mean A i r T e m p e r a t u r e s Mean a i r t e m p e r a t u r e s d u r i n g t h e f l i g h t s e a s o n were e s t i m a t e d f o r e a c h s i t e from n e a r b y l o n g - t e r m E n v i r o n m e n t Canada weather d a t a , s h o r t - t e r m B.C. P r o v i n c i a l Government and E n v i r o n m e n t Canada d a t a , and s h o r t - t e r m d a t a c o l l e c t e d by m y s e l f . The B.C. P r o v i n c i a l Government d a t a were f o r weather s t a t i o n s numbers 116121, 116097, 112213, 112212, 112242, 112101, 112102, 112075, 112071, 112077, and 111055 ( M i n i s t r y of E n v i r o n m e n t , 1980). The E n v i r o n m e n t Canada d a t a were f o r 1971-72, 1976-83 from t h e Keremeos, P e n t i c t o n A, E n d e r b y , Salmon Arm, V e r n o n C o l d s t r e a m , V e r n o n S i l v e r s t a r , Okanagan C e n t e r , Mt. Kobau O b s e r v a t o r y , and L y t t o n weather s t a t i o n s ( E n v i r o n m e n t Canada, 1971-72, 1976-83). I c o l l e c t e d d a t a i n t h e summer of 1983 f o r V e r n o n H i l l a t 1150m, Mt. Apex a t 2247m, G r e e n Mt. a t 1525m, and D i v i d e n d Mt. a t 1920m. Vern o n H i l l i s n e a r V e r n o n , B.C., t h e l a t t e r t h r e e l o c a l i t i e s a r e near P e n t i c t o n , B.C. (A p p e n d i x 1 ) . The p o p u l a t i o n s u s e d were t h o s e from Mt. Kobau, G r e e n Mt., Mt. Apex, D i v i d e n d Mt., V e r n o n H i l l , S i l v e r S t a r Mt., C r a t e r Mt., L y t t o n , and C a t h e d r a l Park ( A p p e n d i x 1 ) . The e x t r a p o l a t i o n p r o c e d u r e was t o e s t a b l i s h a 10 y e a r mean t e m p e r a t u r e p e r 2 week i n t e r v a l u s i n g a v a l l e y b o t t o m E n v i r o n m e n t Canada weather s t a t i o n . A s e c o n d s e t of d a t a ( s h o r t - t e r m ) was u s e d t o e s t a b l i s h t h e change i n a i r t e m p e r a t u r e w i t h e l e v a t i o n (and a s s o c i a t e d f a c t o r s ) between t h e two s t a t i o n s by c o m p a r i n g mean_ t e m p e r a t u r e s f o r t h e d a y s i n common. The s e c o n d s e t o f d a t a was p o s i t i o n e d a t a h i g h e r e l e v a t i o n t h a n t h e l o n g - t e r m s t a t i o n , on t h e same e x p o s u r e as t h e c o l o n y , and ' i n 18 l i n e ' w i t h t h e c o l o n y w i t h r e s p e c t t o c h a n g e s a s s o c i a t e d w i t h t o p o g r a p h y ( e s t i m a t e d c h a n g e s i n c l o u d i n e s s , t e m p e r a t u r e i n v e r s i o n s , and a i r movements). T h i s s l o p e was t h e n u s e d t o e s t i m a t e t h e mean t e m p e r a t u r e i n t h e n e a r b y phoebus c o l o n i e s . O n l y p o p u l a t i o n s f o r w h i c h a v a i l a b l e w e a t h e r d a t a a l l o w e d r e a s o n a b l e e x t r a p o l a t i o n t o t h e a c t u a l c o l o n i e s were u s e d f o r t h i s c a l c u l a t i o n . F l i g h t p e r i o d s ( A p p e n d i x 1) were e s t i m a t e d f r o m t h e amount of wear on t h e a v a i l a b l e s p e c i m e n s . I f a l l s p e c i m e n s were f r e s h , t h e f l i g h t s e a s o n had j u s t s t a r t e d . I f a l l s p e c i m e n s were worn, t h e s e a s o n was e n d i n g . In t h e m i d d l e of t h e s e a s o n t h e r e was a m i x t u r e of worn and f r e s h s p e c i m e n s . A d j u s t m e n t s were made when i t was known t h a t a f l i g h t p e r i o d was a b n o r m a l l y e a r l y o r l a t e . A l l n o r m a l f l i g h t p e r i o d s were assumed t o be 6 weeks l o n g b a s e d on my c o l l e c t i n g e x p e r i e n c e . C o r r e l a t i o n s w i t h P h e n o t y p e (1 C o r r e l a t i o n s between P o p u l a t i o n s M u l t i p l e r e g r e s s i o n s between e a c h p h e n o t y p i c c h a r a c t e r and e l e v a t i o n and l a t i t u d e were done. U s i n g o n l y t h o s e p o p u l a t i o n s f o r w h i c h I c o u l d e s t i m a t e mean n o r m a l a i r t e m p e r a t u r e s d u r i n g t h e f l i g h t p e r i o d , I compared t h e f i t o f e l e v a t i o n "and t e m p e r a t u r e t o mean p h e n o t y p e . In o r d e r t o a l l o w f o r t h e e f f e c t o f d i f f e r e n t sample s i z e s , e a c h mean was w e i g h t e d a c c o r d i n g t o t h e sample s i z e of t h a t p o p u l a t i o n . Mean p h e n o t y p e s were u s e d i n s t e a d of i n d i v i d u a l p h e n o t y p e s i n o r d e r t o r e d u c e t h e v a r i a n c e 19 and t o reduce the e f f e c t of d i f f e r e n c e s i n v a r i a n c e s between p o p u l a t i o n s . I a l s o compared mean p o p u l a t i o n phenotypes between sexes , u s i n g r e g r e s s i o n s of mean male phenotype as a f u n c t i o n of mean female phenotype, w i t h each data p o i n t weighted by the s m a l l e r of the two sample s i z e s . (2 C o r r e l a t i o n s w i t h i n a P o p u l a t i o n The p o p u l a t i o n a t V e r n o n , e l . 1l00-1200m was a n a l y z e d f o r : ( a ) c o r r e l a t i o n s between the p h e n o t y p i c c h a r a c t e r s f o r i n d i v i d u a l s , (b)changes between y e a r s , and (c)changes w i t h i n a f l i g h t season. I n h e r i t a n c e of P h e n o t y p e D i r e c t d e t e r m i n a t i o n of h e r i t a b i l i t y of phenotype t h r o u g h p a r e n t - o f f s p r i n g r e g r e s s i o n was not p o s s i b l e owing to the death of the l a r v a e from d i s e a s e . However, another s e t of data was a v a i l a b l e from r e a r i n g done i n 1980 which p e r m i t s some c o n c l u s i o n s as t o the c o n t r i b u t i o n s of d i f f e r e n c e s i n genotype and d e v e l o p m e n t a l environment i n d e t e r m i n i n g a d u l t phenotype. In 1979 I sampled P. phoebus p o p u l a t i o n s from s o u t h e r n B r i t i s h C o l u m b i a , A l b e r t a , and Montana and o b t a i n e d eggs from 1 or 2 a r b i t r a r i l y s e l e c t e d females from each of 6 p o p u l a t i o n s . The o f f s p r i n g were r e a r e d s i m u l t a n e o u s l y under u n c o n t r o l l e d outdoor c o n d i t i o n s i n 1980. R e c o r d s were kept of the p o p u l a t i o n s , but not the i n d i v i d u a l f e m a l e s , from w h i c h the eggs o r i g i n a t e d . P h e n o t y p e s of a r b i t r a r y samples from the pa r e n t p o p u l a t i o n s and 2 0 of a l l t h e r e a r e d o f f s p r i n g were s c o r e d . F o r t h e c h a r a c t e r s f o r whi c h t h e r e q u i r e d a s s u m p t i o n s were met, an a n a l y s i s of v a r i a n c e was done w i t h PHENOTYPE a f u n c t i o n o f LOCALITY, and ENVIRONMENT ( r e a r e d v s w i l d ) n e s t e d w i t h i n LOCALITY. The ' E n v i r o n m e n t ' t e r m i n c l u d e d b o t h t h e e f f e c t of t h e d e v e l o p m e n t a l e n v i r o n m e n t s ( s i x w i l d and one r e a r e d ) and t h e g e n e t i c e f f e c t s due t o e a c h r e a r e d sample h a v i n g o r i g i n a t e d from t h e eggs of 1-2 f e m a l e s i n s t e a d of from a random sample of eggs from a l l f e m a l e s i n a p o p u l a t i o n . When t h e a s s u m p t i o n s r e q u i r e d f o r an ANOVA were not met, c o m p a r i s o n s o f t h e d i s t r i b u t i o n s o f t h e r e a r e d samples were made u s i n g Duncan's M u l t i p l e Range T e s t o r t h e Mann-Whitney U - T e s t . S i n c e e a c h sample was r e a r e d under t h e same c o n d i t i o n s , any d i f f e r e n c e s between r e a r e d s a m p l e s must be due t o g e n e t i c d i f f e r e n c e s . The 6 l o c a l i t i e s a r e g i v e n i n T a b l e 1, w i t h f u r t h e r d e t a i l s i n A p p e n d i x 1. T a b l e I . L o c a l i t i e s f o r I n h e r i t a n c e D a t a Number L o c a l i t y D e s c r i p t i o n 1 C anada, B.C., L y t t o n , B o t a n i e L a k e , el.1370-1525m 2 U.S.A., Montana, M i s s o u l a , e l . 1525m 3 Canada, A l t a , K a n a n a s k i s Rd., R e g a l Ck., el.1525m 4 Canada, B.C., Manning P a r k , G i b s o n P a s s , el.1370m 5 Canada, B.C., B i g Bar C r e e k , P o i s o n Mt., el.2135-2195m 6 Canada, B.C., P e n t i c t o n , Mt. Apex, el.2190-2247m 21 RESULTS Ph e n o t y p e C o r r e l a t i o n w i t h E l e v a t i o n and L a t i t u d e F o r v a r i a t i o n i n b a s a l and d i s t a l m e l a n i s m i n P. phoebus t o f i t t h e p a t t e r n o f a l p i n e m e l a n i s m , t h e r e must be a p o s i t i v e c o r r e l a t i o n between v a r i a t i o n i n a c h a r a c t e r and e l e v a t i o n and l a t i t u d e . F o r t h e b a s a l wing a r e a , t h e r e i s a good c o r r e l a t i o n between mean b a s a l p a t c h b l a c k n e s s and e l e v a t i o n and l a t i t u d e , w i t h t h e c o r r e l a t i o n b e i n g b e t t e r f o r m a l e s t h a n f e m a l e s ( T a b l e I I ) . The c o r r e l a t i o n w i t h e l e v a t i o n i s p o o r f o r male b a s a l p a t c h w i d t h and b a s a l t r a n s p a r e n c y , and n o n - e x i s t e n t f o r t h e s e c h a r a c t e r s f o r f e m a l e s . F o r t h e d i s t a l wing a r e a , t h e r e i s no c o r r e l a t i o n between e l e v a t i o n o r l a t i t u d e and e i t h e r d i s t a l b l a c k n e s s o r d i s t a l t r a n s p a r e n c y ( T a b l e I I ) . V a r i a t i o n i n mean f r o n t wing l e n g t h shows t h e b e s t c o r r e l a t i o n w i t h e l e v a t i o n and l a t i t u d e of any of th e measured c h a r a c t e r s . T h e r e f o r e , t h e components o f b a s a l m e l a n i s m f i t t h e e x p e c t e d p a t t e r n o f a l p i n e m e l a n i s m ( t o d i f f e r i n g d e g r e e s ) but d i s t a l m e l a n i s m does n o t . V a r i a t i o n i n s i z e f i t s t h e p a t t e r n of a l p i n e d w a r f i s m q u i t e w e l l . P h e n o t y p e C o r r e l a t i o n w i t h A i r T e m p e r a t u r e I f t h e s i g n i f i c a n t c o r r e l a t i o n s between a d u l t p h e n o t y p e and e l e v a t i o n and l a t i t u d e r e s u l t from a d a p t a t i o n t o d i f f e r e n t a i r t e m p e r a t u r e s , t h e y s h o u l d c o r r e l a t e a t l e a s t a s w e l l w i t h mean a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d a s w i t h e l e v a t i o n . 22 T a b l e I I . Changes i n P h e n o t y p e w i t h E l e v a t i o n and L a t i t u d e . M u l t i p l e r e g r e s s i o n o f mean p h e n o t y p i c c h a r a c t e r s as a f u n c t i o n of e l e v a t i o n and l a t i t u d e . R e g r e s s i o n C o e f f i c i e n t s (* 100) r2 V a r i a b l e Sex n E l e v a t ion(m) L a t i t u d e ( d e g r e e s ) BASAL M 61 .02*** 4 1 * * * .25 WIDTH F 53 ns ns BASAL M 61 . 28*** 38*** .59 BLACKNESS F 53 . 16*** 32*** .33 BASAL M 61 .03** ns . 1 5 TRANSPARENCY F 53 ns ns DISTAL M 61 ns ns BLACKNESS F 53 ns ns DISTAL M 56 ns ns TRANSPARENCY F 51 ns ns FRONT WING M 61 -. 35*** .60 LENGTH F 53 - . 3 7 * * * - 4 3 * * * .62 A maximum of two ' p o p u l a t i o n s ' p e r m o u n t a i n (1 h i g h , 1 low e l e v a t i o n ) were i n c l u d e d t o a d v o i d g i v i n g any one a r e a e x t r a w e i g h t . Means f o r a r c s i n t r a n s f o r m e d d a t a were u s e d , and e a c h mean was w e i g h t e d by i t s sample s i z e . **=P<.01; ***=p<.00l. n = number of p o p u l a t i o n s . C o n v e r s e l y , t h o s e c h a r a c t e r s w h i c h a r e n o t c o r r e l a t e d w i t h e l e v a t i o n and l a t i t u d e s h o u l d not be be c o r r e l a t e d w i t h a i r t e m p e r a t u r e e i t h e r . B a s a l p a t c h w i d t h c o r r e l a t e s b e t t e r w i t h mean a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d t h a n w i t h e l e v a t i o n f o r m a l e s , a l t h o u g h v e r y l i t t l e o f t h e v a r i a t i o n i s a c c o u n t e d f o r by e i t h e r c o r r e l a t i o n , and n e i t h e r c o r r e l a t i o n i s s i g n i f i c a n t f o r f e m a l e s ( T a b l e I I I ) . B a s a l p a t c h b l a c k n e s s c o r r e l a t e s a b o u t as w e l l w i t h a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d as w i t h e l e v a t i o n , w i t h t h e c o r r e l a t i o n b e i n g s l i g h t l y worse f o r males and b e t t e r f o r f e m a l e s ( F i g u r e s 3-4). In c o n t r a s t , t h e 23 F i g u r e 3. Male b a s a l p a t c h b l a c k n e s s as a f u n c t i o n of e l e v a t i o n and a i r t e m p e r a t u r e . ( a ) B a s a l p a t c h b l a c k n e s s a g a i n s t e l e v a t i o n ; ( b ) B a s a l p a t c h b l a c k n e s s a g a i n s t mean a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d . See T a b l e I I I f o r r e g r e s s i o n c o e f f i c i e n t s , r - s q u a r e d v a l u e s , and s i g n i f i c a n c e . An a r c s i n e s c a l e i s u s e d on t h e o r d i n a t e a x i s . 25 F i g u r e 4. Female b a s a l p a t c h b l a c k n e s s as a f u n c t i o n o f e l e v a t i o n and a i r t e m p e r a t u r e . ( a ) B a s a l p a t c h b l a c k n e s s a g a i n s t e l e v a t i o n ; ( b ) B a s a l p a t c h b l a c k n e s s a g a i n s t mean a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d . See T a b l e I I I f o r r e g r e s s i o n c o e f f i c i e n t s , r - s q u a r e d v a l u e s , and s i g n i f i c a n c e . An a r c s i n e s c a l e i s u s e d on t h e o r d i n a t e a x i s . 27 T a b l e I I I : Changes i n Ph e n o t y p e w i t h E l e v a t i o n and A i r T e m p e r a t u r e . S i n g l e f a c t o r r e g r e s s i o n s o f mean p h e n o t y p e as a f u n c t i o n of e l e v a t i o n or mean a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d . V a r i a b l e Sex E l e v a t i o n ( m ) A i r T e m p e r a t u r e ( ° C ) R e g r . C o e f f . r2 R e g r . C o e f f . r2 BASAL M ns ns -3.6 * . 10 WIDTH F ns — ns BASAL M . 1 9 *** .66 -70 * * * . 58 BLACKNESS F .10 * * * .29 -54 *** .40 BASAL M .05 * * * .40 -17 * * * .25 TRANSPARENCY F .06 * * * .27 -27 * * * .25 DISTAL M -.09 ** .20 ns BLACKNESS F ns ns DISTAL M ns ns TRANSPARENCY F ns ns FRONT WING M -.35 * * * .70 1 07 * * * . 42 LENGTH F -.23 * * * .53 1 1 4 * * * .40 R e g r e s s i o n c o e f f i c i e n t s (* 10 0) f o r a r c s i n e t r a n s f o r m e d d a t a , means w e i g h t e d by sample s i z e . Number of p o p u l a t i o n s = 44 ( m a l e s ) , 40 ( f e m a l e s ) . *p<.05; **p<.005; * * * p < . 0 0 l . c o r r e l a t i o n s of b a s a l t r a n s p a r e n c y and f r o n t wing l e n g t h w i t h a i r t e m p e r a t u r e a r e much worse t h a n t h e c o r r e l a t i o n s w i t h e l e v a t i o n . In t h e d i s t a l wing a r e a , d i s t a l b l a c k n e s s and d i s t a l t r a n s p a r e n c y a r e not c o r r e l a t e d w i t h a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d , j u s t as t h e y were n o t c o r r e l a t e d w i t h e l e v a t i o n . T h e r e f o r e , some o f t h e v a r i a t i o n i n b a s a l m e l a n i s m , e s p e c i a l l y b a s a l p a t c h b l a c k n e s s , may be an a d a p t a t i o n t o d i f f e r e n t a i r t e m p e r a t u r e s d u r i n g t h e f l i g h t p e r i o d f o r d i f f e r e n t p o p u l a t i o n s . V a r i a t i o n i n t h e o t h e r c h a r a c t e r which i s s t r o n g l y c o r r e l a t e d w i t h e l e v a t i o n , f r o n t wing l e n g t h , i s 28 u n l i k e l y t o be an a d a p t a t i o n t o d i f f e r e n t a i r t e m p e r a t u r e s d u r i n g t h e f l i g h t p e r i o d , nor i s t h e v a r i a t i o n i n b a s a l t r a n s p a r e n c y . As e x p e c t e d , d i s t a l t r a n s p a r e n c y and d i s t a l b l a c k n e s s were no b e t t e r c o r r e l a t e d w i t h a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d t h a n w i t h e l e v a t i o n , and t h e r e f o r e a r e a l s o u n l i k e l y t o be an a d a p t a t i o n t o d i f f e r e n t a i r t e m p e r a t u r e s . P h e n o t y p e V a r i a t i o n w i t h i n I n d i v i d u a l M o u n t a i n s I s e l e c t e d t h e f o u r m o u n t a i n s f o r w h i c h I had t h e g r e a t e s t sample s i z e s and g r e a t e s t e l e v a t i o n a l r a n g e . Mt. Apex and G r e e n Mt. a r e v e r y c l o s e t o g e t h e r i n t h e s o u t h - c e n t r a l Okanagan V a l l e y , w i t h j u s t a f o r e s t e d v a l l e y (5 km wide) s e p a r a t i n g them. S i l v e r S t a r Mt. i s i n t h e n o r t h e r n Okanagan V a l l e y , and B o t a n i e Mt. i s i n t h e w e s t e r n Thompson V a l l e y ( A p p e n d i x 1 ) . P h e n o t y p e s v a r y c o n s i d e r a b l y between t h e m o u n t a i n s , w i t h t h e t h r e e samples from s i m i l a r e l e v a t i o n s showing as much d i f f e r e n c e between t h e m s e l v e s as w i t h t h e sample from a l o w e r e l e v a t i o n ( T a b l e I V ) . The v a r i a t i o n between m o u n t a i n s i s g r e a t e s t f o r f e m a l e s , and t h e d i s t a l wing c h a r a c t e r s a r e more v a r i a b l e t h a n t h e b a s a l c h a r a c t e r s . I f v a r i a t i o n i n mean b a s a l p a t c h w i d t h and b l a c k n e s s a r e a d a p t a t i o n s t o d i f f e r e n t a i r t e m p e r a t u r e s d u r i n g t h e f l i g h t s e a s o n , t h e n t h e y s h o u l d be c o r r e l a t e d w i t h e l e v a t i o n on most, i f not a l l m o u n t a i n s . P o p u l a t i o n s from a l l f o u r m o u n t a i n s show a p o s i t i v e c o r r e l a t i o n between male b a s a l p a t c h b l a c k n e s s and e l e v a t i o n , but f o r f e m a l e s t h e c o r r e l a t i o n i s o n l y s i g n i f i c a n t f o r two o f t h e f o u r m o u n t a i n s ( T a b l e V ) . F r o n t wing l e n g t h i s 29 T a b l e IV. Phenotype D i f f e r e n c e s between M o u n t a i n s . Mt.Apex G r e e n Mt. S i l v e r S t a r B o t a n i e Mt. 1750m 1860m 1800m 1525m V a r i a b l e Mean n mean n mean n mean n (s . e. ) ( s . e . ) ( s . e . ) ( s . e . ) BASAL M 7.48 20 7.42 79 7.45 81 7.52 23 WIDTH ( .08) ( .06) ( .06) ( .09) F 8.32 6 8.35 43 9.39 28 8.93 1 5 ( .40) ( .08) (.29) ( . 13) BASAL M 17.27 1 9 18.61 81 18.32 81 1 9.68 23 BLACKNESS (.46) ( . 19) ( .18) ( .32) F 15.44 6 16.96 43 17.87 28 18.85 1 5 (.71 ) ( .33) (.27) (.29) BASAL M 3.09 1 9 2.98 81 2. 67 81 2.34 23 TRANSPAR. ( .20) ( .10) (.13) ( .17) F 4.73 6 4. 06 43 3.97 28 3.71 1 5 ( .26) ( .22) ( .30) ( .24) DISTAL M 3.90 16 4.54 74 6.00 80 3.64 1 7 BLACKNESS ( .25) ( . 15) (.15) ( .29) F 8.70 5 1 1 .42 42 14.48 28 10.15 1 4 (.27) ( .30) (.17) ( .28) DISTAL M 3.51 7 4.74 69 6.77 69 3.79 8 TRANSPAR. (.22) ( . 15) (.15) (.12) F 8.52 5 9.36 41 11.86 27 9.30 9 ( .31 ) ( .29) ( .28) ( .23) WING M 30.97 1 9 31.41 80 31 .36 81 31.24 2 1 LENGTH ( .23) ( . 18) ( . 14) ( .30) F 32.50 5 32.63 43 33.41 28 32.07 1 5 (1.12) ( .27) (.23) ( .57) The means a r e f o r a r c s i n t r a n s f o r m e d d a t a . n e g a t i v e l y c o r r e l a t e d w i t h e l e v a t i o n i n most o f t h e m o u n t a i n s . D i s t a l b l a c k n e s s and d i s t a l t r a n s p a r e n c y a r e c o r r e l a t e d w i t h e l e v a t i o n on t h e s e f o r m o u n t a i n s , a l t h o u g h t h e y a r e not c o r r e l a t e d when a l l m o u n t a i n s were c o n s i d e r e d t o g e t h e r . T h e r e f o r e d i s t a l m e l a n i s m f o l l o w s t h e p a t t e r n o f a l p i n e m e l a n i s m on a l o c a l l e v e l , but n o t a t t h e g e n e r a l l e v e l e x p e c t e d . The 30 T a b l e V. P h e n o t y p e Changes w i t h i n M o u n t a i n s . R e g r e s s i o n s f o r p h e n o t y p e as a f u n c t i o n o f e l e v a t i o n f o r f o u r m o u n t a i n s . V a r i a b l e Mt. Apex Green Mt. S i l v e r S t a r B o t a n i e Mt. RC r2 n RC r2 n RC r2 n RC r2 n B a s a l M ns 1 1 0 ns 278 ns 500 ns 88 W i d t h F ns 24 ns 1 1 1 .07 .05 184 ns 58 B a s a l M .39 . 1 2 1 08 .48 .12 277 .06 .01 488 .21 . 1 2 80 B l a c k . F .75 .43 24 ns 1 08 ns 181 .28 . 1 5 53 B a s a l M ns 1 08 ns 277 .05 .01 488 .07 .08 80 T r a n s . F ns 24 ns 108 ns 181 ns 53 D i s t a l M ns 1 02 ns 265 .10 .04 472 ns 67 B l a c k . F .71 .50 23 ns 109 .44 .27 169 .22 .22 55 D i s t a l M .29 .20 74 ns 191 .23 . 1 7 387 . 1 5 .20 35 T r a n s . F ns 21 ns 93 . 29 .24 151 . 1 4 . 1 1 39 Wing M -.47 . 15 109 ns 276 -.40 .40 496 -.29 .29 85 L e n g t h F -.44 . 1 2 23 ns 1 1 0 -.34 .29 1 83 -.28 . 1 5 58 M=male, F = f e m a l e ; ns=not s i g n i f i c a n t (p>.05), n=sample s i z e . R C = ( r e g r e s s i o n c o e f f i e n t * 1 0 0 ) f o r a r c s i n e t r a n s f o r m e d d a t a . F o r l o c a l i t y d e t a i l s , e l e v a t i o n s p r e a d , and sample s i z e p e r e l e v a t i o n see A p p e n d i x 1. Mt. Apex and G r e e n Mt. - i n c l u d e o n l y t h o s e s i t e s l a b e l l e d a s s u c h ; S i l v e r S t a r Mt. - i n c l u d e s V e r n o n H i l l & Swan La k e ; B o t a n i e M t . - i n c l u d e s a l l L y t t o n s i t e s . ' same c o r r e l a t i o n s w i t h e l e v a t i o n do not o c c u r - o n G r e e n Mt. as on Mt. Apex, d e s p i t e t h e i r c l o s e n e s s . The low r - s q u a r e d v a l u e s i n d i c a t e h i g h v a r i a n c e w i t h i n e a c h e l e v a t i o n r e l a t i v e t o v a r i a t i o n between e l e v a t i o n s . The s l o p e s of t h e r e g r e s s i o n s v a r y c o n s i d e r a b l y between m o u n t a i n s , s u g g e s t i n g t h a t t h e ' e f f e c t ' of e l e v a t i o n may be d i f f e r e n t f o r e a c h m o u n t a i n . 31 P h e n o t y p e C o r r e l a t i o n s between P o p u l a t i o n s A n o t h e r s o u r c e of e v i d e n c e f o r t h e a d a p t i v e s i g n i f i c a n c e o f t h e p h e n o t y p i c c h a r a c t e r s i s c o r r e l a t i o n s between p o p u l a t i o n s f o r t h e c h a r a c t e r s . I f two c h a r a c t e r s a r e c o r r e l a t e d between p o p u l a t i o n s , t h e n t h e y may be a s s o c i a t e d w i t h t h e same e n v i r o n m e n t a l v a r i a b l e , a l t h o u g h o t h e r e x p l a n a t i o n s a r e a l s o p o s s i b l e . I f t h e r e i s no c o r r e l a t i o n between two c h a r a c t e r s , t h e n t h e y d e f i n i t e l y a r e n o t a s s o c i a t e d w i t h t h e same e n v i r o n m e n t a l v a r i a b l e s . Between p o p u l a t i o n s b a s a l p a t c h w i d t h i s w e l l c o r r e l a t e d w i t h p a t c h b l a c k n e s s i n m a l e s , and b o t h p a t c h w i d t h and b l a c k n e s s a r e p o o r l y c o r r e l a t e d w i t h w i n g l e n g t h i n b o t h s e x e s ( T a b l e V I ) . B a s a l b l a c k n e s s i s weakly c o r r e l a t e d w i t h d i s t a l b l a c k n e s s i n f e m a l e s , but not i n m a l e s , and b a s a l t r a n s p a r e n c y i s f a i r l y w e l l c o r r e l a t e d w i t h d i s t a l t r a n s p a r e n c y i n b o t h s e x e s . D i s t a l b l a c k n e s s i s q u i t e w e l l c o r r e l a t e d w i t h d i s t a l t r a n s p a r e n c y i n b o t h s e x e s , and w i t h wing l e n g t h i n males o n l y . F o r t h e s e c o m b i n a t i o n s of c h a r a c t e r s t h e same e n v i r o n m e n t a l v a r i a b l e s may be a s s o c i a t e d w i t h t h e o b s e r v e d p a t t e r n s o f v a r i a t i o n between p o p u l a t i o n s . A l l t h e o t h e r p o s s i b l e c o r r e l a t i o n s between t h e s i x c h a r a c t e r s f o r e a c h sex a r e n o t s i g n i f i c a n t , and t h e r e f o r e t h e same e n v i r o n m e n t a l v a r i a b l e s a r e d e f i n i t e l y n o t a s s o c i a t e d w i t h v a r i a t i o n i n t h o s e p a i r s o f c h a r a c t e r s . 32 T a b l e V I . I n t e r p o p u l a t i o n C o r r e l a t i o n s between C h a r a c t e r s . P a r t i a l C o r r e l a t i o n s between p h e n o t y p i c c h a r a c t e r s u s i n g p o p u l a t i o n means. SEX BASAL WIDTH Male .69 BASAL Female ns BLACKNESS Male ns ns BASAL Female ns ns TRANSPARENCY Male ns ns ns DISTAL Female ns .37 ns BLACKNESS Male ns ns .54 .67 DISTAL Female ns ns . 40 .65 TRANSPARENCY Male -.28 -.40 ns .52 ns FRONT ' Female -.32 -.38 ns ns ns LENGTH M a l e s : n=53 p o p u l a t i o n s ; F e m a l e s : n=46 p o p u l a t i o n s ; o n l y i n d i v i d u a l s f o r w h i c h no d a t a were m i s s i n g were u s e d ; means w e i g h t e d by sample s i z e ; a maximum o f 2 p o p u l a t i o n s p e r m o u n t a i n were u s e d ; ns = n o t s i g n i f i c a n t ( p>.05). P o p u l a t i o n P h e n o t y p e C o r r e l a t i o n s between S e x e s Many of t h e above c o r r e l a t i o n s a r e sex d e p e n d e n t . I f t h e mean v a l u e s of t h e c h a r a c t e r s a r e not c o r r e l a t e d between t h e s e x e s , t h e same e n v i r o n m e n t a l f a c t o r s c an n o t be a s s o c i a t e d w i t h v a r i a t i o n i n e a c h s e x . When t h e p h e n o t y p e of one sex i s r e g r e s s e d a g a i n s t t h e o t h e r , n ot o n l y i s t h e c o r r e l a t i o n d e t e r m i n e d but a l s o t h e r e l a t i v e r a t e s o f change i n p h e n o t y p e between p o p u l a t i o n s c a n be s e e n . I f t h e r a t e o f change of a c h a r a c t e r between p o p u l a t i o n s i s s i m i l a r f o r t h e s e x e s , t h e n t h e e x t e n t of s e x u a l d i m o r p h i s m i s c o n s t a n t between p o p u l a t i o n s . A l l t h e c h a r a c t e r s a r e c o r r e l a t e d between s e x e s t o some e x t e n t ( T a b l e V I I ) , b ut t h e c o r r e l a t i o n s f o r b a s a l p a t c h w i d t h 33 T a b l e V I I . I n t e r p o p u l a t i o n C o r r e l a t i o n s between S e x e s . R e g r e s s i o n s of mean male p o p u l a t i o n p h e n o t y p e as a f u n c t i o n of mean f e m a l e p h e n o t y p e . V a r i a b l e R e g r . C o e f f . * r2 BASAL WIDTH .37 .24 BASAL BLACKNESS .91 .66 BASAL TRANSPARENCY .48 .46 DISTAL BLACKNESS .40 . 1 9 DISTAL TRANSPARENCY .73 .53 FRONT WING LENGTH .95 .93 Means w e i g h t e d by t h e sample s i z e o f t h e l e a s t numerous sex . A maximum of two samples p e r m o u n t a i n were u s e d , w i t h a t o t a l of 81 p o p u l a t i o n s . * A l l r e g r e s s i o n s s i g n i f i c a n t a t p < . 0 0 l . and d i s t a l b l a c k n e s s a r e s u f f i c i e n t l y poor t h a t i t i s u n l i k e l y t h a t t h e same e n v i r o n m e n t a l v a r i a b l e s a r e a s s o c i a t e d w i t h v a r i a t i o n i n t h o s e c h a r a c t e r s . The r a t e s of change between p o p u l a t i o n s f o r t h o s e c h a r a c t e r s a r e not t h e same f o r e a c h sex, i n d i c a t i n g t h a t t h e e x t e n t of s e x u a l d i m o r p h i s m i s v a r i a b l e . In c o n t r a s t , f r o n t wing l e n g t h and b a s a l p a t c h b l a c k n e s s a r e w e l l c o r r e l a t e d between s e x e s , and t h e r a t e of change between p o p u l a t i o n s i s t h e same f o r t h e s e x e s , i n d i c a t i n g t h a t ( a ) t h e r e l a t i v e s i z e and b a s a l b l a c k n e s s of t h e s e x e s i s c o n s t a n t , and ( b ) i t i s l i k e l y t h a t v a r i a t i o n i n t h o s e c h a r a c t e r s i s a s s o c i a t e d w i t h t h e same e n v i r o n m e n t a l f a c t o r s i n b o t h s e x e s . The o t h e r two c h a r a c t e r s , b a s a l t r a n s p a r e n c y and d i s t a l t r a n s p a r e n c y , a r e not v e r y w e l l c o r r e l a t e d between s e x e s , and t h e r a t e of change between p o p u l a t i o n s i s not t h e same, i n d i c a t i n g t h a t t h e e x t e n t 34 of s e x u a l d i m o r p h i s m i s v a r i a b l e and s u g g e s t i n g t h a t most of t h e t h e i r v a r i a t i o n i s n o t a s s o c i a t e d w i t h t h e same e n v i r o n m e n t a l f a c t o r s i n t h e two s e x e s . P h e n o t y p e C o r r e l a t i o n s w i t h i n a P o p u l a t i o n Two p o s s i b l e ( but not e x c l u s i v e ) e x p l a n a t i o n s of c o r r e l a t i o n s between mean v a l u e s of t h e p h e n o t y p i c c h a r a c t e r s a t th e i n t e r p o p u l a t i o n l e v e l , a r e t h a t v a r i a t i o n i n t h e c h a r a c t e r s i s due t o ( a ) t h e i r b e i n g a f f e c t e d by t h e same changes i n d e v e l o p m e n t a l e n v i r o n m e n t , and ( b ) t h e i r e x p r e s s i o n b e i n g g e n e t i c a l l y a s s o c i a t e d i n some manner. Th e s e p o s s i b i l i t i e s can be l o o k e d a t t o g e t h e r , but n o t d i s t i n g u i s h e d , by e x a m i n i n g t h e c o r r e l a t i o n s between c h a r a c t e r s f o r t h e i n d i v i d u a l s w i t h i n a p o p u l a t i o n . I f t h e c h a r a c t e r s a r e not c o r r e l a t e d t h e n n e i t h e r p o s s i b i l i t y can be t r u e , but i f t h e y a r e c o r r e l a t e d t h e n e i t h e r may be t r u e . The m a j o r i t y of t h e p o s s i b l e c o m p a r i s o n s between c h a r a c t e r s show no s i g n i f i c a n t c o r r e l a t i o n ( T a b l e V I I I ) , and t h e r e f o r e t h e r e c an be no g e n e t i c o r d e v e l o p m e n t a l a s s o c i a t i o n between t h o s e c h a r a c t e r s . T h e r e a r e o n l y t h r e e p o s s i b l y b i o l o g i c a l l y s i g n i f i c a n t , b u t v e r y weak, c o r r e l a t i o n s between c h a r a c t e r s : b a s a l p a t c h w i d t h w i t h p a t c h d a r k n e s s , b a s a l t r a n s p a r e n c y w i t h d i s t a l t r a n s p a r e n c y , and d i s t a l b l a c k n e s s w i t h d i s t a l t r a n s p a r e n c y . T h e r e f o r e , w i t h t h o s e t h r e e p o s s i b l e e x c e p t i o n s , d e t e r m i n a t i o n o f p h e n o t y p e i s c o m p l e t e l y i n d e p e n d e n t f o r a l l t h e c h a r a c t e r s . 35 T a b l e V I I I : C o r r e l a t i o n s between C h a r a c t e r s w i t h i n I n d i v i d u a l s . P a r t i a l C o r r e l a t i o n s between p h e n o t y p i c c h a r a c t e r s w i t h i n a p o p u l a t i o n . SEX YEAR BASAL WIDTH M 1982 ns 1 983 .24 BASAL F 1 982 .24 BLACKNESS 1 983 ns M 1 982 ns ns 1 983 ns ns BASAL F 1 982 ns ns TRANSPARENCY 1 983 ns ns M 1 982 ns ns ns 1 983 ns ns ns DISTAL F 1 982 ns ns ns BLACKNESS 1983 ns ns ns M 1 982 ns ns ns .20 1 983 ns ns . 36 .20 DISTAL F 1 982 ns ns -.27 ns TRANSPARENCY 1 983 ns ns ns .26 M 1 982 ns ns ns ns ns 1 983 ns ns ns ns ns FRONT WING F 1 982 ns ns ns ns ns LENGTH 1 983 ns ns ns ns ns ns = c o r r e l a t i o n s not s i g n i f i c a n t (p>.05). M=male, F=female Sample s i z e s : M a l e ( l 9 8 2 ) = 123; M a l e ( l 9 8 3 ) = 130; F e m a l e d 982) = 44; F e m a l e ( l 9 8 3 ) = 84. O n l y s p e c i m e n s f o r w h i c h a l l s i x c h a r a c t e r s were s c o r e d were u s e d • P h e n o t y p i c V a r i a t i o n o v e r Time One p o s s i b l e p r o b l e m w i t h between p o p u l a t i o n c o r r e l a t i s t h e e r r o r i n e s t i m a t i n g t h e mean p o p u l a t i o n p h e n o t y p e s . A p o s s i b l e s o u r c e of e r r o r i s s i g n i f i c a n t c h a n g e s i n mean p h e n o t y p e w i t h i n o r between y e a r s . I f t h e r e a r e major c h a n g e s i n mean p h e n o t y p e w i t h t i m e , p o o r c o r r e l a t i o n s may be t h e r e s u l t o f t h a t v a r i a n c e . 36 F o r t h e V e r n o n H i l l p o p u l a t i o n , t h e mean v a l u e s of t h e c h a r a c t e r s v a r i e d o n l y s l i g h t l y between y e a r s ( T a b l e I X ) , w i t h c h a n g e s i n b o t h t h e means and t h e s h a p e s o f t h e f r e q u e n c y d i s t r i b u t i o n s c o n t r i b u t i n g t o 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 between y e a r s . The weather d u r i n g most o f t h e 1982 f l i g h t s e a s o n was a b n o r m a l l y warm and d r y w h i l e 1983 was a b n o r m a l l y c o l d and wet ( E n v i r o n m e n t Canada (1982,1983) r e c o r d s f o r V e r n o n , B . C . ) . T h e r e f o r e v a r i a t i o n i n mean p h e n o t y p e between y e a r s , e ven w i t h m a j o r e n v i r o n m e n t a l c h a n g e s , i s not l i k e l y t o a f f e c t p h e n o t y p e c o r r e l a t i o n s when s p e c i m e n s c o l l e c t e d i n d i f f e r e n t y e a r s a r e i n c l u d e d i n t h e c o m p a r i s o n s . Changes i n p h e n o t y p e from 1982 t o 1983 f o r t h e V e r n o n H i l l p o p u l a t i o n were a l m o s t a l w a y s i n t h e same d i r e c t i o n f o r b o t h s e x e s ( T a b l e I X ) , s u g g e s t i n g t h a t t h e same f a c t o r s may a f f e c t male and f e m a l e p h e n o t y p e . V a r i a t i o n w i t h i n a f l i g h t s e a s o n i s a l s o q u i t e s m a l l ( F i g u r e s 5-10), and i s u n l i k e l y t o add a g r e a t d e a l of v a r i a n c e t o t h e c o r r e l a t i o n s of mean p h e n o t y p e between p o p u l a t i o n s . T h e r e i s no p a r t i c u l a r p a t t e r n t o t h e v a r i a t i o n i n mean p h e n o t y p e o f newly emerged m a l e s , i n d i c a t i n g t h a t t h e r e w i l l be no c o r r e l a t i o n between s a m p l i n g t i m e and mean p h e n o t y p e . I n h e r i t a n c e of P h e n o t y p e T h e r e a r e two ways t o a n a l y z e t h e d a t a f o r i n h e r i t a n c e of p h e n o t y p e ( s e e ' m a t e r i a l s and methods' f o r f u r t h e r d e t a i l s ) . The p r e f e r r e d method o f a n a l y s i s i s t o use a n e s t e d ANOVA, f o r w h i c h t h e ' e n v i r o n m e n t ' t e r m i s n e s t e d w i t h i n t h e ' l o c a l i t y ' 37 F i g u r e 5. S e a s o n a l v a r i a t i o n i n male b a s a l p a t c h w i d t h . The means and s t a n d a r d e r r o r s ( v e r t i c a l b a r s ) a r e p l o t t e d f o r a l l males ( s o l i d c i r c l e s ) and f r e s h males o n l y (open c i r c l e s ) . F i g u r e 6. S e a s o n a l v a r i a t i o n i n male b a s a l p a t c h b l a c k n e s s . The means and s t a n d a r d e r r o r s ( v e r t i c a l b a r s ) a r e p l o t t e d f o r a l l m a l e s ( s o l i d c i r c l e s ) and f r e s h males o n l y (open c i r c l e s ) . B a s a l P a t c h C o l o r P r o p o r t i o n B l a c k ( % CO CO CD CD CD co co o ro ^ _ J I I I I I J I B a s a l P a t c h W i d t h (% of HW Ce l l ) ^ ^ ^ ^ 4^  ^. ^ 39 F i g u r e 7. S e a s o n a l v a r i a t i o n i n male b a s a l t r a n s p a r e n c y . The means and s t a n d a r d e r r o r s ( v e r t i c a l b a r s ) a r e p l o t t e d f o r a l l m a l e s ( s o l i d c i r c l e s ) and f r e s h m ales o n l y (open c i r c l e s ) . F i g u r e 8. S e a s o n a l v a r i a t i o n i n male d i s t a l b l a c k n e s s . The means and s t a n d a r d e r r o r s ( v e r t i c a l b a r s ) a r e p l o t t e d f o r a l l m a l e s ( s o l i d c i r c l e s ) and f r e s h m ales o n l y (open c i r c l e s ) . Distal Front Wing Proportion Black (%) - J . _L -A ro ro ro ro Ol s (D -k U Ul >| J I I I I I I I Basal Patch Color Area Without Scales (%) o o -± ro ro co co cn b bi b w b I i i i i i i 41 F i g u r e 9. S e a s o n a l v a r i a t i o n i n male d i s t a l t r a n s p a r e n c y . The means and s t a n d a r d e r r o r s ( v e r t i c a l b a r s ) a r e p l o t t e d f o r a l l m a l e s ( s o l i d c i r c l e s ) and f r e s h m a l e s o n l y (open c i r c l e s ) . F i g u r e 10. S e a s o n a l v a r i a t i o n i n male f r o n t wing l e n g t h . The means and s t a n d a r d e r r o r s ( v e r t i c a l b a r s ) a r e p l o t t e d f o r a l l m a l e s ( s o l i d c i r c l e s ) and f r e s h m a l e s o n l y (open c i r c l e s ) . W i n g l e n g t h (mm) CO CO CO CO CO en en b In b cnH cn «-o-ro I 10 cn CO O CO cn-} CO CO cn o) cn b D i s t a l F r o n t W i n g A r e a W i t h o u t S c a l e s (% 0) *g 00 CO O -* ro ... J ; 1 1 J 1 1 _ j Q • — • -43 T a b l e IX. Changes i n P h e n o t y p e w i t h Sex and Y e a r . C o m p a r i s o n * of male and f e m a l e p h e n o t y p e s w i t h i n and between y e a r s , w i t h i n a p o p u l a t i o n . V a r i a b l e Y e a r Means M a l e ( n ) F e m a l e ( n ) BASAL WIDTH 1 982 7. 62 (128) 9. 07 ( 44) 1 983 7. 51 (218) 8. 76 (103) BASAL BLACKNESS 1 982 17. 53 ( 1 28) 16. 81 ( 44) 1 983 18. 1 5 (217) 17. 53 ( 1 02) BASAL TRANSPARENCY 1 982 2. 87 (128) 4. 38 ( 44) 1983 2. 02 (217) 3. 28 (102) DISTAL BLACKNESS 1 982 4. 26 (128) 1 1 . 57 ( 44) 1 983 4. 62 (207) 10. 67 (103) DISTAL TRANSPARENCY 1 982 5. 78 (123) 10. 05 ( 44) 1 983 4. 1 2 ( 1 30) 9. 65 ( 84) FRONT WING LENGTH 1 982 34. 51 ( 1 28) 35. 95 ( 44) 1 983 34. 1 9 (218) 36. 40 (103) * Mann-Whitney U - T e s t (p<.05) u s e d b e c a u s e v a r i a n c e s were not homogeneous and f r e q u e n c y d i s t r i b u t i o n s h a p e s not n o r m a l f o r many o f t h e s a m p l e s . A l l between y e a r and between sex c o m p a r i s o n s have s i g n i f i c a n t l y d i f f e r e n t d i s t r i b u t i o n s , e x c e p t f o r between y e a r s f o r f e m a l e BASAL WIDTH and FRONT WING LENGTH. Means a r e a r c s i n t r a n s f o r m e d . D i s t a l b l a c k n e s s i s not d i r e c t l y c o m p a r a b l e between s e x e s ( s e e m e t h o d s ) . t e r m . I f t h e l o c a l i t y t erm i s s i g n i f i c a n t i n s u c h an a n a l y s i s , t h e n t h e r e a r e s i g n i f i c a n t d i f f e r e n c e s between r e a r e d s a m p l e s , and t h o s e d i f f e r e n c e s a r e c o r r e l a t e d w i t h t h e d i f f e r e n c e s o b s e r v e d i n t h e w i l d p o p u l a t i o n s . The s e c o n d method of a n a l y s i s i s t o o n l y compare t h e p h e n o t y p e s o f t h e r e a r e d s a m p l e s . S i n c e t h e r e a r i n g c o n d i t i o n s were u n c o n t r o l l e d , but a l l samples r e a r e d s i m u l t a n e o u s l y , t h e mean of e a c h r e a r e d sample s h o u l d not be s i g n i f i c a n t l y d i f f e r e n t even i f t h e p h e n o t y p e s of t h e i n d i v i d u a l s i n e a c h sample were a f f e c t e d by s l i g h t d i f f e r e n c e s 44 i n e n v i r o n m e n t . I f t h e r e a r e s i g n i f i c a n t d i f f e r e n c e s between r e a r e d samples f o r a c h a r a c t e r , t h e n t h e y s h o u l d be due t o g e n e t i c d i f f e r e n c e s f o r t h a t c h a r a c t e r . B a s a l P a t c h W i d t h Male b a s a l p a t c h w i d t h i s h i g h l y h e r i t a b l e . B a s a l p a t c h w i d t h v a r i e d s i g n i f i c a n t l y between w i l d p o p u l a t i o n s , and t h e r e a r e d s a m p l e s showed the same p a t t e r n of v a r i a t i o n ( F i g u r e 11). The r e a r e d samples were not s i g n i f i c a n t l y d i f f e r e n t f r o m t h e i r r e s p e c t i v e w i l d samples. T h e r e i s no e v i d e n c e f o r h e r i t a b i l i t y of f e m a l e b a s a l p a t c h w i d t h . B a s a l P a t c h B l a c k n e s s T h e r e i s v e r y l i t t l e e v i d e n c e t h a t v a r i a t i o n i n b a s a l p a t c h b l a c k n e s s i s h e r i t a b l e . The o n l y e v i d e n c e i s t h a t r e a r e d l o c a l i t y 4 males a r e s i g n i f i c a n t l y l i g h t e r t h a n l o c a l i t i e s 3, 5, and 6 (Duncan's M u l t i p l e Range T e s t , p<.05). T h e r e i s however s t r o n g e v i d e n c e t h a t b a s a l p a t c h b l a c k n e s s i s a f f e c t e d by d e v e l o p m e n t a l e n v i r o n m e n t , s i n c e v i r t u a l l y a l l t h e r e a r e d s amples a r e s u b s t a n t i a l l y d a r k e r t h a n th e w i l d s a m p l e s ( F i g u r e 1 2 ) . B a s a l T r a n s p a r e n c y Some of t h e v a r i a t i o n i n male b a s a l t r a n s p a r e n c y i s h e r i t a b l e , w i t h r e a r e d s a m p l e s 3, 5, and 6 s i g n i f i c a n t l y l e s s t r a n s p a r e n t t h a n l o c a l i t i e s 2 and 4 (Mann-Whitney U - T e s t , 45 F i g u r e 11. I n h e r i t a n c e o f b a s a l p a t c h w i d t h . V a r i a t i o n between p o p u l a t i o n s f o r r e a r e d ( d a r k b a r s ) and w i l d ( l i g h t b a r s ) s a m p l e s . The s i g n i f i c a n c e l e v e l s of t h e n e s t e d ANOVA f a c t o r s a r e i n d i c a t e d (ns = p>.05). Basal Patch Width Basal Patch Width (X of HW Cell) (% of HW Cell) i t . 47 F i g u r e 12. I n h e r i t a n c e o f b a s a l p a t c h b l a c k n e s s . V a r i a t i o n between p o p u l a t i o n s f o r r e a r e d ( d a r k b a r s ) and w i l d ( l i g h t b a r s ) s a m p l e s . V e r t i c a l b a r s i n d i c a t e one s t a n d a r d e r r o r . The s i g n i f i c a n c e l e v e l s of t h e n e s t e d ANOVA f a c t o r s a r e i n d i c a t e d (ns = p>.05). Basal Patch Color Proportion Black {%) Basal Patch Color Proportion Black(^) 49 p < . 0 1 ) . S i n c e v i r t u a l l y a l l t h e r e a r e d s a m p l e s a r e l e s s t r a n s p a r e n t t h a n t h e w i l d p o p u l a t i o n s , d e v e l o p m e n t a l e n v i r o n m e n t a l s o a f f e c t s b a s a l t r a n s p a r e n c y ( F i g u r e 1 3 ) . D i s t a l B l a c k n e s s T h e r e i s v e r y l i t t l e e v i d e n c e f o r d i s t a l b l a c k n e s s b e i n g h e r i t a b l e , p o s s i b l y b e c a u s e o f r e l a t i v e l y l i t t l e d i f f e r e n c e b e t w een most of t h e w i l d p o p u l a t i o n s ( F i g u r e 1 4 ) . R e a r e d l o c a l i t y 2 m a l e s were l i g h t e r t h a n t h e o t h e r l o c a l i t i e s , a n d r e a r e d l o c a l i t y 6 f e m a l e s were l i g h t e r t h a n t h o s e f r o m l o c a l i t i e s 3, 4, a n d 5 ( D u n c a n ' s M u l t i p l e Range T e s t , p < . 0 5 ) . D i s t a l T r a n s p a r e n c y T h e r e i s a s t r o n g g e n e t i c component t o v a r i a t i o n i n d i s t a l t r a n s p a r e n c y , w i t h s i g n i f i c a n t ' l o c a l i t y ' e f f e c t s f o r b o t h s e x e s ( F i g u r e 1 5 ) . The s i g n i f i c a n t ' e n v i r o n m e n t ' t e r m f o r m a l e s may be due t o e i t h e r a r e a l e f f e c t o f d e v e l o p m e n t a l e n v i r o n m e n t o r t h e e f f e c t o f g e n e t i c d i f f e r e n c e s i n t h e p a r e n t s w h i c h do n o t r e f l e c t t h e d i f f e r e n c e s b e t w e e n w i l d p o p u l a t i o n s . F r o n t Wing L e n g t h V a r i a t i o n i n f r o n t w i n g l e n g t h i s h i g h l y h e r i t a b l e ( F i g u r e 1 6 ) . The l o c a l i t y t e r m i s s i g n i f i c a n t f o r m a l e s , and t h e r e a r 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 r e a r e d f e m a l e s a m p l e s (6 & 2 > 3 > 4; M a n n - W h i t n e y U - T e s t , p < . 0 l ) . S i n c e v i r t u a l l y a l l t h e r e a r e d s a m p l e s a r e s m a l l e r t h a n t h e w i l d p o p u l a t i o n s , u r e 13. I n h e r i t a n c e o f b a s a l t r a n s p a r e n c y . V a r i a t i o n between p o p u l a t i o n s f o r r e a r e d ( d a r k b a r s ) and w i l d ( l i g h t b a r s ) s a m p l e s . V e r t i c a l b a r s i n d i c a t e one s t a n d a r d e r r o r . Basal Patch Color Area Without Scales (%) Basal Patch Color Area Without Scales(/C) 52 F i g u r e 14. I n h e r i t a n c e of d i s t a l b l a c k n e s s . V a r i a t i o n between p o p u l a t i o n s f o r r e a r e d ( d a r k b a r s ) and w i l d ( l i g h t b a r s ) s a m p l e s . V e r t i c a l b a r s i n d i c a t e one s t a n d a r d e r r o r . The s i g n i f i c a n c e l e v e l s of t h e n e s t e d ANOVA f a c t o r s a r e i n d i c a t e d (ns = p>.05). 53 F e m a le L o c a l i t y n.s. E n v i r o n m e n t : n.s. 1 2 3 4 5 6 L o c a l i t y g u r e 15. I n h e r i t a n c e of d i s t a l t r a n s p a r e n c y . V a r i a t i o n between p o p u l a t i o n s f o r r e a r e d ( d a r k b a r s ) and w i l d ( l i g h t b a r s ) s a m p l e s . V e r t i c a l b a r s i n d i c a t e one s t a n d a r d e r r o r . The s i g n i f i c a n c e l e v e l s o f t h e n e s t e d ANOVA f a c t o r s a r e i n d i c a t e d (ns = p>.05). 5 5 M a l e 17-i 1 3 -> O e = I I o L J <D < L o c a l i t y E n v i r o n m e n t : 3 4 L o c a l i t y P< .06 P< .005 F e m a l e L o c a l i t y = p<.03 E n v i r o n m e n t : n.s. 3 4 L o c a l i t y 56 F i g u r e 16. I n h e r i t a n c e o f f r o n t wing l e n g t h . V a r i a t i o n between p o p u l a t i o n s f o r r e a r e d ( d a r k b a r s ) and w i l d ( l i g h t b a r s ) s a m p l e s . V e r t i c a l b a r s i n d i c a t e one s t a n d a r d e r r o r . The s i g n i f i c a n c e l e v e l s of t h e n e s t e d ANOVA f a c t o r s a r e i n d i c a t e d f o r m a l e s . Male v a r i a n c e s were s l i g h t l y non-homogeneous ( B a r t l e t t ' s C h i - S q u a r e d , p>.03), w h i c h was not c o n s i d e r e d t o be g r e a t enough t o s i g n i f i c a n t l y a f f e c t t h e r e s u l t s . 57 1 2 3 4 5 6 L o c a l i t y 58 d e v e l o p m e n t a l e n v i r o n m e n t a l s o a f f e c t s s i z e . T h e r e f o r e , of t h e s i x p h e n o t y p i c c h a r a c t e r s , t h e r e i s s t r o n g e v i d e n c e of a g e n e t i c b a s i s t o v a r i a t i o n i n f o u r , and weak e v i d e n c e f o r t h e o t h e r two c h a r a c t e r s . T h e r e i s a l s o e v i d e n c e f o r d e v e l o p m e n t a l e n v i r o n m e n t a f f e c t i n g p h e n o t y p e f o r t h r e e c h a r a c t e r s . F u r t h e r e v i d e n c e o f an e f f e c t of d e v e l o p m e n t a l e n v i r o n m e n t on p h e n o t y p e c o u l d be o b t a i n e d by c o r r e l a t i n g t h e c h a n g e s i n p h e n o t y p e d u r i n g t h e f l i g h t p e r i o d w i t h an e n v i r o n m e n t a l f a c t o r d u r i n g d e v e l o p m e n t . The a v a i l a b l e knowledge o f t h e l i f e - h i s t o r y o f P a r n a s s i u s phoebus i s l i m i t e d , b u t newly emerged a d u l t s d e f i n i t e l y s p e n t a t l e a s t t h e p r e v i o u s two weeks as pupae. The p h e n o t y p e o f newly emerged males was- n o t s i g n i f i c a n t l y c o r r e l a t e d w i t h t h e mean a i r t e m p e r a t u r e of t h e 7-10 d a y s p r i o r t o emergence (p>.05, n=98). T h e r e f o r e v a r i a t i o n i n p h e n o t y p e o v e r t h e f l i g h t p e r i o d i s not due t o v a r i a t i o n i n t h e mean a i r t e m p e r a t u r e d u r i n g l a t e p u p a l d e v e l o p m e n t . 59 DISCUSSION I n v e s t i g a t i o n s of a l p i n e m e l a n i s m i n b u t t e r f l i e s have f o c u s s e d on t h e genus C o l i a s , a l l t h e s p e c i e s of w h i c h have s i m i l a r v e n t r a l wing s u r f a c e s . In t h e s e s t u d i e s measurements of m e l a n i s m were made on t h e b a s a l a r e a o f t h e v e n t r a l h i n d wing, w h i c h i s a p p a r e n t l y r e p r e s e n t a t i v e of t h e e n t i r e v e n t r a l b a s k i n g s u r f a c e ( J e n s R o l a n d , pers.comm.). However, e a c h o f t h e c o l o r p a t t e r n components of a b u t t e r f l y wing a r e p o t e n t i a l l y a b l e t o v a r y i n d e p e n d e n t l y ( N i j h o u t , 1981). As a r e s u l t , i n i n v e s t i g a t i n g t h e s i g n i f i c a n c e o f v a r i a t i o n i n wing c o l o r i n c o m p l e x l y p a t t e r n e d b u t t e r f l i e s , e a c h wing p a t t e r n component s h o u l d be c o n s i d e r e d s e p a r a t e l y . In P. phoebus i t seemed r e a s o n a b l e t o use an i n t e r m e d i a t e l e v e l of r e s o l u t i o n by a s s u m i n g t h a t t h e v a r i a t i o n i n p a r t of a g r o u p o f m a r k i n g s i s r e p r e s e n t a t i v e of t h e v a r i a t i o n i n t h e e n t i r e g r o u p , as i n E u p h y d r y a s (Mason e t a l . , 1968). S u b j e c t i v e e v a l u a t i o n s s u g g e s t e d t h a t t h e h i n d wing d i s c a l c e l l i s r e p r e s e n t a t i v e of b a s a l h i n d wing c o l o r a t i o n , and t h a t f r o n t wing c e l l s ML and CuJ a r e r e p r e s e n t a t i v e of mean d i s t a l f r o n t wing m e l a n i s m ( m a r g i n a l and s u b m a r g i n a l m a r k i n g s ) i n P a r n a s s i u s p h oebus. V a r i a t i o n i n m e l a n i s m i n P, phoebus i s complex, w i t h o n l y one of t h e c h a r a c t e r s ( b a s a l p a t c h b l a c k n e s s ) a p p r o x i m a t e l y f o l l o w i n g t h e p a t t e r n o f a l p i n e m e l a n i s m f o u n d i n C o l i a s ( R o l a n d , 1978). Even f o r b a s a l p a t c h b l a c k n e s s t h e r e i s an e s s e n t i a l d i f f e r e n c e from C o l i a s b e c a u s e t h e s e x e s a r e d i m o r p h i c , n o t j u s t i n c o l o r a t i o n b u t i n how w e l l t h e v a r i a t i o n i n c o l o r i s c o r r e l a t e d w i t h e l e v a t i o n and o t h e r c h a r a c t e r s 60 between p o p u l a t i o n s . B a s a l p a t c h b l a c k n e s s i s e s s e n t i a l l y t h e same measurement t h a t was us e d f o r i n v e s t i g a t i n g t h e a d a p t i v e s i g n i f i c a n c e of a l p i n e m e l a n i s m i n C o l i a s ( R o l a n d , 1982). I t i s t h e r e f o r e r e a s o n a b l e t o p r e d i c t t h a t v a r i a t i o n i n b a s a l p a t c h b l a c k n e s s i n b o t h s e x e s o f P. phoebus w i l l a f f e c t body t e m p e r a t u r e and a c t i v i t y i n t h e same ways as i n C o l i a s . The o t h e r f o u r measures of m e l a n i s m f o r P. phoebus a r e not c o m p a r a b l e t o any p o s s i b l e measurement f o r C o l i a s . I f t h e c o r r e l a t i o n between b a s a l b l a c k n e s s and e l e v a t i o n i s owing t o t h e a d a p t i v e s i g n i f i c a n c e of i t s e f f e c t on body t e m p e r a t u r e , as i s t h e c a s e i n C o l i a s , t h e n t h e o t h e r b a s a l c h a r a c t e r s s h o u l d a l s o be c o r r e l a t e d w i t h e l e v a t i o n owing t o t h e i r e f f e c t s on body t e m p e r a t u r e . B a s a l p a t c h w i d t h does show a weak c o r r e l a t i o n w i t h e l e v a t i o n i n m a l e s , b ut does not i n f e m a l e s , i n d i c a t i n g t h a t t h e r e i s p r o b a b l y l i t t l e t h e r m o r e g u l a t o r y e f f e c t of t h e n o r m a l v a r i a t i o n i n p a t c h w i d t h . T h i s r e s u l t s u g g e s t s t h a t t h e o u t e r a r e a of t h e b a s a l p a t c h , where change i n w i d t h o c c u r s , has l i t t l e e f f e c t on body t e m p e r a t u r e . I t i s p o s s i b l e t h a t t h e o u t e r edge of t h e b a s a l p a t c h i s u s u a l l y t o o f a r from t h e body t o s i g n i f i c a n t l y a f f e c t body t e m p e r a t u r e . S i n c e t h e b a s a l p a t c h w i d t h i s n o r m a l l y w i d e r i n f e m a l e s t h a n i n m a l e s , one would e x p e c t t h a t v a r i a t i o n i n b a s a l p a t c h w i d t h w i l l s l i g h t l y a f f e c t body t e m p e r a t u r e i n m a l e s , but n o t a t a l l i n f e m a l e s . C o n s e q u e n t l y , v a r i a t i o n i n b a s a l p a t c h w i d t h s h o u l d a f f e c t t h e b e h a v i o r of males but not f e m a l e s . An u n s e a l e d b u t t e r f l y wing r e s u l t s i n a lo w e r body 61 t e m p e r a t u r e t h a n a s c a l e d wing ( D o u g l a s , 1978), i n d i c a t i n g t h a t b a s a l t r a n s p a r e n c y s h o u l d a l s o a f f e c t body t e m p e r a t u r e . I f more s c a l e s r e s u l t i n h i g h e r body t e m p e r a t u r e s , t h e r e s h o u l d be an i n v e r s e c o r r e l a t i o n between b a s a l t r a n s p a r e n c y and e l e v a t i o n . I n s t e a d t h e r e i s a weak p o s i t i v e c o r r e l a t i o n i n t h e o p p o s i t e d i r e c t i o n . T h i s s u g g e s t s t h a t normal v a r i a t i o n i n b a s a l t r a n s p a r e n c y w i l l have o n l y a minor a f f e c t on body t e m p e r a t u r e , and any b e h a v i o r a l e f f e c t s w i l l be m i n o r . M e l a n i s m i n t h e d i s t a l wing a r e a shows a more complex p a t t e r n o f v a r i a t i o n t h a n t h a t i n t h e b a s a l wing a r e a . L o c a l c o r r e l a t i o n s between d i s t a l b l a c k n e s s and e l e v a t i o n a r e c l e a r l y p r e s e n t , b u t t h e r e i s no c o r r e l a t i o n when a l l a r e a s a r e c o n s i d e r e d t o g e t h e r . T h i s i s a p p a r e n t l y owing t o t h e l a r g e v a r i a t i o n i n d i s t a l b l a c k n e s s w i t h i n e l e v a t i o n s when c o m p a r i n g p o p u l a t i o n s between m o u n t a i n s ( T a b l e I V ) . The same p a t t e r n o c c u r s f o r d i s t a l t r a n s p a r e n c y . As a r e s u l t v a r i a t i o n i n d i s t a l b l a c k n e s s and t r a n s p a r e n c y a r e u n l i k e l y t o be t h e r e s u l t o f a d a p t a t i o n t o d i f f e r e n t t h e r m a l e n v i r n o n m e n t s , b e c a u s e a i r t e m p e r a t u r e c h a n g e s w i l l be g r e a t e r between t h a n w i t h i n e l e v a t i o n s f o r m o u n t a i n s n e a r e a c h o t h e r . In f a c t D o u g l a s (1978) h a s shown t h a t even t h e p r e s e n c e of t h e d i s t a l p a r t of t h e wing has no e f f e c t on body t e m p e r a t u r e , and t h e r e f o r e i t i s u n l i k e l y t h a t v a r i a t i o n i n d i s t a l m e l a n i s m has a t h e r m o r e g u l a t o r y s i g n i f i c a n c e . B o t h s e x e s show i n c r e a s e d * d i s t a l b l a c k n e s s and t r a n s p a r e n c y w i t h i n c r e a s e d e l e v a t i o n a t t h e l o c a l l e v e l , r a t h e r t h a n a p o s i t i v e c o r r e l a t i o n f o r f e m a l e s and a n e g a t i v e c o r r e l a t i o n f o r 62 m a l e s a s f o u n d by F e r r i s (1976) and C l i f f o r d ( 1 9 8 1 ) . However, t h e i r 'melanism' i n c l u d e d many s p o t s o t h e r t h a n t h e a r e a w i t h i n my ' d i s t a l m elanism' a r e a , and my d i s t a l m e l a n i s m i s p a r t l y b a s e d on t h e m a r g i n a l m a r k i n g s w h i c h C l i f f o r d d oes not i n c l u d e i n h er a n a l y s i s . F o r t h e m a r k i n g s w h i c h b o t h C l i f f o r d and I i n c l u d e d i n our measurements, t h e s u b m a r g i n a l ' s p o t s ' , she fo u n d no d i f f e r e n c e between h e r t h r e e p o p u l a t i o n s , w h i c h d i f f e r e d c o n s i d e r a b l y i n e l e v a t i o n . S i z e , as i n d i c a t e d by f r o n t wing l e n g t h , i s b e t t e r c o r r e l a t e d w i t h e l e v a t i o n and l a t i t u d e t h a n any o t h e r c h a r a c t e r m easured. Body s i z e of b u t t e r f l i e s a f f e c t s b o t h e q u i l i b r i u m t e m p e r a t u r e s and r a t e s of h e a t i n g and c o o l i n g ( D o u g l a s , 1978), and t h e r e f o r e may have a t h e r m o r e g u l a t o r y s i g n i f i c a n c e . R a p i d h e a t i n g r a t e s ( s m a l l b u t t e r f l i e s ) m ight be a d v a n t a g e o u s under t h e b r o k e n c l o u d c o n d i t i o n s w h i c h f r e q u e n t l y o c c u r a t h i g h e l e v a t i o n s ( D o u g l a s , 1978). H i g h e r e q u i l i b r i u m t e m p e r a t u r e s and s l o w e r c o o l i n g r a t e s ( l a r g e b u t t e r f l i e s ) m i g h t a l s o be a d v a n t a g e o u s a t h i g h e r e l e v a t i o n s , p l a c i n g a lo w e r l i m i t on s i z e a t h i g h e l e v a t i o n s ( D o u g l a s , 1978). T h e r e f o r e I p r e d i c t t h a t v a r i a t i o n i n s i z e i n P. phoebus s h o u l d have an e f f e c t on t h e r m o r e g u l a t i o n , and s h o u l d a l s o a f f e c t a c t i v i t y . However, s i n c e t h e c o r r e l a t i o n between s i z e and a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d i s s u b s t a n t i a l l y p o o r e r t h a n t h e c o r r e l a t i o n w i t h e l e v a t i o n , t h e r e must be a much more i m p o r t a n t f a c t o r t h a n a i r t e m p e r a t u r e a c t i n g t o d e t e r m i n e v a r i a t i o n i n s i z e . The v a r i a t i o n i n any one c h a r a c t e r i s e s s e n t i a l l y i n d e p e n d e n t of t h e v a r i a t i o n i n t h e o t h e r c h a r a c t e r s . The 63 a l m o s t c o m p l e t e l a c k of s i g n i f i c a n t c o r r e l a t i o n s between c h a r a c t e r s w i t h i n i n d i v i d u a l s ( T a b l e V I I I ) i n d i c a t e s t h a t t h e y a r e d e v e l o p m e n t a l l y and g e n e t i c a l l y e n t i r e l y , o r a l m o s t e n t i r e l y , i n d e p e n d e n t . T h e r e a r e few c o r r e l a t i o n s between c h a r a c t e r s f o r mean p o p u l a t i o n p h e n o t y p e s ( T a b l e V I ) , i n d i c a t i n g t h a t w i t h t h e f o l l o w i n g p o s s i b l e e x c e p t i o n s t h e c h a r a c t e r s a r e a l s o e v o l u t i o n a r i l y i n d e p e n d e n t . B a s a l p a t c h w i d t h and b l a c k n e s s a r e w e l l c o r r e l a t e d i n m a l e s , s u p p o r t i n g t h e s u g g e s t i o n t h a t t h e y b o t h a f f e c t male t h e r m o r e g u l a t i o n . They a r e not c o r r e l a t e d i n f e m a l e s , s u g g e s t i n g t h a t f o r f e m a l e s t h e y have d i f f e r e n t a d a p t i v e s i g n i f i c a n c e s . D i s t a l b l a c k n e s s and d i s t a l t r a n s p a r e n c y a r e w e l l c o r r e l a t e d between p o p u l a t i o n s , s u g g e s t i n g t h a t t h e i r v a r i a t i o n has a s i m i l a r a d a p t i v e s i g n i f i c a n c e . The few o t h e r c o r r e l a t i o n s between p o p u l a t i o n s a r e s u f f i c i e n t l y p o or t h a t t h e y p r o b a b l y have r e l a t i v e l y l i t t l e s i m i l a r i t y i n a d a p t i v e s i g n i f i c a n c e . F o r most o f t h e c h a r a c t e r s f e m a l e s a r e more v a r i a b l e between p o p u l a t i o n s t h a n m a l e s , a d i f f e r e n c e w h i c h a l s o o c c u r s i n some o t h e r b u t t e r f l i e s ( P e a r s e and M u r r a y 1982). The c o r r e l a t i o n s w i t h ' e n v i r o n m e n t a l ' v a r i a b l e s ( e l e v a t i o n , l a t i t u d e , and a i r t e m p e r a t u r e ) a r e weaker f o r f e m a l e s t h a n m a l e s , i n c o n t r a s t t o t h e S a t y r i d b u t t e r f l y H e r teronympha merope ( P e a r s e and M u r r a y , 1982), a l t h o u g h t h i s may be t h e r e s u l t of u s i n g t o o few e n v i r o n m e n t a l v a r i a b l e s . Wing l e n g t h , on t h e o t h e r hand, has s i m i l a r c o r r e l a t i o n s w i t h e l e v a t i o n , l a t i t u d e , and a i r t e m p e r a t u r e f o r t h e two s e x e s . T h e r e i s a v e r y s t r o n g c o r r e l a t i o n between t h e s e x e s f o r wing l e n g t h , and i t c h a n g e s a t 64 t h e same r a t e between p o p u l a t i o n s , i n d i c a t i n g t h a t t h e r e l a t i v e s i z e ' of m a l e s and f e m a l e s i s c o n s t a n t . B a s a l p a t c h b l a c k n e s s i s t h e o n l y o t h e r c h a r a c t e r w h i c h shows a good c o r r e l a t i o n between s e x e s , s u g g e s t i n g t h a t e i t h e r i t s t h e r m o r e g u l a t o r y e f f e c t s a r e i m p o r t a n t i n b o t h s e x e s or t h a t t h e g e n e t i c c o n t r o l r e s u l t s i n i t b e i n g e x p r e s s e d i n b o t h s e x e s even i f s e l e c t i o n i s o n l y a c t i n g on one s e x . The r e s u l t s f o r t h e i n h e r i t a n c e of p h e n o t y p e a r e s u f f i c i e n t t o i n d i c a t e t h a t v a r i a t i o n i n most, i f n o t a l l , of t h e c h a r a c t e r s i s i n p a r t g e n e t i c a l l y b a s e d , and i n p a r t owing t o d i f f e r e n t d e v e l o p m e n t a l e n v i r o n m e n t s . I t i s t h e r e f o r e r e a s o n a b l e t o assume t h a t d i f f e r e n c e s i n p h e n o t y p e w i t h i n p o p u l a t i o n s a r e owing t o d i f f e r e n c e s i n b o t h g e n o t y p e and d e v e l o p m e n t a l e n v i r o n m e n t , w i t h many of t h e d i f f e r e n c e s between p o p u l a t i o n s a r i s i n g f r o m l o c a l a d a p t a t i o n s t o e n v i r o n m e n t a l d i f f e r e n c e s . T h e r e a r e a number of e n v i r o n m e n t a l f a c t o r s o t h e r t h a n a i r t e m p e r a t u r e w h i c h m i g h t a c t on t h e p h e n o t y p e of a P a r n a s s i u s  phoebus t o change i t s f i t n e s s . F o r wing c o l o r ( b a s a l and d i s t a l ) , p r e d a t i o n and i n t r a - s p e c i f i c i n t e r a c t i o n s a r e t h e most l i k e l y . F o o d q u a n t i t y , f o o d q u a l i t y , and l e n g t h o f t i m e a v a i l a b l e f o r d e v e l o p m e n t and r e p r o d u c t i o n i n a s e a s o n may a f f e c t s i z e , b o t h t h r o u g h d e v e l o p m e n t a l e f f e c t s and e v o l u t i o n . I p r o p o s e t h e f o l l o w i n g h y p o t h e s e s , b a s e d on t h e p r e c e d i n g c o r r e l a t i o n s and c a s u a l o b s e r v a t i o n s , t o a c c o u n t f o r v a r i a t i o n s i z e and m e l a n i s m : ( 1 ) I n t h e b a s a l wing a r e a o f b o t h s e x e s , n o r m a l v a r i a t i o n i n 65 c o l o r ( a l l components) a f f e c t s body t e m p e r a t u r e . O n l y v a r i a t i o n i n b a s a l p a t c h b l a c k n e s s ( i n b o t h s e x e s ) and v a r i a t i o n i n b a s a l p a t c h w i d t h (males o n l y ) has a s u f f i c i e n t l y l a r g e e f f e c t on body t e m p e r a t u r e t o s i g n i f i c a n t l y a f f e c t i n d i v i d u a l f i t n e s s . (2) l n t h e d i s t a l wing a r e a f o r b o t h s e x e s , c o l o r has no e f f e c t on body t e m p e r a t u r e , and t h e r e f o r e has no e f f e c t t h r o u g h i t on a c t i v i t y . T h e s e f i r s t two h y p o t h e s e s a r e b a s e d on t h e p r e c e d i n g c o r r e l a t i o n s . The f o l l o w i n g s u g g e s t i o n s a r e b a s e d on p u b l i s h e d h y p o t h e s e s o r u n q u a n t i f i e d and f r a g m e n t a r y o b s e r v a t i o n s , and a r e i n t e n d e d as a b a s i s f o r f u t u r e r e s e a r c h . T h e s e h y p o t h e s e s a r e p r i m a r i l y c o n c e r n e d w i t h d i s t a l wing c o l o r , a l t h o u g h t h e y may a l s o h o l d t r u e t o some e x t e n t f o r t h e b a s a l a r e a . (3) S i z e w i l l a f f e c t body t e m p e r a t u r e s , and t h e r e f o r e a c t i v i t y i n b o t h s e x e s ( D o u g l a s , 1978). (4) l t i s a d v a n t a g e o u s f o r b o t h s e x e s t o have a w h i t e g r o u n d c o l o r t o i n c r e a s e a p o s a m a s i s . O b s e r v a t i o n : P. phoebus i s a p p a r e n t l y d i s t a s t e f u l t o b i r d s (Thomas E i s n e r , _iri l i t t . ) , a l t h o u g h my o b s e r v a t i o n s i n d i c a t e t h a t t h e y a r e h i g h l y p a l a t a b l e t o c h i p m u n k s . (5) l t i s a d v a n t a g e o u s f o r m a l e s t o have a w h i t e g r o u n d c o l o r t o i n c r e a s e t h e i r v i s i b i l i t y t o v i r g i n f e m a l e s . O b s e r v a t i o n : P. phoebus m a l e s o n l y c h a s e l i g h t c o l o r e d b u t t e r f l i e s ( C o l i a s , P i e r i s , E u c h l o e , P l e b e j u s , and o t h e r P a r n a s s i u s ) , and n o t d a r k b u t t e r f l i e s ( S p e y e r i a , B o l o r i a , E u p h y d r y a s , and E r e b i a ) ( c a s u a l o b s e r v a t i o n s ) . T h i s i n d i c a t e s t h a t l i g h t n e s s i n wing c o l o r i s a component i n s p e c i e s r e c o g n i t i o n i n t h i s s p e c i e s . 66 (6) I t i s a d v a n t a g e o u s f o r f e m a l e s t o have a d a r k g r o u n d c o l o r t o d e c r e a s e t h e i r v i s i b i l i t y t o m a l e s . O b s e r v a t i o n s : F e m a l e s o n l y mate o n c e , a f t e r w h i c h a l a r g e g e n i t a l p l u g p r e v e n t s f u r t h e r c o p u l a t i o n . M a l e s w i l l a t t e m p t v i o l e n t l y and p r o t r a c t e d l y t o mate w i t h p r e v i o u s l y mated f e m a l e s i f t h e y can f o r c e them t o t h e g r o u n d . The o n l y o b s e r v e d ' s u c c e s s f u l ' s e c o n d m a t i n g r e s u l t e d i n t h e s e c o n d g e n i t a l p l u g b e i n g p l a c e d o v e r t h e o v i p o s i t o r , p r e v e n t i n g o v i p o s t i o n . (7) The c o r r e l a t i o n between t h e d i s t a l wing c h a r a c t e r s and e l e v a t i o n i s t h e r e s u l t o f a c o r r e l a t i o n w i t h an e n v i r o n m e n t a l f a c t o r ( p r e d a t i o n ? ) w h i c h v a r i e s g r e a t l y between m o u n t a i n s , but w i t h i n a m o u n t a i n t e n d s t o be c o r r e l a t e d w i t h e l e v a t i o n . (8) The d e c r e a s e i n s i z e w i t h i n c r e a s e d e l e v a t i o n i s owing t o t h e s h o r t e r d e v e l o p m e n t a l and r e p r o d u c t i v e t i m e a v a i l a b l e . S m a l l e r i n d i v i d u a l s d e v e l o p f a s t e r , l e a v i n g more t i m e t o r e p r o d u c e . O b s e r v a t i o n : At h i g h e l e v a t i o n s ( o v e r 1800m i n s o u t h e r n B.C.) snow f r e q u e n t l y f a l l s b e f o r e a l l f e m a l e s have f i n i s h e d l a y i n g t h e i r e g g s . (9) The d e c r e a s e i n s i z e w i t h i n c r e a s e d e l e v a t i o n may a l s o be owing t o i n c r e a s e d wind v e l o c i t i e s ( D o u g l a s , 1978). (10) L a r g e r s i z e i s a d v a n t a g e o u s owing t o i n c r e a s e d egg and sperm p r o d u c t i o n , and owing t o more p r o l o n g e d a c t i v i t y when p a s s i n g c l o u d s i n t e r r u p t s o l a r h e a t i n g . (11) The r e a s o n f o r t h e p h e n o t y p i c s e x u a l d i m o r p h i s m i n P. phoebus i s t h a t t h e s e x e s i n h a b i t d i f f e r e n t m i c r o h a b i t a t s , and t h e r e f o r e e x p e r i e n c e d i f f e r e n t s e l e c t i v e p r e s s u r e s . 6 7 THE ADAPTIVE SIGNIFICANCE OF ALPINE MELANISM INTRODUCTION The o v e r a l l l e v e l of m e l a n i s m f o r t h e b a c k g r o u n d c o l o r of P a r n a s s i u s phoebus wings i s d e t e r m i n e d by t h e sum of t h e e f f e c t s of a t l e a s t s i x i n d e p e n d e n t c h a r a c t e r s ( C h a p t e r 1). A l l t h e c h a r a c t e r s a r e d e t e r m i n e d i n d e p e n d a n t l y , b o t h on t h e g e n e t i c and d e v e l o p m e n t a l l e v e l , and i t i s l i k e l y t h a t d i f f e r e n t s e l e c t i v e p r e s s u r e s a c t on e a c h c h a r a c t e r . B a s a l p a t c h b l a c k n e s s i s w e l l c o r r e l a t e d w i t h e l e v a t i o n and l a t i t u d e , as w e l l as a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d . The w i d t h of t h e b a s a l b l a c k p a t c h i s p o o r l y c o r r e l a t e d w i t h e l e v a t i o n and l a t i t u d e , b u t s l i g h t l y b e t t e r c o r r e l a t e d w i t h a i r t e m p e r a t u r e . D i s t a l m e l a n i s m i s e s s e n t i a l l y u n c o r r e l a t e d w i t h e l e v a t i o n when a l l m o u n t a i n s a r e c o n s i d e r e d t o g e t h e r , but i s w e l l c o r r e l a t e d w i t h e l e v a t i o n on some i n d i v i d u a l m o u n t a i n s . S i z e shows t h e b e s t c o r r e l a t i o n w i t h e l e v a t i o n , but i s more p o o r l y c o r r e l a t e d w i t h a i r t e m p e r a t u r e d u r i n g t h e f l i g h t p e r i o d . The o b s e r v e d v a r i a t i o n , a t l e a s t between p o p u l a t i o n s , i s a p p a r e n t l y due t o b o t h g e n e t i c d i f f e r e n c e s and d i f f e r e n c e s i n d e v e l o p m e n t a l e n v i r o n m e n t . From t h e s e o b s e r v a t i o n s I d e v e l o p e d a number o f h y p o t h e s e s as t o t h e a d a p t i v e s i g n i f i c a n c e o f v a r i a t i o n i n t h e components of m e l a n i s m i n P. phoebus, t h r e e of which were t e s t e d and a r e d e s c r i b e d i n t h i s c h a p t e r . ( 1 ) I n t h e b a s a l wing a r e a of b o t h s e x e s , c o l o r ( a l l components) a f f e c t s body t e m p e r a t u r e . O n l y b a s a l p a t c h b l a c k n e s s ( i n b o t h 68 s e x e s ) and b a s a l p a t c h w i d t h ( m a l e s o n l y ) has a s u f f i c i e n t l y l a r g e e f f e c t on body t e m p e r a t u r e t o s i g n i f i c a n t l y a f f e c t i n d i v i d u a l f i t n e s s . The f o l l o w i n g p r e d i c t i o n s f o l l o w from t h i s h y p o t h e s i s: (a) I n d i v i d u a l s w i t h d a r k e r b a s a l wing a r e a s have h i g h e r body t e m p e r a t u r e s under e q u i v a l e n t e n v i r o n m e n t a l c o n d i t i o n s . Body t e m p e r a t u r e w i l l be p o s i t i v e l y c o r r e l a t e d w i t h b a s a l p a t c h w i d t h and b a s a l p a t c h b l a c k n e s s , and n e g a t i v e l y c o r r e l a t e d w i t h b a s a l t r a n s p a r e n c y . (b) At low a i r t e m p e r a t u r e s a n d / o r low l e v e l s of s o l a r r a d i a t i o n , m ales w i t h g r e a t e r b a s a l p a t c h w i d t h and b l a c k n e s s w i l l have g r e a t e r f l i g h t a c t i v i t y , f e e d i n g , and m a t i n g s u c c e s s . D a r k e r m a l e s w i l l spend l e s s t i m e b a s k i n g . A c t i v i t y of f e m a l e s w i l l a l s o be c o r r e l a t e d t o b a s a l p a t c h b l a c k n e s s , but n o t t o n o r m a l v a r i a t i o n i n b a s a l p a t c h w i d t h . Normal v a r i a t i o n i n b a s a l t r a n s p a r e n c y w i l l not a f f e c t a c t i v i t y i n e i t h e r s e x . (c) At h i g h e r a i r t e m p e r a t u r e s a n d / o r s o l a r r a d i a t i o n s m a l e s w i t h g r e a t e r b a s a l p a t c h b l a c k n e s s and w i d t h w i l l t e n d t o o v e r h e a t and f l i g h t a c t i v i t y , f e e d i n g , and r e p r o d u c t i o n w i l l d e c r e a s e , w h i l e ' n o n - p r o d u c t i v e ' h e a t a d v o i d a n c e b e h a v i o r s w i l l i n c r e a s e . ' N o n - p r o d u c t i v e ' h e a t a d v o i d a n c e a c t i v i t i e s a r e t h o s e w h i c h i n c r e a s e s u r v i v a l w h i l e r e d u c i n g immediate r e p r o d u c t i v e o p p o r t u n i t i e s , and may i n c l u d e s h a d e - s e e k i n g , v e r t i c a l b a s k i n g , o r i n c r e a s e d n e c t a r or water i n t a k e . T h e s e p r e d i c t i o n s w i l l h o l d t r u e f o r f e m a l e b a s a l p a t c h b l a c k n e s s , but n o t f o r n o r m a l v a r i a t i o n i n 69 b a s a l p a t c h w i d t h . Normal v a r i a t i o n i n b a s a l t r a n s p a r e n c y w i l l have no e f f e c t f o r e i t h e r s e x . ( d ) M a l e s w i t h g r e a t e r b a s a l p a t c h b l a c k n e s s or b a s a l p a t c h w i d t h w i l l have a g r e a t e r p r o b a b i l i t y of d i s p e r s i n g , b e c a u s e of t h e i r i n c r e a s e d a c t i v i t y . In f e m a l e s o n l y b a s a l p a t c h b l a c k n e s s w i l l a f f e c t d i s p e r s a l r a t e s , and n o r m a l v a r i a t i o n i n b a s a l p a t c h w i d t h w i l l n o t . Normal v a r i a t i o n i n b a s a l t r a n s p a r e n c y w i l l have no d e t e c t a b l e e f f e c t on d i s p e r s a l i n e i t h e r s e x . 2. V a r i a t i o n i n d i s t a l t r a n s p a r e n c y and d i s t a l b l a c k n e s s w i l l have no t h e r m a l a d a p t i v e s i g n i f i c a n c e . The a d a p t i v e s i g n i f i c a n c e of v a r i a t i o n i n t h e s e c h a r a c t e r s i s p r o b a b l y due t o t h e o n l y o t h e r two g r o u p s of f a c t o r s w h i c h a r e l i k e l y t o i n t e r a c t w i t h wing c o l o r , i n t r a s p e c i f i c i n t e r a c t i o n s and p r e d a t i o n . The p r e d i c t i o n s a r e : (a) d i s t a l t r a n s p a r e n c y and b l a c k n e s s w i l l have no e f f e c t on body t e m p e r a t u r e i n e i t h e r s e x . T h i s p r e d i c t i o n does not need t o be t e s t e d f o r P. phoebus s p e c i f i c a l l y b e c a u s e i t has been f o u n d t o be t r u e f o r b u t t e r f l i e s i n g e n e r a l by D o u g l a s ( 1 9 7 8 ) . (b) d i s t a l t r a n s p a r e n c y and b l a c k n e s s i n e i t h e r sex w i l l have no e f f e c t on c h a n g e s i n a c t i v i t y t h r o u g h e f f e c t s on body t e m p e r a t u r e . ( 3 ) V a r i a t i o n i n . s i z e has a t h e r m o r e g u l a t o r y a d a p t i v e s i g n i f i c a n c e i n b o t h s e x e s . The p r e d i c t i o n s a r e : ( a ) I n b o t h s e x e s l a r g e r b u t t e r f l i e s w i l l have h i g h e r e q u i l i b r i u m body t e m p e r a t u r e s and l o w e r h e a t i n g and c o o l i n g 70 r a t e s . T h i s h y p o t h e s i s a l s o does n o t need t o be s p e c i f i c a l l y t e s t e d f o r P. phoebus b e c a u s e i t has been f o u n d t o be t r u e f o r b u t t e r f l i e s i n g e n e r a l by D o u g l a s (1978) . ( b ) l n b o t h s e x e s l a r g e r b u t t e r f l i e s w i l l have g r e a t e r f l i g h t a c t i v i t y , and f e m a l e s w i l l have i n c r e a s e d f e c u n d i t y , but l a r g e r b u t t e r f l i e s w i l l a l s o have a g r e a t e r p r o b a b i l i t y of d i s p e r s i n g b e c a u s e of t h e i n c r e a s e d f l i g h t a c t i v i t y . In t h i s c h a p t e r I c o n c e n t r a t e on t e s t s of t h e t h e r m o r e g u l a t o r y h y p o t h e s e s of t h e a d a p t i v e s i g n i f i c a n c e of v a r i a t i o n i n e a c h p h e n o t y p i c c h a r a c t e r , a l t h o u g h I have not been a b l e t o t e s t a l l the p r e d i c t i o n s . Many of t h e f i e l d methods and methods of a n a l y s i s a r e s i m i l a r t o t h o s e u s e d f o r s t u d y i n g t h e a d a p t i v e s i g n i f i c a n c e o f a l p i n e m e l a n i s m i n C o l i a s n a s t e s and C. meadi i by R o l a n d (1982) i n o r d e r t o f a c i l i t a t e c o m p a r i s o n s . 71 MATERIALS AND METHODS Study S i t e s Two p o p u l a t i o n s of P a r n a s s i u s phoebus n e a r V e r n o n , B r i t i s h C o l u m b i a were u s e d f o r t h i s s t u d y ( s e e A p p e n d i x 1 f o r l o c a t i o n d e t a i l s ) : 1. V e r n o n H i l l , V e r n o n , B.C., e l . 1l00-1200m: In t h i s a r e a a number o f d r y g r a s s y s l o p e s o c c u r a t t h e t r a n s i t i o n from t h e low e l e v a t i o n g r a s s l a n d s t o t h e h i g h e r e l e v a t i o n f o r e s t e d a r e a s e a s t of t h e town of V e r n o n . The main s t u d y p o p u l a t i o n ("Area 2") i s on a s t e e p , t h i n l y v e g e t a t e d , r o c k r u b b l e and d i r t s o u t h - f a c i n g s l o p e . On t h e e a s t s i d e of t h e main s l o p e i s a s m a l l a r e a ("Area 3") a l m o s t e n t i r e l y s e p a r a t e d from t h e main s l o p e by t r e e s and s h r u b s . On t h e west s i d e i s a n o t h e r s m a l l a r e a ("Area 1") s e p a r a t e d from t h e main s l o p e by a narrow band (5 m e t e r s wide) of c o n i f e r o u s t r e e s . T h e s e t h r e e a r e a s a r e c o m p l e t e l y s u r r o u n d e d by c o n i f e r o u s f o r e s t e x c e p t f o r a narrow (4 meter) o p e n i n g a t t h e b o t t o m o f t h e west s l o p e w h i c h c o n n e c t s t o e x t e n s i v e l o w e r g r a s s l a n d s w h i c h do n o t s u p p o r t a P. phoebus p o p u l a t i o n . The t h r e e a r e a s a r e e s s e n t i a l l y an i s l a n d i n t h e m i d d l e o f f o r e s t . The main s l o p e c o v e r s a b o u t 9 h e c t a r e s , of w h i c h o n e - h a l f c o m p r i s e s t h e b r e e d i n g a r e a f o r P. p h o e b u s . On t h e upper s i d e o f t h e s e t h r e e a r e a s t h e f o r e s t i s c o n t i n u o u s u n t i l t h e open summit of S i l v e r S t a r Mt., 15 km away. The l o w e r s i d e of t h e a r e a s i s i s d e t e r m i n e d by a band of f o r e s t a b o u t 200m ( s t r a i g h t l i n e ) w ide, on t h e o t h e r s i d e of w h i c h i s more h a b i t a t f o r P. p h oebus. I s a m p l e d s e v e r a l n e a r b y p o r t i o n s o f 72 t h a t P. phoebus h a b i t a t , w h i c h a r e d e s i g n a t e d A r e a s 4-7. The t o t a l p o p u l a t i o n s i z e o f V e r n o n H i l l i s p r o b a b l y s e v e r a l t h o u s a n d i n d i v i d u a l s , a l t h o u g h t h e s t u d y a r e a s c o n t a i n o n l y a p o r t i o n of t h e p o p u l a t i o n . 2. S i l v e r S t a r Mt., V e r n o n , B.C., e l . 1700-I800m: The summit of S i l v e r S t a r Mt. has an open s o u t h s l o p e about 100 h e c t a r e s i n a r e a and s u r r o u n d e d by f o r e s t . Of t h a t a r e a , one r o c k y r i d g e (2-3 h e c t a r e s ) i s u s e d by P. phoebus, w i t h a s e c o n d r i d g e o f a b o u t h a l f t h a t a r e a u s e d s p o r a d i c a l l y by a s m a l l p o r t i o n of t h e p o p u l a t i o n . The two r i d g e s a r e s e p a r a t e d by a r a v i n e 200~300m wide f i l l e d w i t h s h r u b s and s c a t t e r e d low t r e e s . The r i d g e s a r e m o s t l y l o o s e r o c k w i t h v e r y s p a r s e , low v e g e t a t i o n . The t o t a l p o p u l a t i o n s i z e was not measured, but i s p r o b a b l y about 300 i n d i v i d u a l s . The n e a r e s t g r a s s l a n d h a b i t a t i s a b o u t 10 km away a t 1200m e l e v a t i o n , and t h e n e a r e s t known permanent p o p u l a t i o n of P. phoebus i s t h e V e r n o n H i l l p o p u l a t i o n d e s c r i b e d a b o v e . The c o r r e l a t i o n s o f p h e n o t y p e w i t h e l e v a t i o n f o r V e r n o n H i l l t o t h e summit of S i l v e r S t a r Mt. a r e g i v e n i n T a b l e V, w i t h t h e means f o r S i l v e r S t a r Mt. g i v e n i n T a b l e IV ( C h a p t e r 1 ) . M e l a n i s m and Body T e m p e r a t u r e The e f f e c t of b a s a l wing m e l a n i s m on body t e m p e r a t u r e was d e t e r m i n e d u s i n g a method s i m i l a r t o R o l a n d ' s ( 1 9 8 2 ) . The e f f e c t of d i s t a l wing m e l a n i s m was n o t measured b e c a u s e i t has been w e l l e s t a b l i s h e d t h a t t h e d i s t a l wing a r e a has no e f f e c t on the- body t e m p e r a t u r e of d o r s a l b a s k i n g b u t t e r f l i e s , whether 73 a l i v e o r dead ( D o u g l a s , 1978). Dead, d r i e d b u t t e r f l i e s w i t h t h e wings i n d o r s a l b a s k i n g p o s i t i o n were u s e d , w i t h a l l s p e c i m e n s b e i n g s i z e - m a t c h e d m a l e s . The e f f e c t of s i z e was a l s o n o t measured s i n c e i t has been w e l l e s t a b l i s h e d ( D o u g l a s , 1978). T h e r m o c o u p l e s were i n s e r t e d i n t o 4 d r i e d abdomens, and t h e a bdomen/thermocouple c o m b i n a t i o n s u s e d t o r e p l a c e t h e abdomens of t h e t e s t s p e c i m e n s . The new abdomens were a t t a c h e d w i t h a s m a l l d r o p o f h o u s e h o l d w h i t e g l u e . The t h e r m o c o u p l e l e a d s , p l u s one c a l i b r a t i o n t h e r m o c o u p l e , were a t t a c h e d t o a f i v e - w a y s i n g l e - p o l e s w i t c h c o n n e c t e d t o a Doran M i n i T h e r m o c o u p l e P o t e n t i o m e t e r Model E422. The b u t t e r f l i e s were s u s p e n d e d i n m i d - a i r by t h e t h e r m o c o u p l e l e a d , a n g l e d t o w a r d s t h e p o s i t i o n of t h e sun a t s o l a r noon, w i t h t h e c a l i b r a t i o n t h e r m o c o u p l e immersed i n an i c e b a t h a t 0°C. A i r t e m p e r a t u r e , s o l a r r a d i a t i o n ^ and body t e m p e r a t u r e were r e c o r d e d from 30 m i n u t e s b e f o r e dawn t o 30 m i n u t e s a f t e r s u n s e t e v e r y 15 m i n u t e s . T h e s e measurements were made on 3 s e t s of 4 b u t t e r f l i e s , u s i n g t h e same abdomen/thermocouple c o m b i n a t i o n s e a c h t i m e . S o l a r r a d i a t i o n was m easured u s i n g a Weather M e a s u r e Model R401 m e c h a n i c a l p y r a n o m e t e r , and a i r t e m p e r a t u r e was measured u s i n g a thermometer s u s p e n d e d i n s h a d e . Wind s p e e d was v e r y low a t a l l t i m e s . M u l t i p l e r e g r e s s i o n a n a l y s i s was u s e d t o d e t e r m i n e t h e e f f e c t s of s o l a r r a d i a t i o n , a i r t e m p e r a t u r e , b a s a l p a t c h w i d t h , b a s a l p a t c h d a r k n e s s , and b a s a l t r a n s p a r e n c y on body t e m p e r a t u r e . In o r d e r t o e l i m i n a t e e r r o r due t o d i f f e r e n c e s between t h e h e a t i n g c h a r a c t e r i s t i c s o f t h e 4 abdomens u s e d , t h e y 74 were i n c l u d e d a s dummy v a r i a b l e s i n t h e r e g r e s s i o n a n a l y s i s . Minimum Body T e m p e r a t u r e R e q u i r e d f o r F l i g h t The body t e m p e r a t u r e s o f t w e l v e males and one f e m a l e were t a k e n by t h e 'gr a b and s t a b ' method ( D o u g l a s , 1978) d u r i n g t h e o n e - h a l f hour a f t e r t h e f i r s t i n d i v i d u a l was seen i n f l i g h t . As e a c h s p e c i m e n a p p e a r e d i n f l i g h t i t was n e t t e d and h e l d by t h e wings i n t h e n e t w h i l e a c o p p e r c o n s t a n t a n h y p o d e r m i c t h e r m o c o u p l e was i n s e r t e d v e n t r a l l y t o t h e c e n t e r of t h e t h o r a x , and t h e t e m p e r a t u r e r e c o r d e d on t h e p o t e n t i o m e t e r d e s c r i b e d above. The e n t i r e p r o c e d u r e t o o k l e s s t h an 20 s e c o n d s . Two s p e c i m e n s w h i c h were c a p t u r e d r i g h t b e s i d e t h e p o t e n t i o m e t e r were o b s e r v e d f o r t h e p e r i o d f r o m 10 s e c o n d s t o 60 s e c o n d s a f t e r c a p t u r e , t o d e t e r m i n e r a t e of d e c r e a s e i n body t e m p e r a t u r e a f t e r c a p t u r e . An e l e c t r o n i c c o l d - j u n c t i o n (Omega MCJ) was u s e d f o r c a l i b r a t i o n . The a i r t e m p e r a t u r e was a c o n s t a n t 15 d e g r e e s t h r o u g h o u t , t h e sky was c l e a r , and t h e r e was no win d . Most of t h e c a p t u r e s were made l e s s t h a n 3 m i n u t e s a f t e r an i n d i v i d u a l i n i t i a t e d f l i g h t , and a l l s p e c i m e n s showed c o n t r o l l e d f l i g h t f o r a t l e a s t 30 s e c o n d s b e f o r e c a p t u r e . The measurements were made between 0625-0655 PST a t 1800m nea r C o r n w a l l L o o k o u t (Cache C r e e k ) , B.C. i n e a r l y A u g u s t 1984. 75 D e t e r m i n a t i o n of F l i g h t T h r e s h o l d s D e t e r m i n a t i o n of t h e c o m b i n a t i o n s of a i r t e m p e r a t u r e and s o l a r r a d i a t i o n under w h i c h f l i g h t o c c u r r e d was done u s i n g t h e method of R o l a n d ( 1 9 8 2 ) , a t t h e V e r n o n H i l l s t u d y s i t e i n J u n e and J u l y 1983. S pecimens were c a p t u r e d i n f l i g h t , t h e t i m e r e c o r d e d , and t h e n t h e y were marked w i t h an i n d e l i b l e f e l t t i p pen on t h e v e n t r a l r e a r m a r g i n of t h e f r o n t wing w i t h an i d e n t i f y i n g number. They were t h e n d r o p p e d o n t o a p i e c e of s t y r o f o a m b o a r d c o v e r e d i n 'bubble p a c k ' , w h i c h i n d u c e d them t o open t h e i r wings i n t o a wide s p r e a d p o s i t i o n (as a d e f e n s i v e r e a c t i o n ) . When a s q u a r e of p l a t e g l a s s was p l a c e d on t o p , t h e body of t h e b u t t e r f l y was between t h e b u b b l e s o f t h e b u b b l e pack and t h e wings were h e l d between t h e g l a s s and t h e b u b b l e s . T hus t h e b u t t e r f l y was i m m o b i l i z e d w i t h o u t any damage. The p h e n o t y p e measurements d e s c r i b e d i n C h a p t e r 1 were t h e n made, and t h e b u t t e r f l y r e l e a s e d a g a i n . I f t h e b u t t e r f l y was a l r e a d y marked and measured, i t was s i m p l y r e l e a s e d a g a i n a f t e r r e c o r d i n g t i m e of c a p t u r e . B e f o r e r e l e a s e , t h e b u t t e r f l i e s were c o o l e d i n t h e shade and r e l e a s e d i n t h e base of h e r b a c e o u s v e g e t a t i o n i n o r d e r t o f o r c e them t o c r a w l up t o t h e t o p of t h e v e g e t a t i o n b e f o r e a t t e m p t i n g f l i g h t . T h i s p r e v e n t e d any ' p a n i c ' e s c a p e f l i g h t upon r e l e a s e , and a l l i n d i v i d u a l s a p p e a r e d t o r e j o i n t h e p o p u l a t i o n w i t h n ormal b e h a v i o r w i t h i n 5 m i n u t e s of r e l e a s e . 76 E f f e c t of P h e n o t y p e and E n v i r o n m e n t on A c t i v i t y In e a r l y A u g u st 1982 a t t h e S i l v e r S t a r Mt. c o l o n y , i n d i v i d u a l s p e c i m e n s were f o l l o w e d f o r 10 m i n u t e i n t e r v a l s , w h i l e a l l a c t i v i t i e s and a r a n k e d e s t i m a t e of wind s p e e d (none, g e n t l e , s t r o n g ) were r e c o r d e d on t a p e . A i r t e m p e r a t u r e and s o l a r r a d i a t i o n were r e c o r d e d s e p a r a t e l y u s i n g a thermometer and r e c o r d i n g m e c h a n i c a l p y r a n o m e t e r . O n l y p e r i o d s i n w h i c h t h e r e was no v i s i b l e e f f e c t o f wind on a c t i v i t y , and i n w h i c h t h e r e were no c l o u d s , were u s e d . A l l s p e c i m e n s were i n f l i g h t a t t h e s t a r t of e a c h o b s e r v a t i o n p e r i o d , and were c a p t u r e d a t t h e end of t h e t e n m i n u t e s f o r l a t e r s c o r i n g o f p h e n o t y p e . B e f o r e a n a l y s i s e a c h e n v i r o n m e n t a l and p h e n o t y p i c v a r i a b l e was d i v i d e d i n t o ' h i g h ' and 'low' c l a s s e s a t t h e median. The sample s i z e f o r f e m a l e s was t o o s m a l l f o r u s e f u l a n a l y s i s . An a n a l y s i s of v a r i a n c e was used t o d e t e r m i n e t h e e f f e c t of t h e t h r e e e n v i r o n m e n t a l and two p h e n o t y p i c v a r i a b l e s on a v e r a g e t i m e s p e n t i n f l i g h t , t i m e s p e n t f e e d i n g , and t i m e s p e n t b a s k i n g w i t h o u t f e e d i n g . The p h e n o t y p i c c h a r a c t e r s were t e s t e d two a t a t i m e , f i r s t b a s a l p a t c h w i d t h and d a r k n e s s , t h e n b a s a l t r a n s p a r e n c y and f r o n t wing l e n g t h , and t h e n d i s t a l b l a c k n e s s and t r a n s p a r e n c y . O n l y t h e main f a c t o r s and t h e i r f i r s t o r d e r i n t e r a c t i o n s were i n c l u d e d i n t h e a n a l y s i s due t o t h e s m a l l sample s i z e (32 i n d i v i d u a l s ) . 77 P o p u l a t i o n P a r a m e t e r s A t t h e V e r n o n H i l l s t u d y s i t e ( A r e a 2 ) , I p e r i o d i c a l l y c o l l e c t e d , numbered, and s c o r e d t h e p h e n o t y p e of as many i n d i v i d u a l s as p o s s i b l e , i n a d d i t i o n t o t h e i n f o r m a t i o n c o l l e c t e d f o r d e t e r m i n a t i o n of f l i g h t t h r e s h o l d s . T h i s i n c r e a s e d t h e number o f r e c a p t u r e s p e r day, r e s u l t i n g i n i n c r e a s e d p r e c i s i o n i n t h e e s t i m a t e s of p o p u l a t i o n s i z e , s u r v i v a l r a t e s , and number of m a l e s j o i n i n g t h e p o p u l a t i o n . The f i n a l sample was t a k e n on J u l y 18, a f t e r w h i c h i n c l e m e n t w eather p r e v e n t e d f u r t h e r b u t t e r f l y a c t i v i t y f o r t h e n e x t 10 d a y s . No P a r n a s s i u s phoebus were seen i n f l i g h t on t h e n e x t sunny day ( J u l y 2 9 ) . D i s p e r s a l _ . I n f o r m a t i o n on d i s p e r s a l was c o l l e c t e d on V e r n o n H i l l i n 1983. P r e l i m i n a r y o b s e r v a t i o n s i n d i c a t e d t h a t i n a 10-15 m i n u t e i n t e r v a l , a male would f l y o v e r a t l e a s t h a l f o f A r e a 2, and f r e q u e n t l y much more. I assumed t h e r e f o r e t h a t e a c h of t h e s e v e n a r e a s were e s s e n t i a l l y p a n m i c t i c on a d a i l y b a s i s , s i n c e a l l were w i t h o u t b a r r i e r s and a l l ( e x c e p t a r e a 7) were s m a l l e r t h a n A r e a 2. Measurement o f movement w i t h i n an a r e a was t h e r e f o r e p o i n t l e s s , and o n l y a r e a of c a p t u r e and n o t p o i n t of c a p t u r e was r e c o r d e d . Most s a m p l i n g was done i n A r e a 2 as p a r t of t h e s t u d i e s a l r e a d y d e s c r i b e d . On June 22 A r e a s 1, 2, and 3 were sampled i n t e n s i v e l y . On June 28 and J u l y 18 a l l s e v e n a r e a s were sampl e d i n t e n s i v e l y , e x c e p t f o r a r e a 7 on June 28 and 78 A r e a 6 on J u l y 18. A l l i n d i v i d u a l s c a p t u r e d were marked w i t h an i d e n t i f y i n g number ( i f not a l r e a d y marked) as d e s c r i b e d p r e v i o u s l y , and r e l e a s e d a g a i n i n t h e same a r e a i n w h i c h t h e y were c a p t u r e d . 79 RESULTS B a s a l M e l a n i s m and Body T e m p e r a t u r e B a s a l p a t c h w i d t h and b a s a l p a t c h b l a c k n e s s a r e b o t h p o s i t i v e l y c o r r e l a t e d w i t h a b d o m i n a l t e m p e r a t u r e i n dead b u t t e r f l i e s , and b a s a l t r a n s p a r e n c y shows a weak n e g a t i v e c o r r e l a t i o n ( T a b l e X ) . S i n c e t h e r e l a t i v e e f f e c t s o f wing c o l o r on t h e e q u i l i b r i u m t e m p e r a t u r e s o f l i v e and d e a d b u t t e r f l i e s a r e th e same ( D o u g l a s , 1978), t h i s c o n c l u s i o n i s a p p l i c a b l e t o l i v e b u t t e r f l i e s . Minimum Body T e m p e r a t u r e f o r F l i g h t The minimum body t e m p e r a t u r e f o r - m a l e s i n f l i g h t was 17.6°C, w i t h t h e r a n g e of t e m p e r a t u r e s d u r i n g t h e f i r s t h a l f hour a f t e r t h e f i r s t i n d i v i d u a l was o b s e r v e d i n f l i g h t i n t h e m o r n i n g b e i n g 17.7-20.1°C (mean=!9.1; s.e.=0.22; n=12). O n l y a s i n g l e f e m a l e was m e asured, h e r body t e m p e r a t u r e was 17.4°C, s u g g e s t i n g t h a t minimum f l i g h t t e m p e r a t u r e s f o r f e m a l e s a r e a t l e a s t as low as f o r m a l e s . The body t e m p e r a t u r e s were 3-5 d e g r e e s above t h e a i r t e m p e r a t u r e of 15°C. The body t e m p e r a t u r e s of two m a l e s were r e c o r d e d from 10 s e c o n d s t o 60 s e c o n d s a f t e r c a p t u r e and l o s t 0.7 and 0.8 d e g r e e s i n t h a t t i m e . T h e r e f o r e t h e e r r o r on t h e o b s e r v e d t e m p e r a t u r e s i s under 0.5°C. S i n c e f e m a l e s have v e r y l i t t l e h a i r on t h e i r b o d i e s compared t o m a l e s , a g r e a t e r d r o p i n body t e m p e r a t u r e c o u l d have o c c u r r e d between c a p t u r e and measurement of t h e f e m a l e t e m p e r a t u r e t h a n 80 T a b l e X. B a s a l M e l a n i s m and Body T e m p e r a t u r e . M u l t i p l e r e g r e s s i o n o f body t e m p e r a t u r e (°C) as a f u n c t i o n o f s o l a r r a d i a t i o n ( g m . c a l / c m 2 / m i n ) , ambient t e m p e r a t u r e ( ° C ) , b a s a l p a t c h w i d t h , b a s a l p a t c h b l a c k n e s s , and b a s a l t r a n s p a r e n c y . V a r i a b l e P a r t i a l R e g r e s . C o e f f . P a r t i a l C o r r . C o e f f . P.phoebus C . n a s t e s * P.phoebus C . n a s t e s * S o l a r R a d i a t i o n A i r T e m p e r a t u r e B a s a l B l a c k n e s s B a s a l W i d t h B a s a l T r a n s p a r . C o n s t a n t 36.76 17.45*** 1.03 .625 .832** (.047) 1.43** -.473** -25.75 .776 .508 .615 .395 .240** .362 .263** -.109** P . phoebus r2=0.825; n=444; a l l c o e f f i c i e n t s p < . 0 0 0 l , e x c e p t f o r b a s a l t r a n s p a r e n c y , p<.01. C . n a s t e s r2=0.798; n=l56; a l l c o e f f i c i e n t s p<.00! * V a l u e s f o r C o l i a s n a s t e s from R o l a n d (1982) a r e i n c l u d e d f o r c o m p a r i s o n . * * P h e n o t y p e v a r i a b l e s f o r phoebus a r c s i n t r a n s f o r m e d ( C h a p t e r 1) t o i n s u r e n o r m a l i t y a r o u n d e a c h d a t a p o i n t i n t h e r e g r e s s i o n . ***The r e g r e s s i o n c o e f f i c i e n t f o r s o l a r r a d i a t i o n i n R o l a n d (1982) was a c t u a l l y ' i n w a t t s / m 2 r a t h e r t h a n t h e gm.cal/cm 2/min s t a t e d ( R o l a n d , pers.comm.). In t h i s t a b l e I have c o n v e r t e d t h e v a l u e t o gm.cal/cm 2/min (1 gm.cal/cm 2/min = 697.8 W/m 2). The d a t a f o r b a s a l b l a c k n e s s i n R o l a n d were u n t r a n s f o r m e d , t h e r e f o r e t h e r e g r e s s i o n c o e f f i c i e n t s a r e not d i r e c t l y c o m p a r a b l e f o r t h a t t e r m . o c c u r r e d f o r t h e m a l e s . Minimum F l i g h t T h r e s h o l d The l e v e l s of s o l a r r a d i a t i o n and a i r t e m p e r a t u r e above w h i c h f l i g h t o c c u r s a r e shown i n F i g u r e s 17-18. The minimum f l i g h t t h r e s h o l d i s c u r v e d and r e s u l t s f r o m t h e combined e f f e c t s of a i r t e m p e r a t u r e and s o l a r r a d i a t i o n . Below 0.2 gm.cal/cm 2/min s o l a r r a d i a t i o n a l l f l i g h t c e a s e s r e g a r d l e s s of a i r t e m p e r a t u r e . The f e m a l e f l i g h t t h r e s h o l d may be t h e same as 81 t h a t o f t h e male, a l t h o u g h i t was n o t p o s s i b l e t o be a b s o l u t e l y c e r t a i n of when f e m a l e f l i g h t began or ended. The l a c k of f e m a l e c a p t u r e s a t h i g h a i r t e m p e r a t u r e s i s t h e r e s u l t o f t h e r e b e i n g v e r y l i t t l e s a m p l i n g t i m e a t t h a t t e m p e r a t u r e . F l i g h t w i l l n e v e r o c c u r below 4°C a t t h e l a t i t u d e of V e r n o n , B.C. b e c a u s e s o l a r r a d i a t i o n i s n e v e r s u f f i c i e n t l y h i g h t o compensate f o r t e m p e r a t u r e s below t h a t . When a minimum f l i g h t t e m p e r a t u r e of 18°C i s us e d i n t h e body t e m p e r a t u r e r e g r e s s i o n of T a b l e X, a minimum f l i g h t t h r e s h o l d c an be p r e d i c t e d f o r b u t t e r f l i e s w i t h d i f f e r e n t l e v e l s of b a s a l m e l a n i s m ( F i g u r e 19). Th e s e p r e d i c t e d t h r e s h o l d s a r e a l l w e l l below t h e a c t u a l t h r e s h o l d . When t h e c a p t u r e s of males a r e s p l i t i n t o l i g h t and d a r k g r o u p s a l o n g t h e median f o r b a s a l p a t c h b l a c k n e s s ( F i g u r e 20) o r b a s a l p a t c h w i d t h ( F i g u r e 21) t h e r e i s no a p p a r e n t dependence of f l i g h t t h r e s h o l d on wing c o l o r . Even when o n l y i n d i v i d u a l s w h i c h were l i g h t f o r b o t h c h a r a c t e r s o r d a r k f o r b o t h c h a r a c t e r s a r e c o n s i d e r e d t h e r e i s no e f f e c t of wing c o l o r on f l i g h t t h r e s h o l d ( F i g u r e 2 2 ) . I c o n s i d e r e d t h e p o s s i b i l i t y o f a g r a d u a l change i n mean m e l a n i s m of t h e i n d i v i d u a l s i n f l i g h t as a f u n c t i o n o f d i s t a n c e from t h e t h r e s h o l d . F i r s t I d i d a m u l t i p l e r e g r e s s i o n of e a c h m e l a n i s m measure ( s e p a r a t e l y ) a s a f u n c t i o n of s o l a r r a d i a t i o n and a i r t e m p e r a t u r e , and f o u n d no s i g n i f i c a n t c o r r e l a t i o n (p>.05). S i n c e i t was p o s s i b l e t h a t v a r i a t i o n i n one mel a n i s m measure was c o n f o u n d i n g d e t e c t i o n of an e f f e c t o f t h e o t h e r measure, I c a l c u l a t e d t h e d i s t a n c e of e a c h p o i n t f r o m t h e f l i g h t 82 F i g u r e 17. Male minimum f l i g h t t h r e s h o l d . The c o m b i n a t i o n s o f a i r t e m p e r a t u r e and s o l a r r a d i a t i o n (1982 & 1983) a t w h i c h a t l e a s t one male f l e w ( s o l i d c i r c l e s ) o r no males f l e w (open c i r c l e s ) . 83 g u r e 18. Female minimum f l i g h t t h r e s h o l d . The c o m b i n a t i o n s of a i r t e m p e r a t u r e and s o l a r r a d i a t i o n (1983) a t w h i c h a t l e a s t one f e m a l e f l e w ( s o l i d c i r c l e s ) . The d o t t e d l i n e i n d i c a t e s t h e a p p a r e n t f e m a l e f l i g h t t h r e s h o l d , t h e s o l i d l i n e i s t h e male f l i g h t t h r e s h o l d from F i g u r e 17. 85 Temperature °C 86 F i g u r e 19. P r e d i c t e d minimum f l i g h t t h r e s h o l d s . U s i n g t h e r e g r e s s i o n model from T a b l e X, w i t h b a s a l t r a n s p a r e n c y o m i t t e d and minimum body t e m p e r a t u r e s e t a t 18 C, p r e d i c t e d f l i g h t t h r e s h o l d s were c a l c u l a t e d . L i n e a = t h e a c t u a l male minimum f l i g h t t h r e s h o l d ; l i n e b = t h e p r e d i c t e d t h r e s h o l d f o r an i n d i v i d u a l w i t h the l e a s t b a s a l p a t c h w i d t h and b a s a l p a t c h b l a c k n e s s o b s e r v e d i n t h e Vernon H i l l p o p u l a t i o n ; l i n e c = t h e p r e d i c t e d t h r e s h o l d u s i n g t h e median v a l u e s o f p a t c h w i d t h and b l a c k n e s s ; l i n e d = t h e p r e d i c t e d t h r e s h o l d u s i n g t h e g r e a t e s t v a l u e s of p a t c h w i d t h and b l a c k n e s s . In t h e r e g r e s s i o n model, bt = body t e m p e r a t u r e ; b1 = t h e r e g r e s s i o n c o e f f i c i e n t f o r s o l a r r a d i a t i o n ( s r ) ; b2 = t h e r e g r e s s i o n c o e f f i c i e n t f o r a i r t e m p e r a t u r e ( a t ) ; b3 = t h e r e g r e s s i o n c o e f f i c i e n t f o r b a s a l p a t c h w i d t h ( m l ) ; and b4 = t h e r e g r e s s i o n c o e f f i c i e n t , f o r b a s a l p a t c h b l a c k n e s s (m2). 87 Temperature (°C) 88 F i g u r e 20. E f f e c t of b a s a l p a t c h b l a c k n e s s on f l i g h t t h r e s h o l d . The c o m b i n a t i o n s of s o l a r r a d i a t i o n and a i r t e m p e r a t u r e (1983) f o r w h i c h one o r more d a r k ( s o l i d c i r c l e s ) o r l i g h t (open c i r c l e s ) m a l e s were f l y i n g . Dark and l i g h t g r o u p s were s p l i t a t t h e median p o p u l a t i o n p h e n o t y p e . The minimum male f l i g h t t h r e s h o l d i s i n d i c a t e d . 89 1.4T 1.2+ 1.0 + CD c (0 c \ CO o CO . CO .8-.4+ .2+ % T g-. 4 1 1 r- H »-8 12 16 20 24 Temperature (°C) 28 30 90 F i g u r e 21. E f f e c t of b a s a l p a t c h w i d t h on f l i g h t t h r e s h o l d . The c o m b i n a t i o n s of s o l a r r a d i a t i o n and a i r t e m p e r a t u r e f o r w h i c h one o r more d a r k ( s o l i d c i r c l e s ) o r l i g h t (open c i r l c e s ) m a l e s were f l y i n g . Dark and l i g h t g r o u p s were s p l i t a t t h e median p o p u l a t i o n p h e n o t y p e . The minimum male f l i g h t t h r e s h o l d i s i n d i c a t e d . 91 92 F i g u r e 22. Combined e f f e c t of b a s a l p a t c h w i d t h and b l a c k n e s s on f l i g h t t h r e s h o l d . The c o m b i n a t i o n s of s o l a r r a d i a t i o n and a i r t e m p e r a t u r e (1983) f o r w h i c h one o r more d a r k ( s o l i d c i r c l e s ) or l i g h t (open c i r c l e s ) m ales were f l y i n g . The l i g h t g r o u p c o n s i s t s of t h o s e i n d i v i d u a l s w h i c h were below t h e median f o r b o t h b a s a l p a t c h b l a c k n e s s and w i d t h ; t h e d a r k g r o u p c o n s i s t s of t h o s e i n d i v i d u a l s above t h e median f o r b o t h c h a r a c t e r s . The minimum male f l i g h t t h r e s h o l d i s i n d i c a t e d . S o l a r Radiat ion gm c a l / c m 2 - min 94 t h r e s h o l d , u s i n g t h e i n v e r s e o f t e m p e r a t u r e t o make t h e t h r e s h o l d l i n e a r . I t h e n d i d a m u l t i p l e r e g r e s s i o n of d i s t a n c e from t h e t h r e s h o l d as a f u n c t i o n o f a l l s i x p h e n o t y p i c measures f o r e a c h i n d i v i d u a l . A v e r y weak c o r r e l a t i o n between d i s t a n c e from t h e t h r e s h o l d and b a s a l p a t c h b l a c k n e s s was f o u n d (p=.05; n=400). However by u s i n g d i f f e r e n t t r a n s f o r m a t i o n s of t h e d i s t a n c e from t h e t h r e s h o l d v a l u e s , any one of t h e p h e n o t y p i c m e a sures c o u l d be made s t a t i s t i c a l l y ' s i g n i f i c a n t ' . T h e r e f o r e i t c a n o n l y be c o n c l u d e d t h a t t h e r e i s no b i o l o g i c a l l y s i g n i f i c a n t e f f e c t of p h e n o t y p e on t h e d i s t a n c e f r o m th e f l i g h t t h r e s h o l d a t w h i c h an i n d i v i d u a l i s l i k e l y t o be c a p t u r e d . When t h e d a t a f o r f e m a l e s were a n a l y z e d i n t h e same way as f o r t h e m a l e s , t h e r e was a l s o no d i f f e r e n c e i n a p p a r e n t f l i g h t t h r e s h o l d . T h e r e was a l s o no c o r r e l a t i o n between d i s t a n c e from t h e t h r e s h o l d and p h e n o t y p e f o r f e m a l e s . M e l a n i s m and A c t i v i t y Once above t h e minimum f l i g h t t h r e s h o l d , t h e p r o p o r t i o n of t i m e m a l e s spend i n f l i g h t i s a f u n c t i o n of a i r t e m p e r a t u r e , s o l a r r a d i a t i o n , b a s a l p a t c h w i d t h , and b a s a l p a t c h b l a c k n e s s ( F i g u r e 2 3 a - c ) . The p r o p o r t i o n o f t i m e s p e n t i n f l i g h t i s g r e a t e r f o r h i g h e r v a l u e s of b o t h e n v i r o n m e n t a l v a r i a b l e s and b o t h m e l a n i s m m e a s u r e s . Wind has no e f f e c t on f l i g h t a c t i v i t y o v e r t h e r a n g e u s e d (p>.16 f o r i n t e r a c t i o n s and main e f f e c t ) . F l i g h t a c t i v i t y i n c r e a s e s w i t h i n c r e a s e d s o l a r r a d i a t i o n (p<.002) and i n c r e a s e d a i r t e m p e r a t u r e (p<.0005) under a l l c o n d i t i o n s , but t h e e f f e c t of a change i n e i t h e r s o l a r r a d i a t i o n 9 5 F i g u r e 23. P r o p o r t i o n of t i m e males spend i n f l i g h t . The ANOVA f a c t o r s w h i c h s i g n i f i c a n t l y a f f e c t e d t h e p r o p o r t i o n of t i m e s p e n t i n f l i g h t a r e p l o t t e d . The method o f a n a l y s i s i s d e s c r i b e d i n t h e t e x t . V e r t i c a l b a r s = one s t a n d a r d e r r o r ; t h e numbers above e a c h e r r o r bar = sample s i z e . ( a ) T h e e f f e c t of s o l a r r a d i a t i o n and a i r t e m p e r a t u r e on f l i g h t a c t i v i t y ; ( b ) T h e e f f e c t of b a s a l p a t c h w i d t h and s o l a r r a d i a t i o n on f l i g h t a c t i v i t y ; ( c ) T h e e f f e c t of b a s a l p a t c h b l a c k n e s s and a i r t e m p e r a t u r e on f l i g h t a c t i v i t y . 100-t 80H ON? x 60 O) •- 40 E 20H (a)Temp.xSolar Interaction p=.05 9 I lo hi lo hi LOW HIGH Solar Radiation (b)Basal Patch WidthxSolar Interact ion p<.04 1 It dk It dk LOW HIGH Solar Radiation COBasal Patch Blackness xTemp. Interaction p<.004 9 7 or a i r t e m p e r a t u r e i s g r e a t e s t when t h e o t h e r f a c t o r i s h i g h e r . I n c r e a s e d b a s a l p a t c h w i d t h i n c r e a s e s f l i g h t a c t i v i t y a t low s o l a r r a d i a t i o n s o n l y , and i n c r e a s e d b a s a l p a t c h b l a c k n e s s i n c r e a s e s f l i g h t a c t i v i t y a t low t e m p e r a t u r e s o n l y . The p r o p o r t i o n o f t i m e m a l e s s p e n d f e e d i n g d e c r e a s e s w i t h i n c r e a s e d s o l a r r a d i a t i o n o r i n c r e a s e d t e m p e r a t u r e ( F i g u r e 24a-b ) . T h e r e i s no e f f e c t o f b a s a l p a t c h w i d t h , b a s a l p a t c h b l a c k n e s s , o r wind ( a l l p>.35 f o r i n t e r a c t i o n s and main e f f e c t s ) on p r o p o r t i o n of t i m e s p e n t f e e d i n g . The p r o p o r t i o n of t i m e males spend b a s k i n g but n o t f e e d i n g may be d e p e n d e n t on a l l f i v e v a r i a b l e s . Time s p e n t b a s k i n g d e c r e a s e s w i t h i n c r e a s e s i n b o t h s o l a r r a d i a t i o n (p<.0005) and a i r t e m p e r a t u r e (p<.0005) r e g a r d l e s s o f t h e v a l u e s o f t h e o t h e r v a r i a b l e s . A t low s o l a r r a d i a t i o n i n c r e a s e d b a s a l p a t c h w i d t h d e c r e a s e s t h e t i m e s p e n t b a s k i n g , but a t h i g h s o l a r r a d i a t i o n i t has no e f f e c t ( F i g u r e 2 5 a ) . I n c r e a s e d b a s a l p a t c h b l a c k n e s s d e c r e a s e s t h e amount o f b a s k i n g a t low a i r t e m p e r a t u r e s , but not a t h i g h t e m p e r a t u r e s ( F i g u r e 2 5 c ) . V a r i a n c e s a r e n o t homogeneous f o r t h e b a s a l p a t c h b l a c k n e s s by s o l a r r a d i a t i o n i n t e r a c t i o n ( F i g u r e 25b) and t h e b l a c k n e s s by wind i n t e r a c t i o n ( F i g u r e 2 5 d ) , t h e r e f o r e a l t h o u g h t h e r e s u l t s a r e p r e s e n t e d t h e s i g n i f i c a n c e l e v e l s of t h e d i f f e r e n c e s i n t h e means a r e not c o n s i d e r e d t o be r e l i a b l e . A p p a r e n t l y c h a n g e s i n e i t h e r p a t c h w i d t h o r b l a c k n e s s have a g r e a t e r e f f e c t on t i m e s p e n t b a s k i n g when t h e o t h e r f a c t o r i s low ( F i g u r e 2 5 e ) , a l t h o u g h t h e e f f e c t i s m i n o r . I a l s o t e s t e d f o r t h e e f f e c t s of t h e o t h e r p h e n o t y p i c u r e 24. P r o p o r t i o n of t i m e s p e n t f e e d i n g . The ANOVA f a c t o r s w h i c h s i g n i f i c a n t l y a f f e c t e d t h e p r o p o r t i o n of t i m e s p e n t f e e d i n g a r e p l o t t e d . The method o f a n a l y s i s i s d e s c r i b e d i n t h e t e x t . V e r t i c a l b a r s = one s t a n d a r d e r r o r ; t h e numbers above e a c h e r r o r bar = sample s i z e . (a) The e f f e c t o f s o l a r r a d i a t i o n on f e e d i n g a c t i v i t y . (b) The e f f e c t of a i r t e m p e r a t u r e on f e e d i n g a c t i v i t y . O n l y o b s e r v a t i o n s i n w h i c h a t l e a s t some f e e d i n g o c c u r r e d were i n c l u d e d i n t h e a n a l y s i s . (a)Temperature p<.02 100-1 C 80 C 1 60-o LL © E 40 20 10 LOW HIGH Air Temperature (b)Solar Radiation p < .005 10 I 13 LOW HIGH Solar Radiation 1 00 F i g u r e 25. P r o p o r t i o n o f t i m e s p e n t b a s k i n g . The ANOVA f a c t o r s w h i c h s i g n i f i c a n t l y a f f e c t e d t h e p r o p o r t i o n of t i m e s p e n t b a s k i n g but n o t f e e d i n g a r e p l o t t e d . The method of a n a l y s i s i s d e s c r i b e d i n t h e t e x t . V e r t i c a l b a r s = one s t a n d a r d e r r o r ; t h e numbers above e a c h e r r o r bar = sample s i z e . O n l y o b s e r v a t i o n s i n w h i c h o v e r 5% o f t h e t o t a l t i m e was s p e n t i n f l i g h t were i n c l u d e d i n t h e a n a l y s i s . ( a ) T h e e f f e c t of b a s a l p a t c h w i d t h and s o l a r r a d i a t i o n , ( b ) T h e e f f e c t of b a s a l p a t c h b l a c k n e s s and s o l a r r a d i a t i o n , ( c ) T h e e f f e c t of b a s a l p a t c h b l a c k n e s s and a i r t e m p e r a t u r e , ( d ) T h e e f f e c t of b a s a l p a t c h b l a c k n e s s and wind, ( e ) T h e e f f e c t of b a s a l p a t c h w i d t h and b a s a l p a t c h b l a c k n e s s . (a)Basal Patch WidthxSolar Interaction P<005 100-t ^ 80 cn c w CO C D 60H <D E 40-20 It dk It dk LOW HIGH Solar Radiation (b)Basal Patch BlacknessxSolar Interaction P<.04 Homogeneity P>.01 9 T 8 (c)Basal Patch BlacknessxTemp. Interaction p<.0005 8 It dk It dk LOW HIGH Solar Radiation It dk It dk LOW HIGH Air Temperature (d)Basal Patch BlacknessxWind Interaction P<.03 100 Homogeneity n p>.02 80-(e)Basal Patch Widthx Blackness Interaction p<.03 10 1_ It dk It dk LIGHT DARK B L A C K N E S S 103 c h a r a c t e r s on f l i g h t a c t i v i t y , t e s t i n g f i r s t b a s a l t r a n s p a r e n c y and f r o n t wing l e n g t h , and t h e n d i s t a l t r a n s p a r e n c y and b l a c k n e s s . None of t h e s e c h a r a c t e r s s i g n i f i c a n t l y a f f e c t e d t i m e s p e n t i n f l i g h t (p>.05). M e l a n i s m and S u r v i v a l The f r e q u e n c y d i s t r i b u t i o n of male r e s i d e n c e t i m e ( F i g u r e 26) a p p r o x i m a t e s a n o r m a l d i s t r i b u t i o n s l i g h t l y skewed tow a r d s l o w e r r e s i d e n c e t i m e , w i t h t h e e x c e p t i o n of a l a r g e c l a s s of i n d i v i d u a l s c a p t u r e d once and n e v e r a g a i n . T h e r e f o r e d i s a p p e a r a n c e i s p r i m a r i l y age d e p e n d e n t . Of t h e 45 one t i m e o n l y c a p t u r e s , 14 were i n t h e f i n a l s ample. The 14 o n e - t i m e -o n l y c a p t u r e s i n t h e f i n a l sample were l e f t i n t h e a n a l y s e s b e c a u s e t h e i r f a i l u r e t o be r e c a p t u r e d was t h e r e s u l t of a b i o l o g i c a l e v e n t (bad w e a t h e r ) r a t h e r t h a n l a c k of s a m p l i n g e f f o r t . When the number of d a y s i n r e s i d e n c e i s p l o t t e d as a c u m u l a t i v e f r e q u e n c y d i s t r i b u t i o n ( F i g u r e 2 7 ) , t h e d i s t r i b u t i o n i s l i n e a r o v e r most o f t h e r a n g e , i n d i c a t i n g t h a t age-i n d e p e n d e n t l o s s e s from t h e p o p u l a t i o n a r e n e g l i g i b l e , e x c e p t f o r t h e c a p t u r e d o n e - t i m e - o n l y g r o u p . T h e r e i s no s i g n i f i c a n t d i f f e r e n c e i n r e s i d e n c e t i m e ( o n e - t a i l e d K o l m o g o r o v - S m i r n o v Two-sample T e s t , p>.05) f o r m a l e s w h i c h a r e l i g h t or d a r k f o r e i t h e r b a s a l p a t c h b l a c k n e s s ( F i g u r e 28) o r b a s a l p a t c h w i d t h ( F i g u r e 2 9 ) . Even when o n l y m a l e s w h i c h a r e l i g h t or d a r k f o r b o t h p a t c h w i d t h and b l a c k n e s s a r e c o n s i d e r e d t h e r e i s no d i f f e r e n c e i n r e s i d e n c e t i m e ( F i g u r e 3 0 ) . I f t h e 14 o n e - t i m e - o n l y c a p t u r e s a r e e l i m i n a t e d from t h e a n a l y s e s , t h e r e i s s t i l l no p h e n o t y p e 1 04 F i g u r e 26. F r e q u e n c y d i s t r i b u t i o n of r e s i d e n c e t i m e . The number of days from f i r s t t o l a s t c a p t u r e a t V e r n o n H i l l i n 1983 f o r a l l males i n A r e a 2. 105 o m co I-s o m o o CO i r o CM CM 0) o e <D C O (1) cc .in C O S|Bnp!A|pu| 106 d e p e n d e n t s u r v i v a l f o r t h e s e c h a r a c t e r s . When I l o o k e d a t t h e o t h e r c h a r a c t e r s , t h e r e was a l s o no e f f e c t of b a s a l t r a n s p a r e n c y , d i s t a l t r a n s p a r e n c y , o r d i s t a l b l a c k n e s s on r e s i d e n c e t i m e , w i t h t h e c u m u l a t i v e r e s i d e n c e c u r v e s e s s e n t i a l l y t h e same a s f o r b a s a l p a t c h b l a c k n e s s . However s m a l l e r i n d i v i d u a l s d i s a p p e a r f a s t e r t h a n l a r g e i n d i v i d u a l s ( F i g u r e 31; t w o - t a i l e d K o l m o g o r o v - S m i r n o v T e s t , p<.05). S a m p l i n g of t h e f e m a l e s was not i n t e n s i v e enough f o r m e a n i n g f u l a n a l y s i s of s u r v i v a l r a t e s . When t h e 14 o n e - t i m e -o n l y c a p t u r e s i n t h e l a s t sample were e l i m i n a t e d from t h e a n a l y s e s t h e c o n c l u s i o n s were u n c h a n g e d . P o p u l a t i o n P a r a m e t e r s The m a r k / r e c a p t u r e d a t a were u s e d t o c a l c u l a t e s u r v i v a l r a t e s , p o p u l a t i o n s i z e s , and a d d i t i o n s t o t h e male p o p u l a t i o n of A r e a 2 f o r e a c h sample p e r i o d u s i n g t h e method o f J o l l y (1965) as m o d i f i e d by Seber ( 1 9 7 3 ) . A h i g h p r o p o r t i o n of t h e p o p u l a t i o n was marked y i e l d i n g p r e c i s e s u r v i v a l e s t i m a t e s ( T a b l e X I ) . S e v e n t y - s i x p e r c e n t of t h e m a l e s were r e c a p t u r e d a t l e a s t once i n A r e a 2, 82% i f t h e o n e - t i m e c a p t u r e s i n t h e f i n a l sample a r e e x c l u d e d , and 88% i f males w h i c h were c a p t u r e d o u t s i d e A r e a 2 b e f o r e o r a f t e r c a p t u r e i n A r e a 2 a r e i n c l u d e d . T h e r e were i n s u f f i c i e n t d a t a t o c a l c u l a t e f e m a l e p o p u l a t i o n p a r a m e t e r s mean i n g f u l l y . 107 F i g u r e 27. C u m u l a t i v e f r e q u e n c y d i s t r i b u t i o n o f r e s i d e n c e t i m e . The number of d a ys from f i r s t t o l a s t c a p t u r e a t V e r n o n H i l l i n 1983 f o r a l l m a l e s i n A r e a 2. Note t h e l i n e a r s c a l e f o r the o r d i n a t e . 108 20CH 160 i2 120 CU 3 73 • MM > a £ 80H 40 1 1 1 1 1 10 20 30 40 Days in R e s i d e n c e 109 F i g u r e 28. B a s a l p a t c h b l a c k n e s s and r e s i d e n c e t i m e . C u m u l a t i v e f r e q u e n c y d i s t r i b u t i o n f o r l i g h t (open c i r c l e s ) and d a r k ( s o l i d c i r c l e s ) m a l e s . L i g h t and d a r k c l a s s e s were s p l i t a t t h e m edian. 100 1 80 60 • 0 • 00 40 o 20 T I I : I I I 10 20 30 Days in R e s i d e n c e 111 F i g u r e 29. B a s a l p a t c h w i d t h and r e s i d e n c e t i m e . C u m u l a t i v e f r e q u e n c y d i s t r i b u t i o n f o r l i g h t (open c i r c l e s ) and d a r k ( d a r k c i r c l e s ) m a l e s . L i g h t and d a r k c l a s s e s were s p l i t a t t h e median. 1 0% 80 60 c \ o 40-I 20 H O 10 20 30 40 Days in R e s i d e n c e 1 1 3 F i g u r e 30. B a s a l p a t c h w i d t h and b l a c k n e s s and r e s i d e n c e t i m e . C u m u l a t i v e f r e q u e n c y d i s t r i b u t i o n f o r l i g h t (open c i r c l e s ) and d a r k (dark c i r c l e s ) m a l e s . L i g h t i n d i v i d u a l s a r e l i g h t f o r b o t h b a s a l p a t c h w i d t h and b l a c k n e s s ; d a r k i n d i v i d u a l s a r e d a r k f o r b o t h c h a r a c t e r s . L i g h t and d a r k c l a s s e s were s p l i t a t t h e m e dian. 114 50 ^  40 00. o j2 30 CU 3 £ 20H o. °o o G O G O O 10 H o •CD 00. GO 000 10 20 30 Days in R e s i d e n c e 40 1 1 5 F i g u r e 31. F r o n t wing l e n g t h and r e s i d e n c e t i m e . C u m u l a t i v e f r e q u e n c y d i s t r i b u t i o n f o r s m a l l (open c i r c l e s ) and l a r g e ( s o l i d c i r c l e s ) m a l e s . 100 80 (f) CD 3 C C) _ ° ° o o • o V , • t o • •• I I I I I 10 20 30 Days in R e s i d e n c e 1 17 T a b l e X I . M a l e P o p u l a t i o n P a r a m e t e r s f o r V e r n o n H i l l , 1983. D a t e Marked P o p u l a t i o n P r o b . of S u r v i v a l Number Number (% of S i z e Between J o i n i n g Caught Sample) Samples Days ( s . e . ) ( s . e . ) ( s . e . ) June 3 5 1 6 38 9 29 1 04 1 1 52 1 1 1 1 4 51 131 18 77 1 28 19 77 1 30 21 76 142 22 86 1 19 24 83 1 1 7 25 82 121 27 85 100 28 81 131 J u l y 3 82 38 18 64 .77 (.16) (8) .94 ( .07) (22) 1 .03 ( .08) (14) .80 ( .07) (11) .92 ( .09) (20) .99 ( .09) (11) .98 ( . 10) (19) .89 ( .09) (10) .89 ( . 10) (18) .98 (.11) (12) .79 ( .07) (8) 1.11 ( .19) (24) • .26 ( .08) (10) .88 8 .98 68 (22) 31 1.01 5 (24) 38 .93 42 (13) 52 .98 8 (18) 83 .99 4 (17) 1 7 .99 1 8 (15) 57 .89 8 (13) 29 .94 1 3 (12) 64 .98 6 (13) 18 .89 6 (6) 55 1.11 20 (7) 74 .76 4 (4) 75 1 7 22 D i s p e r s a l At l e a s t 6 males and 1 f e m a l e moved from A r e a s 1-3 t o A r e a s 4-7 ( F i g u r e 3 2 ) . The two g r e a t e s t d i s t a n c e s moved were 1.5 and 1.7 km i n a s t r a i g h t l i n e . S i n c e i t i s l i k e l y t h a t t h e f l i g h t r o u t e s from A r e a 2 t o A r e a 7 f o l l o w e d t h e f o r e s t o p e n i n g s c o n n e c t i n g A r e a s 4-7, t h e a c t u a l d i s t a n c e moved was p r o b a b l y 2-3 km. Of t h e s i x males w h i c h moved from A r e a 2, f o u r were i n t h e ' c a p t u r e d o n e - t i m e - o n l y ' g r o u p f o r A r e a 2. The o t h e r two r e m a i n e d a t l e a s t 16 and 21 days i n A r e a 2. F o r t h e m a l e s which were c a p t u r e d on June 28 and J u l y 18, t h e r a t i o of ' c a p t u r e d once p r e v i o u s l y i n A r e a 2' t o ' c a p t u r e d more t h a n once p r e v i o u s l y i n A r e a 2' f o r A r e a s 4-7 (4/2) i s s i g n i f i c a n t l y d i f f e r e n t from t h a t f o r A r e a 2 (8/65) ( F i s h e r ' s E x a c t T e s t , p=.004). The r a t i o s were n o t s i g n i f i c a n t l y d i f f e r e n t (p>.1) when c o m p a r i n g t h e d a t a from A r e a s 1 and 3 t o A r e a 2 f o r June 1 18 F i g u r e 32. D i s p e r s a l . T h i n l i n e s i n d i c a t e s i n g l e movements ( s o l i d = male; d o t t e d = f e m a l e ) ; t h i c k l i n e s i n d i c a t e a t l e a s t f o u r movements o f m a l e s , w i t h f e m a l e s a l s o moving between a r e a s 1, 2, and 3. The numbers r e f e r t o A r e a s 1-7. S t i p p l e d a r e a s i n d i c a t e f o r e s t , w h i t e a r e a s i n d i c a t e g r a s s l a n d . 120 22, June 28, and J u l y 18. T h i r t y - t w o m a l e s moved between A r e a s 1, 2, and 3, w i t h e i t h e r 1 o r 2 s u c h movements r e c o r d e d f o r e a c h i n d i v i d u a l . The p h e n o t y p e was s c o r e d f o r a l l but f i v e of t h e 32. T h e r e were no s i g n i f i c a n t d i f f e r e n c e s i n p h e n o t y p e when c o m p a r i n g t h e 'never moved from A r e a 2' g r o u p o f m a l e s w i t h t h e 'moved a t l e a s t once between a r e a s ' g r o u p (p>.05). However, t h e males which moved t w i c e between a r e a s had s i g n i f i c a n t l y g r e a t e r b a s a l p a t c h b l a c k n e s s (mean=95% b l a c k s c a l e s , s.e.=1%, n=8) t h a n t h o s e w h i c h moved o n l y once (mean=89%, s.e.=approx.2%, d i s t r i b u t i o n s l i g h t l y l e p t o k u r t i c , n=l9) ( o n e - t a i l e d Mann-Whitney U - T e s t , p<.025). T h e r e f o r e m a l e s w i t h g r e a t e r b a s a l p a t c h b l a c k n e s s have a g r e a t e r t e n d e n c y t o d i s p e r s e , a t l e a s t f o r s h o r t d i s t a n c e s . T h e r e was no e f f e c t on movement between a r e a s f o r any of t h e o t h e r m e a s u r e s of m e l a n i s m o r f o r f r o n t wing l e n g t h (p>.05). T h e r e was no s i g n i f i c a n t d i f f e r e n c e between t h e p h e n o t y p i c c h a r a c t e r s of t h e males w h i c h moved from A r e a 2 t o A r e a s 4-7 and t h o s e of t h e A r e a 2 sample on June 28 and J u l y 18 ( o n e - t a i l e d Mann-Whitney U - T e s t s , p>.05). T h e r e f o r e t h e r e i s no e v i d e n c e f o r p h e n o t y p e a f f e c t i n g t h e t e n d e n c y t o d i s p e r s e ' l o n g ' d i s t a n c e s . F o r t h e i n d i v i d u a l s c a p t u r e d i n A r e a s 1-3 on .June 22, June 28, and J u l y 18 w h i c h had been c a p t u r e d a t l e a s t once b e f o r e i n A r e a 2, t h e r a t i o of 'moved from A r e a 2 t o A r e a s 1 or 3' t o ' r e m a i n e d i n A r e a 2' f o r f e m a l e s (11/15) was s i g n i f i c a n t l y g r e a t e r t h a n t h e r a t i o f o r m a l e s (25/118) ( F i s h e r ' s E x a c t T e s t , p< i 0 1 ) . T h e r e f o r e f e m a l e s have a g r e a t e r t e n d e n c y t o d i s p e r s e 121 t h a n m a l e s . T h e r e were t o o few f e m a l e s w h i c h moved t w i c e between A r e a s 1-3, o r wh i c h moved t o A r e a s 4-7, t o make most of t h e c o m p a r i s o n s made f o r m a l e s . When t h e 'never moved from A r e a 2' g r o u p o f f e m a l e s was compared w i t h t h e 'moved a t l e a s t once between a r e a s ' g r o u p t h e o n l y s i g n i f i c a n t d i f f e r e n c e i n p h e n o t y p e was f o r w i n g l e n g t h ( t w o - t a i l e d Mann-Whitney U - T e s t , p<.02). The f e m a l e s w h i c h moved (mean s i z e = 36.1mm, s.e.=.38mm, n=12) were l a r g e r t h a n t h o s e w h i c h d i d not (mean s i z e = 35.0mm, s . e . = . 2 l , n=!5). 122 DISCUSSION B a s a l M e l a n i s m and Body T e m p e r a t u r e Body t e m p e r a t u r e i s a f u n c t i o n of b o t h b a s a l p a t c h w i d t h and b l a c k n e s s i n P a r n a s s i u s phoebus, as b a s a l b l a c k n e s s i s i n C o l i a s . T h e r e f o r e i t was r e a s o n a b l e t o p r e d i c t t h a t t h e b e h a v i o r of b o t h s p e c i e s would be a f f e c t e d s i m i l a r i l y by m e l a n i s m . B a s a l t r a n s p a r e n c y has a s m a l l n e g a t i v e c o r r e l a t i o n w i t h body t e m p e r a t u r e , b e c a u s e i f l i g h t does n o t e n c o u n t e r a s c a l e i t j u s t p a s s e s t h r o u g h t h e t r a n s p a r e n t wing membrane and t h e r e f o r e can not h e a t t h e body. The e f f e c t of b a s a l t r a n s p a r e n c y on body t e m p e r a t u r e i s s u f f i c i e n t l y weak t h a t i t i s u n l i k e l y t o have a m e a s u r a b l e e f f e c t on a c t i v i t y . The a b s o r p t i v i t i e s and e m i s s i v i t i e s of t h e b a s a l wing a r e a a r e n e a r l y i d e n t i c a l f o r i n f r a - r e d w a v e l e n g t h s i n a l l b u t t e r f l i e s , r e g a r d l e s s of v i s i b l e b a s a l wing c o l o r ( D o u g l a s , 1978). T h e r e f o r e t h e e f f e c t on body t e m p e r a t u r e of b a s a l wing c o l o r i s p r o b a b l y due t o v a r i a t i o n i n a b s o r p t i v i t y of l i g h t w a v e l e n g t h s i n t h e v i s i b l e r e g i o n of t h e s p e c t r u m , r a t h e r t h a n i n t h e i n f r a -r e d r e g i o n . The a d d i t i v e r e g r e s s i o n model u s e d f o r t h e e f f e c t of m e l a n i s m on body t e m p e r a t u r e ( T a b l e X) gave r e g r e s s i o n c o e f f i c i e n t s f o r t h e m e l a n i s m v a r i a b l e s -which a r e a v e r a g e e f f e c t s of t h o s e v a r i a b l e s o v e r t h e r a n g e o f s o l a r r a d i a t i o n t e s t e d . A more c o r r e c t model would have t h e m e l a n i s m t e r m s i n t h e e q u a t i o n m u l t i p l i e d by s o l a r r a d i a t i o n and t h e m s e l v e s , s o m e t h i n g s i m i l a r t o ' t r a n s p a r e n c y * w i d t h * s o l a r ' and 123 ' b l a c k n e s s * w i d t h * s o l a r 1 a c c o r d i n g t o K i n g s o l v e r ' s b i o m e c h a n i c a l model o f t h e r m o r e g u l a t i o n f o r l a t e r a l b a s k i n g C o l i a s ( K i n g s o l v e r , 1983a; K i n g s o l v e r , 1983b; K i n g s o l v e r and M o f f a t , 1982). I t e s t e d a number of s u c h m o d e l s , most of w h i c h f i t t h e d a t a as w e l l o r b e t t e r t h a n t h e a d d i t i v e model, but w i t h o u t a s p e c i f i c b i o m e c h a n i c a l model of t h e r m o r e g u l a t i o n f o r d o r s a l b a s k i n g b u t t e r f l i e s I c o u l d n o t j u s t i f y c h o o s i n g any one of them i n s t e a d of t h e a d d i t i v e m odel. A l l t h e m u l t i p l i c a t i v e models of t h e e f f e c t of b a s a l m e l a n i s m on body t e m p e r a t u r e r e s u l t i n a c u r v e d p r e d i c t e d f l i g h t t h r e s h o l d , w h i c h f i t s t h e o b s e r v e d t h r e s h o l d shape b e t t e r t h a n t h e l i n e a r m o d e l . However, b o t h t h e l i n e a r and m u l t i p l i c a t i v e m o d e l s p r e d i c t a minimum f l i g h t t h r e s h o l d w e l l below t h a t o b s e r v e d ( F i g u r e 19). D r i e d b u t t e r f l i e s h e a t more r a p i d l y and a c h i e v e e i t h e r a p p r o x i m a t e l y t h e same e q u i l i b r i u m t e m p e r a t u r e s as l i v e b u t t e r f l i e s ( W a s s e r t h a l , 1975) o r s u b s t a n t i a l l y l o w e r e q u i l i b r i u m t e m p e r a t u r e s ( D o u g l a s , 1978) under c o n s t a n t s o l a r r a d i a t o n . T h e r e f o r e t h e minimum f l i g h t t h r e s h o l d f o r l i v e b u t t e r f l i e s s h o u l d be e i t h e r t h e same as p r e d i c t e d or l o w e r t h a n t h a t p r e d i c t e d , w i t h t h e d i f f e r e n c e between f l i g h t t h r e s h o l d s f o r l i g h t and d a r k i n d i v i d u a l s a s p r e d i c t e d . S i n c e t h e o b s e r v e d f l i g h t t h r e s h o l d i s not p h e n o t y p e d e p e n d e n t and i s c o n s i d e r a b l y above t h e p r e d i c t e d t h r e s h o l d s , e i t h e r : (1) t h e f l i g h t t h r e s h o l d i s not d e t e r m i n e d by body t e m p e r a t u r e , o r (2) t h e b a s k i n g c o n d i t i o n s f o r t h e d ead, d r i e d b u t t e r f l i e s were not a r e a s o n a b l e a p p r o x i m a t i o n of t h o s e e x p e r i e n c e d by l i v e 124 b u t t e r f l i e s . Minimum Body T e m p e r a t u r e f o r F l i q h t A minimum body t e m p e r a t u r e of a b o u t 18°C i s r e q u i r e d f o r v o l u n t a r y f l i g h t i n P. phoebus m a l e s , w h i c h i s s u b s t a n t i a l l y h i g h e r t h a n t h e 10°C minimum f o r t h e Monarch (Danaus p l e x i p p u s L.) ( C h a p l i n and W e l l s , 1982), but much lo w e r t h a n t h e 28-30°C f o r C o l i a s m a l e s and 35°C f o r C o l i a s f e m a l e s (Watt, 1968). Low body t e m p e r a t u r e s r e s u l t i n low wing b e a t f r e q u e n c i e s i n b u t t e r f l i e s (Kammer, 1970), p r o b a b l y due t o d e c r e a s e d s p e e d of m u s c l e c o n t r a c t i o n ( N e v i l l e and W eis-Fogh, 1963). T h e r e f o r e slow f l y i n g s p e c i e s s u c h as Danaus and P a r n a s s i u s w i l l be a b l e t o f l y w i t h a l o w e r minimum body t e m p e r a t u r e t h a n t h e more r a p i d f l y i n g C o l i a s . Body T e m p e r a t u r e and t h e Minimum F l i g h t T h r e s h o l d At below 20°C t h e f l i g h t t h r e s h o l d i s d e f i n e d by b o t h a i r t e m p e r a t u r e and s o l a r r a d i a t i o n ( F i g u r e 1 7 ) . T h i s p o r t i o n of t h e f l i g h t t h r e s h o l d i s s i m i l a r t o t h a t f o u n d f o r A r g y n n i s  p a p h i a ( N y m p h a l i d a e ) ( V i e l m e t t e r , 1958). Heodes v i r g a u r a e ( L y c a e n i d a e ) a l s o has a s i m i l a r f l i g h t t h r e s h o l d f o r t h e mid-day p e r i o d , but f l i g h t i n t h e e a r l y m o r n i n g and l a t e a f t e r n o o n i s d e t e r m i n e d by f a c t o r s o t h e r t h a n a i r t e m p e r a t u r e and s o l a r r a d i a t i o n (Douwes, 1976). Above 20°C t h e f l i g h t t h r e s h o l d i n P. phoebus i s d e f i n e d o n l y by s o l a r r a d i a t i o n , a r e s p o n s e w h i c h was not d e t e c t e d i n e i t h e r A. p a p h i a o r H. v i r g a u r a e ( V i e l m e t t e r , 1 25 1958; Douwes, 1976). Below t h e t r a n s i t i o n p o i n t , t h e f l i g h t t h r e s h o l d i s a p p a r e n t l y d e t e r m i n e d by body t e m p e r a t u r e , and above i t t h e t h r e s h o l d i s i n d e p e n d e n t of body t e m p e r a t u r e . I f a i r t e m p e r a t u r e were t h e o n l y f a c t o r a f f e c t i n g body t e m p e r a t u r e , t h e n t h e t r a n s i t i o n p o i n t i n t h e f l i g h t t h r e s h o l d would be t h e p o i n t a t w h i c h a i r t e m p e r a t u r e e q u a l s t h e minimum body t e m p e r a t u r e r e q u i r e d f o r f l i g h t . I f t h e r e i s an o v e r a l l h e a t g a i n from s o l a r r a d i a t i o n r e l a t i v e t o h e a t l o s s e s t o t h e s u b s t r a t e and t h r o u g h r e s p i r a t o r y e v a p o r a t i o n , t h e t r a n s i t i o n p o i n t s h o u l d be a t a s l i g h t l y l ower a i r t e m p e r a t u r e t h a n t h e minimum body t e m p e r a t u r e r e q u i r e d f o r f l i g h t . I f h e a t l o s s e s a r e g r e a t e r t h a n h e a t g a i n from s o l a r r a d i a t i o n , t h e n t h e t r a n s i t i o n p o i n t s h o u l d be h i g h e r . S i n c e t h e a i r t e m p e r a t u r e a t th e t r a n s i t i o n p o i n t i s a p p r o x i m a t e l y 2°C above minimum body t e m p e r a t u r e f o r f l i g h t , h e a t i n p u t from s o l a r r a d i a t i o n of 0.2 gm.cal/cm 2/min c o n t r i b u t e s a b o u t 2°C l e s s t h a n i s l o s t . T h i s c l o s e c o r r e l a t i o n between t h e a i r t e m p e r a t u r e a t t h e t r a n s i t i o n p o i n t and t h e minimum body t e m p e r a t u r e r e q u i r e d f o r f l i g h t , s t r o n g l y s u g g e s t s t h a t t h e c u r v e d p o r t i o n o f t h e f l i g h t t h r e s h o l d i s d e t e r m i n e d by body t e m p e r a t u r e . B a s k i n g C o n d i t i o n s and t h e F l i g h t T h r e s h o l d The minimum f l i g h t t h r e s h o l d f o r P a r n a s s i u s phoebus i s a p p a r e n t l y d e t e r m i n e d by body t e m p e r a t u r e o v e r most of t h e ra n g e of e n v i r o n m e n t a l c o n d i t i o n s . S i n c e b a s a l wing m e l a n i s m has no e f f e c t on t h e f l i g h t t h r e s h o l d , i t does n o t a f f e c t body t e m p e r a t u r e w h i l e b a s k i n g under f i e l d c o n d i t i o n s . The major 126 d i f f e r e n c e s between t h e f i e l d and l a b o r a t o r y c o n d i t i o n s were: (a) i n t h e f i e l d t h e b u t t e r f l i e s were a l i v e r a t h e r t h a n dead, w h i c h m i g h t a f f e c t mean body t e m p e r a t u r e but s h o u l d n ot change the r e l a t i v e e f f e c t of d i f f e r e n t wing c o l o r s , and (b) i n t h e f i e l d b a s k i n g b u t t e r f l i e s r e s t on a s u b s t r a t e i n s t e a d of b e i n g s u s p e n d e d i n m i d - a i r , w h i c h i s p r o b a b l y t h e f a c t o r w h i c h r e s u l t s i n wing c o l o r h a v i n g no e f f e c t on t h e minimum f l i g h t t h r e s h o l d . The l a t t e r p o s s i b i l i t y f i t s t h e o b s e r v a t i o n t h a t t h e wings of b u t t e r f l i e s d o r s a l b a s k i n g on a s u b s t r a t e have l i t t l e or no e f f e c t on body t e m p e r a t u r e ( H e i n r i c h , 1972; Kammer and B r a c c h i , 1973). I f t h i s i s t r u e , t h e n t h e b a s k i n g c o n d i t i o n s of t h e dead, d r i e d b u t t e r f l i e s ( s u s p e n d e d ) were not a r e a s o n a b l e a p p r o x i m a t i o n o f t h e e f f e c t s o f s o l a r r a d i a t i o n , a i r t e m p e r a t u r e , and wing c o l o r on t h e body t e m p e r a t u r e of a l i v e b u t t e r f l y b a s k i n g on a s u b s t r a t e . B a s k i n g and Body T e m p e r a t u r e The p r e s e n c e of wings on a d o r s a l b a s k i n g b u t t e r f l y r e s t i n g on a s u b s t r a t e a p p a r e n t l y d o e s n o t r e s u l t i n t h e i n c r e a s e d t r a n s f e r o f e n e r g y from s o l a r r a d i a t i o n t o t h e body compared t o t h a t s p e c i m e n w i t h t h e wings removed, p o s s i b l y due t o h e a t t r a n s f e r t o t h e s u b s t r a t e ( D o u g l a s , 1978; H e i n r i c h , 1972; Kammer and B r a c c h i , 1972). The s i g n i f i c a n c e of t h i s o b s e r v a t i o n has a p p a r e n t l y n o t been u n d e r s t o o d , s i n c e r e c e n t p u b l i s h e d t e s t s f o r the e f f e c t of w i n g s , o r wing c o l o r , on body t e m p e r a t u r e of d o r s a l b a s k i n g b u t t e r f l i e s have been done on s p e c i m e n s s u s p e n d e d 127 i n m i d - a i r , as I d i d ( t o ' e l i m i n a t e t h e e f f e c t s of s u b s t r a t e ' ) , and t h e r e s u l t s a p p l i e d t o b u t t e r f l i e s d o r s a l b a s k i n g on a s u b s t r a t e ( D o u g l a s , 1978; W a s s e r t h a l , 1975). A l t h o u g h i t has been known f o r o v e r a d e c a d e t h a t t h e wings o f a d o r s a l b a s k i n g b u t t e r f l y may have l i t t l e or no e f f e c t on body t e m p e r a t u r e w h i l e on t h e g r o u n d , but do have an e f f e c t i n t h e a i r , a c o n n e c t i o n has n ot been made between t h e two p i e c e s of i n f o r m a t i o n . The l o g i c a l c o n c l u s i o n i s t h a t wings o n l y a f f e c t t h e body t e m p e r a t u r e of a d o r s a l b a s k i n g b u t t e r f l y when i t i s i n m i d - a i r , and n o t when i t i s r e s t i n g on a s u b s t r a t e . I f t h i s i s t r u e t h e n body c o l o r may a f f e c t body t e m p e r a t u r e w h i l e b a s k i n g , as do t h e e l y t r a of b e e t l e s (Benham. e t a l . , 1974), b ut wing c o l o r w i l l n o t . In a d o r s a l b a s k i n g b u t t e r f l y t h e a i r warmed under t h e wings has a v e r y l a r g e ' s u r f a c e a r e a a g a i n s t t h e g r o u n d ' t o 'volume' r a t i o , as w e l l a s a l a r g e ' s u r f a c e a r e a a g a i n s t t h e g r o u n d ' t o ' s u r f a c e a r e a a g a i n s t t h e b u t t e r f l y body' r a t i o . T h i s would r e s u l t i n t h e m a j o r i t y o f t h e h e a t t r a p p e d by t h e wings b e i n g t r a n s f e r r e d t o t h e g r o u n d b e f o r e i t r e a c h e s t h e body. I t would n o t make any d i f f e r e n c e i f t h e s u b s t r a t e t e m p e r a t u r e were above o r below t h e a i r t e m p e r a t u r e a t t h e moment a b u t t e r f l y l a n d s . A f t e r a b u t t e r f l y commenced b a s k i n g , t h e t h i n l a y e r of a i r between t h e s u b s t r a t e and t h e wing would r a p i d l y change t o t h e s u b s t r a t e t e m p e r a t u r e due t o t h e l a r g e s u r f a c e a r e a t o volume r a t i o . Any h e a t a d d e d t o t h a t l a y e r o f a i r by t h e e f f e c t of t h e wings would r e s u l t i n t h e s u b s t r a t e t e m p e r a t u r e b e i n g l o w e r t h a n t h e a i r above i t , and t h e e x t r a 1 28 h e a t would t r a n s f e r i n t o t h e s u b s t r a t e h e a t s i n k . Why s h o u l d a b u t t e r f l y d o r s a l bask a t a l l t h e n ? When t h e s u b s t r a t e t e m p e r a t u r e i s l o w e r t h a n t h e a i r t e m p e r a t u r e ( e a r l y m o r n i n g and l a t e e v e n i n g ) , t h e wings w i l l be s p r e a d i n o r d e r t o maximize s o l a r r a d i a t i o n a b s o r p t i o n d i r e c t l y by t h e body (He i n r i c h , 1972; Kammer and B r a c c h i , 1973; R a w l i n s , 1980), and h e a t l o s s w i l l be m i n i m i z e d by r e d u c i n g d i r e c t c o n t a c t w i t h t h e s u b s t r a t e by s u p p o r t i n g t h e body on t h e l e g s . When s u b s t r a t e t e m p e r a t u r e s a r e above a i r t e m p e r a t u r e ( m i d - d a y ) , d i r e c t s o l a r r a d i a t i o n a b s o r p t i o n by t h e body w i l l s t i l l be maximized by d o r s a l b a s k i n g , but a h i g h mass s u b s t r a t e s u c h a s r o c k o r d i r t s h o u l d be c h o s e n and t h e body p r e s s e d t o i t i n o r d e r t o maximize h e a t g a i n f r o m t h e s u b s t r a t e ( C l e n c h , 1966; D o u g l a s , 1978; K e v i n and S h o r t h o u s e , 1970). C a s u a l o b s e r v a t i o n s s u g g e s t t h a t t h i s may be t h e p a t t e r n of s u b s t r a t e c h o i c e and b a s k i n g b e h a v i o r i n P a r n a s s i u s p h oebus. Heat l o s s t o t h e s u b s t r a t e w i l l i n c r e a s e a s t h e d i f f e r e n c e between body t e m p e r a t u r e and s u b s t r a t e t e m p e r a t u r e i n c r e a s e s . S i n c e t h e minimum body t e m p e r a t u r e r e q u i r e d f o r f l i g h t i s c o n s t a n t , t h e low e r t h e s u b s t r a t e t e m p e r a t u r e t h e h i g h e r t h e h e a t l o s s t o i t . T h e r e f o r e t h e l o w e r t h e s u b s t r a t e t e m p e r a t u r e t h e g r e a t e r t h e i n c r e m e n t s i n s o l a r r a d i a t i o n w h i c h a r e needed t o compensate f o r t h e i n c r e a s e d h e a t l o s s . A l s o , t h e low e r t h e a i r t e m p e r a t u r e , t h e l o w e r t h e s u b s t r a t e t e m p e r a t u r e w i l l be under a g i v e n l e v e l of s o l a r r a d i a t i o n . T h e r e f o r e as t h e a i r t e m p e r a t u r e d r o p s , s u b s t r a t e t e m p e r a t u r e w i l l d r o p , r e s u l t i n g i n l a r g e r i n c r e m e n t s i n s o l a r r a d i a t i o n b e i n g needed t o r a i s e t h e 129 body t e m p e r a t u r e t o f l i g h t l e v e l s . T h i s w i l l r e s u l t i n a c u r v e d f l i g h t t h r e s h o l d , s u c h as o c c u r s i n F i g u r e 17, w i t h t h e wings not h a v i n g any e f f e c t on body t e m p e r a t u r e . The i m p o r t a n c e of s u b s t r a t e t e m p e r a t u r e i n t h e d e t e r m i n a t i o n of t h e body t e m p e r a t u r e of b a s k i n g i n s e c t s was a p p a r e n t l y f i r s t r e c o g n i z e d by Gunn ( 1 9 4 2 ) . In R o l a n d ' s (1982) r e g r e s s i o n f o r f a c t o r s a f f e c t i n g body t e m p e r a t u r e of C o l i a s , t h e r e g r e s s i o n c o e f f i c i e n t of 0.625 f o r t h e e f f e c t of a i r t e m p e r a t u r e on body t e m p e r a t u r e was s u b s t a n t i a l l y below t h e 1.0 w h i c h i s e x p e c t e d i f a i r t e m p e r a t u r e and s o l a r r a d i a t i o n a r e t h e o n l y e n v i r o n m e n t a l f a c t o r s a f f e c t i n g body t e m p e r a t u r e . H i s s p e c i m e n s r e s t e d on t h e g r o u n d , and t h e r e f o r e h e a t l o s s t o t h e ground, c o u l d o c c u r . I n c r e a s e s i n body t e m p e r a t u r e due t o t h e g r o u n d t e m p e r a t u r e b e i n g r a i s e d by i n c r e a s e d s o l a r r a d i a t i o n would c o n t r i b u t e t o t h e s o l a r r a d i a t i o n r e g r e s s s i o n c o e f f i c i e n t . The d i f f e r e n c e between g r o u n d t e m p e r a t u r e (minus t h e e f f e c t of s o l a r r a d i a t i o n on i t ) and a i r t e m p e r a t u r e would i n c r e a s e a s a i r t e m p e r a t u r e i n c r e a s e d , and h e a t l o s s t o t h e g r o u n d would t h e r e f o r e i n c r e a s e a s a i r t e m p e r a t u r e i n c r e a s e d . T h i s would r e s u l t i n t h e r e g r e s s i o n c o e f f i c i e n t f o r a i r t e m p e r a t u r e b e i n g s u b s t a n t i a l l y l e s s t h a n 1.0. R o l a n d s t i l l o b t a i n e d p o s i t i v e e f f e c t s f o r wing m e l a n i s m b e c a u s e i n t h e l a t e r a l b a s k i n g p o s i t i o n o f C o l i a s t h e a i r warmed by t h e wings i s above t h e body, and c a n n o t be d i r e c t l y c o o l e d by th e g r o u n d b e f o r e i t t r a n s f e r s i n t o t h e body. 130 F l i g h t and Body T e m p e r a t u r e Once i n f l i g h t an i n s e c t i s s t i l l g a i n i n g h e a t from s o l a r r a d i a t i o n , as w e l l as g a i n i n g h e a t from m u s c u l a r t h e r m o g e n e s i s , and l o s i n g h e a t t h r o u g h c o n v e c t i o n , r e - r a d i a t i o n , and e v a p o r a t i o n ( R a i n e y , 1974). Most l a r g e b u t t e r f l i e s have s l o w , g l i d i n g f l i g h t p a t t e r n s and t r a v e l a t under 9 km/hr ( D o u g l a s , 1978), w i t h ' c o l d ' P. phoebus f l y i n g a t under 6 km/hr ( e s t i m a t e d ) , a l l o w i n g maximum a b s o r p t i o n of s o l a r r a d i a t i o n w i t h l i t t l e d i s t u r b a n c e o f t h e b o u n d a r y l a y e r o f s t i l l a i r n e x t t o t h e base of t h e w i n g s . The h a i r y t h o r a x and wing b a s e s w h i c h a r e p r e s e n t i n many b u t t e r f l i e s , and a r e p a r t i c u l a r i l y p r o n o u n c e d i n P a r n a s s i u s phoebus m a l e s , w i l l widen t h e b o u n d a r y l a y e r of s t i l l a i r a t t h e base of t h e w i n g s . I t i s known t h a t a t wind s p e e d s of a t l e a s t 1.6 km/hr, wing m e l a n i s m a f f e c t s t h e body t e m p e r a t u r e of b u t t e r f l i e s s u s p e n d e d i n m i d - a i r ( D o u g l a s , 1978; D o u g l a s and G r u l a , 1978). T h e r e f o r e a b s o r p t i o n of s o l a r r a d i a t i o n w h i l e i n f l i g h t may d e p e n d i n p a r t on b a s a l wing m e l a n i s m i n P. phoebus and o t h e r s p e c i e s w i t h a slow, g l i d i n g f l i g h t p a t t e r n . In l a t e r a l b a s k i n g s p e c i e s s u c h as C o l i a s , t h e b a s k i n g s u r f a c e i s t h e l o w e r wing s u r f a c e and t h e r e f o r e c a n n o t a b s o r b s o l a r r a d i a t i o n d u r i n g f l i g h t . I f a b s o r p t i o n of s o l a r r a d i a t i o n by b u t t e r f l y wings w h i l e i n f l i g h t i s i m p o r t a n t , v a r i a t i o n i n wing c o l o r may have a g r e a t e r i n f l u e n c e on t h e r m o r e g u l a t i o n w h i l e i n f l i g h t t h a n when d o r s a l b a s k i n g . T h i s h y p o t h e s i s c o u l d be t e s t e d i n t h e f o l l o w i n g ways: ( l ) r e p e a t t h e body t e m p e r a t u r e e x p e r i m e n t a t d i f f e r e n t wind 131 s p e e d s (2-10 km/hr) b o t h when t h e b u t t e r f l i e s a r e on a s u b s t r a t e and when t h e y a r e s u s p e n d e d ; (2) d e t e r m i n e whether d a r k e r i n d i v i d u a l s have l o n g e r f l i g h t s between l a n d i n g s , and t h e r e f o r e a b s o r b more s o l a r r a d i a t i o n d u r i n g f l i g h t t h a n l i g h t i n d i v i d u a l s ; and (3) d e t e r m i n e whether d a r k e r i n d i v i d u a l s have s h o r t e r b a s k i n g p e r i o d s , and t h e r e f o r e a b s o r b more s o l a r r a d i a t i o n w h i l e b a s k i n g t h a n l i g h t e r i n d i v i d u a l s . E i t h e r o f t h e l a t t e r two p o s s i b i l i t i e s would r e s u l t i n d a r k e r male P. phoebus s p e n d i n g a g r e a t e r p r o p o r t i o n o f t o t a l t i m e i n f l i g h t a t low s o l a r r a d i a t i o n and a i r t e m p e r a t u r e , and l e s s t o t a l t i m e b a s k i n g , as I f o u n d . D e t e c t i n g t h e E f f e c t of B a s a l M e l a n i s m on A c t i v i t y An a p p a r e n t anomaly i n my r e s u l t s i s t h a t a l t h o u g h I f o u n d t h a t b a s a l l y d a r k e r m a l e s spend a g r e a t e r p r o p o r t i o n o f t h e i r t i m e i n f l i g h t t h a n l i g h t m a l e s , p a r t i c u l a r i l y a t low s o l a r r a d i a t i o n and a i r t e m p e r a t u r e , t h i s d i d n o t show up as a change i n t h e p r o p o r t i o n of l i g h t t o d a r k m a l e s c a p t u r e d as d i s t a n c e f r o m t h e f l i g h t t h r e s h o l d i n c r e a s e d . T h i s i s b e c a u s e i n o r d e r t o d e t e c t s u c h a change I would have t o sample t h e b u t t e r f l i e s i n f l i g h t i n s t a n t a n e o u s l y , under a g i v e n s e t of c o n d i t i o n s ("scan s a m p l i n g " , A ltmann 1974). S i n c e I was l o o k i n g f o r an a b s o l u t e d i f f e r e n c e i n f l i g h t ( e i t h e r f l y i n g o r not f l y i n g ) , I d i d n ot i n s t a n t a n e o u s l y sample, b ut c a u g h t e v e r y i n d i v i d u a l i n f l i g h t r e g a r d l e s s of how g r e a t a p r o p o r t i o n o f t i m e e a c h i n d i v i d u a l f l e w . S i n c e P a r n a s s i u s a r e e x t r e m e l y e a s y t o see i n il 32 f l i g h t , and v e r y e a s y t o c a p t u r e once s i g h t e d , I e f f e c t i v e l y c a u g h t e v e r y i n d i v i d u a l w h i c h was c a p a b l e o f f l i g h t and was w i t h i n c a p t u r e r a n g e . F e e d i n g F e e d i n g by a d u l t b u t t e r f l i e s i n c r e a s e s l o n g e v i t y and r e p r o d u c t i v e s u c c e s s , and t h e r e f o r e i n d i v i d u a l f i t n e s s (Murphy e t a l . , 1983, 1984). T h e r e f o r e i f wing m e l a n i s m a f f e c t s t h e a b i l i t y of a b u t t e r f l y t o f e e d , i t w i l l a l s o a f f e c t r e l a t i v e f i t n e s s . F e e d i n g a c t i v i t y by C o l i a s i s l i m i t e d by b a s a l m e l a n i s m b e c a u s e the minimum f l i g h t t h r e s h o l d i s d e t e r m i n e d i n p a r t by wing m e l a n i s m ( R o l a n d , 1982), and t h e r e f o r e d a r k e r i n d i v i d u a l s c a n f l y between f l o w e r s under c o l d e r c o n d i t i o n s t han l i g h t i n d i v i d u a l s . In c o n t r a s t , f e e d i n g i n P. phoebus i s n o t a f u n c t i o n of p h e n o t y p e d e p e n d e n t f l i g h t t h r e s h o l d s . The e f f e c t o f b a s a l m e l a n i s m on p r o p o r t i o n of t i m e s p e n t i n f l i g h t i s n o t r e f l e c t e d i n d i f f e r e n c e s i n t h e amount o f t i m e s p e n t f e e d i n g ( F i g u r e 2 4 ) , b e c a u s e n e c t a r i n g i s combined w i t h d o r s a l b a s k i n g , and d o r s a l b a s k i n g a p p a r e n t l y does not r e s u l t i n p h e n o t y p e d e p e n d e n t body t e m p e r a t u r e s . The o n l y p o s s i b l e e f f e c t t h a t m e l a n i s m w i l l have on body t e m p e r a t u r e (and t h e r e f o r e a c t i v i t y ) w h i l e f e e d i n g i s t h a t d a r k e r i n d i v i d u a l s may be warmer when t h e y i n i t i a l l y l a n d on a f l o w e r . The t e m p e r a t u r e of t h e b u t t e r f l i e s s h o u l d c o n v e r g e on t h e t e m p e r a t u r e of t h e f l o w e r , b o t h t h r o u g h d i r e c t h e a t t r a n s f e r ( c o n d u c t i o n , c o n v e c t i o n , r a d i a t i o n ) and t h r o u g h t h e e f f e c t of t a k i n g i n n e c t a r . S i n c e n e c t a r i n t a k e by b u t t e r f l i e s 133 may be up t o 43-46% of t h e p r e - f e e d i n g body w e i g h t ( N i e l s e n and N i e l s e n , 1959), i t has t h e p o t e n t i a l f o r l a r g e e f f e c t s on body t e m p e r a t u r e . I n d i v i d u a l s f o r w h i c h t h e r e i s a g r e a t e r d i f f e r e n c e between body and f l o w e r t e m p e r a t u r e s w i l l change t e m p e r a t u r e most r a p i d l y , and any i n i t i a l d i f f e r e n c e due t o e x t r a wing m e l a n i s m w i l l be l o s t . S t a r v e d P. phoebus w i l l s p o n t a n e o u s l y c r a w l a r o u n d and f e e d w i t h body t e m p e r a t u r e s as low as 6°C ( l a b o r a t o r y o b s e r v a t i o n ) , t h e r e f o r e f e e d i n g w i l l n ot be l i m i t e d by any body t e m p e r a t u r e s above t h e minimum r e q u i r e d f o r f l i g h t . S c o t t (1973a) f o u n d t h a t P. phoebus f e e d i n g p e a k e d between 10:30 and 13:30 h o u r s , and l i t t l e f e e d i n g was done i n t h e e a r l y m o r n i n g . S u c c e s s f u l n e c t a r i n g r e q u i r e s t h a t b u t t e r f l i e s f l y between f l o w e r s , w h i c h i n t u r n r e q u i r e s a body t e m p e r a t u r e above the minimum r e q u i r e d f o r f l i g h t . In t h e e a r l y m o r n i n g , t h e body t e m p e r a t u r e w i l l be below t h e minimum needed f o r f l i g h t much of t h e t i m e , r e s u l t i n g i n c o m p a r a t i v e l y low f l i g h t a c t i v i t y and t h e r e f o r e l i m i t e d f e e d i n g . However, P. phoebus i s l i k e l y t o be most hungry i n t h e m o r n i n g , w h i c h s h o u l d r e s u l t i n most f e e d i n g o c c u r r i n g b e f o r e mid-day p r o v i d e d a i r t e m p e r a t u r e s a r e h i g h enough. I f o u n d t h a t more f e e d i n g o c c u r r e d a t l o w e r a i r t e m p e r a t u r e s and s o l a r r a d i a t i o n s , a s would be e x p e c t e d i f most f e e d i n g does o c c u r b e f o r e mid-day. 1 34 S u r v i v a l The s u r v i v a l r a t e f o r m a l e s i n t h e V e r n o n H i l l c o l o n y was e x t r e m e l y h i g h f o r P. phoebus, w i t h a mean r e s i d e n c e t i m e of 11 d a y s (15 d a ys i f t h e g r o u p of o n e - t i m e - o n l y c a p t u r e s i s o m i t t e d ) . T h i s i s d o u b l e t h e e s t i m a t e s of a b o u t 6 d a ys by S c o t t (1973a,b) and 5-9 d a y s by C l i f f o r d ( 1 9 8 1 ) , and i s p r o b a b l y t h e r e s u l t o f t h e c o n t i n u o u s f o r e s t b a r r i e r i n my a r e a i n h i b i t i n g d i s p e r s a l , whereas t h e i r a r e a s d i d not have c o n t i n u o u s f o r e s t b a r r i e r s . C l i f f o r d (1981) f o u n d a maximum r e s i d e n c e t i m e of 21 d a y s , compared w i t h my 39 d a y s . In an unbounded p o p u l a t i o n , a g e - i n d e p e n d e n t d i s p e r s a l , r a t h e r t h a n a g e - d e p e n d e n t m o r t a l i t y may be t h e dominant ' m o r t a l i t y ' f a c t o r . I p r e d i c t t h a t i n an unbounded p o p u l a t i o n c u m u l a t i v e s u r v i v a l c u r v e s f o r P a r n a s s i u s w i l l d e c a y e x p o n e n t i a l l y as f o r C o l i a s ( R o l a n d , 1982), r a t h e r t h a n l i n e a r l y . S u r v i v a l r a t e s w i l l depend on b a s a l wing b l a c k n e s s i n males and s i z e i n f e m a l e s , s i n c e t h o s e c h a r a c t e r s a f f e c t d i s p e r s a l r a t e s . S m a l l e r males had l o w e r s u r v i v a l r a t e s t h a n l a r g e r m a l e s . The r e a s o n f o r t h i s i s n o t c l e a r f r o m t h e d a t a , s i n c e t h e most l i k e l y r e a s o n , i n c r e a s e d d i s p e r s a l by s m a l l m a l e s , a p p a r e n t l y d o e s n o t o c c u r . A n o t h e r p o s s i b l e r e a s o n f o r t h e more r a p i d d i s a p p e a r a n c e of s m a l l m a l e s i s t h a t t h e y had a h i g h e r m o r t a l i t y r a t e , w h i c h c o u l d not be t e s t e d w i t h t h e a v a i l a b l e d a t a . 1 35 D i s p e r s a l I f h a b i t a t s u i t a b l e f o r a s p e c i e s ' r e p r o d u c t i o n i s p a t c h y b u t s t a b l e , d i s p e r s a l r a t e s s h o u l d be low. F o r s u c h h a b i t a t s , i f a d i s p e r s e r f i n d s a n o t h e r c o l o n y i t s r e p r o d u c t i v e s u c c e s s i s u n l i k e l y t o be any b e t t e r t h e r e t h a n a t t h e o r i g i n a l s i t e , and i t w i l l have undergone t h e r i s k s a s s o c i a t e d w i t h d i s p e r s a l . I f s u i t a b l e h a b i t a t f o r r e p r o d u c t i o n i s e p h e m e r a l , o r t h e p r o b a b i l i t y of f i n d i n g a new p a t c h of s u i t a b l e h a b i t a t i s h i g h , d i s p e r s a l s h o u l d be h i g h i n o r d e r t o i n s u r e t h a t when t h e o l d h a b i t a t d i s a p p e a r s t h e r e a r e o f f s p r i n g of t h e p r e v i o u s g e n e r a t i o n i n o t h e r a r e a s ( B a k e r , 1969; Southwood, 1962). S i n c e t h e r e i s a p o s s i b i l i t y of even a ' s t a b l e ' h a b i t a t d i s a p p e a r i n g , some d i s p e r s a l s h o u l d be a d v a n t a g e o u s f o r a l l s p e c i e s . The o v e r a l l d i s p e r s a l r a t e o f P. phoebus i s a p p a r e n t l y q u i t e low, a p p a r e n t l y b e c a u s e a r e a s o f s u i t a b l e h a b i t a t a r e u s u a l l y s t a b l e and i s o l a t e d f r o m e a c h o t h e r . E u p h y d r a s e d i t h a , w h i c h i n h a b i t s s i m i l a r h a b i t a t s , a l s o has a low d i s p e r s a l r a t e ( E h r l i c h e t a l . , 1984). In c o n t r a s t , s p e c i e s s u c h as P i e r i s  r a p a e and P i e r i s p r o t o d i c e ( w h i c h i n h a b i t e p h e m e r a l h a b i t a t s ) , o r E r e b i a e p i p s o d e a ( f o r w h i c h t h e p r o b a b i l i t y of a d i s p e r s e r f i n d i n g s u i t a b l e h a b i t a t i s h i g h ) , have h i g h d i s p e r s a l r a t e s ( B a k e r , 1968; B r u s s a r d and E h r l i c h , 1970; Emmel, 1973). T h e r e a r e s e v e r a l p o s s i b l e r e a s o n s f o r t h e g r e a t e r d i s p e r s a l o f f e m a l e t h a n male P. p h o e b u s . F e m a l e s of s p e c i e s i n h a b i t i n g p a t c h y e n v i r o n m e n t s s h o u l d have a g r e a t e r t e n d e n c y t o d i s p e r s e t h a n m a l e s b e c a u s e t h e y o n l y need t o f i n d s u i t a b l e h a b i t a t t o o v i p o s i t . M a l e s s h o u l d t e n d t o r e m a i n i n a r e a s w h i c h 136 have ( v i r g i n ) f e m a l e s w i t h w h i c h t o mate, w h i c h g e n e r a l l y means th e a r e a i n w h i c h t h e y emerged. O c c a s i o n a l l y a 'temporary' h a b i t a t may open up, s u c h as t h e summit of W h i t e Rocks Mtn n e a r Kelowna, B.C., w h i c h has r e c e n t l y been b u r n e d o f f l e a v i n g an u n c o l o n i z e d i s l a n d of s u i t a b l e h a b i t a t i n t h e m i d d l e of e x t e n s i v e f o r e s t . The p r o b a b i l i t y of a d i s p e r s i n g f e m a l e f i n d i n g s u c h a new, u n c o l o n i z e d h a b i t a t w i l l be v e r y low, but a f e m a l e w h i c h e s t a b l i s h e s a new c o l o n y may be r e p r o d u c t i v e l y v e r y s u c c e s s f u l , s i n c e t h e e n t i r e new c o l o n y w i l l be composed of h e r o f f s p r i n g ( H a r r i s o n , 1980). T h i s may a l s o be t h e c a s e f o r E u p h y d r a s e d i t h a (Nymphal i d a e ) ( E h r l i c h e_t a_l. , 1984). Female P. phoebus a r e f r e q u e n t l y h a r a s s e d by males a t t e m p t i n g t o mate w i t h them, even t h o u g h a s e c o n d m a t i n g i s n o t p o s s i b l e due t o t h e g e n i t a l p l u g d e p o s i t e d d u r i n g t h e f i r s t m a t i n g . T h i s h a r a s s m e n t may r e s u l t i n i n c r e a s e d f e m a l e d i s p e r s a l , as has been p r o p o s e d f o r P i e r i s p r o t o d i c e ( P i e r i d a e ) ( S h a p i r o , 1970), and Danaus p l e x i p p u s ( D a n i d a e ) (R. K i t c h i n g , pers.comm.). Mean r e s i d e n c e t i m e on V e r n o n H i l l was o v e r t w i c e as l o n g as f o u n d by S c o t t (1973b) and C l i f f o r d (1981) i n C o l o r a d o , p r o b a b l y b e c a u s e o f d i f f e r e n c e s i n d i s p e r s a l r a t e s . V a r i a t i o n i n d i s p e r s a l c h a r a c t e r i s t i c s a p p a r e n t l y o c c u r s between p o p u l a t i o n s of many o t h e r b u t t e r f l y s p e c i e s ( G i l b e r t and S i n g e r , 1975), due t o e i t h e r g e n e t i c , e n v i r o n m e n t a l , or h a b i t a t boundary d i f f e r e n c e s ( G i l b e r t and S i n g e r , 1975; Southwood, 1962). U n q u a n t i f i e d o b s e r v a t i o n s i n d i c a t e t h a t P. phoebus t r e a t s d ense f o r e s t as a b o u n d a r y , but u n s u i t a b l e g r a s s l a n d and open f o r e s t 1 37 do n o t g r e a t l y l i m i t d i s p e r s a l ( C D . F e r r i s , _in 1 i t t . ) . T h e r e f o r e h a b i t a t b oundary d i f f e r e n c e s between t h e C o l o r a d o p o p u l a t i o n s and t h e V e r n o n H i l l p o p u l a t i o n p r o b a b l y a c c o u n t s f o r most o f t h e d i f f e r e n c e i n r e s i d e n c e t i m e between t h e two a r e a s , a l t h o u g h g e n e t i c d i f f e r e n c e s c a n n o t be r u l e d o u t . The r e l a t i v e d i s p e r s a l r a t e s of t h e s e x e s may a l s o v a r y between p o p u l a t i o n s of b u t t e r f l i e s . The P i e r i d b u t t e r f l y P i e r i s r a p a e shows no e f f e c t o f sex on d i s p e r s a l i n t h e e a s t e r n U.S.A. (Emmel, 1973), but i n A u s t r a l i a f e m a l e s a p p a r e n t l y d i s p e r s e more t h a n males (R.E. J o n e s , pers.comm.). M a l e s w i t h g r e a t e r b a s a l p a t c h b l a c k n e s s t e n d t o d i s p e r s e more t h a n l i g h t e r m a l e s , p r o b a b l y b e c a u s e t h e y spend a g r e a t e r p r o p o r t i o n of t i m e i n f l i g h t , and t h e r e f o r e w i l l t e n d t o f l y f a r t h e r . I p r e d i c t t h a t t h e p r o b a b i l i t y o f d i s p e r s a l a c r o s s a m ajor f o r e s t b a r r i e r w i l l not be d ependent on b a s a l p a t c h b l a c k n e s s , a s s u m i n g t h a t t h e ' d e c i s i o n ' t o c r o s s i s i n d e p e n d e n t of t h e p r o p o r t i o n of t i m e i n f l i g h t . T h i s a s s u m p t i o n t h a t movement a c r o s s major f o r e s t b a r r i e r s i s q u a l i t a t i v e l y d i f f e r e n t from movement between d i f f e r e n t a r e a s of g r a s s l a n d i s r e a s o n a b l e , b e c a u s e : (a) most o f t h o s e m ales w h i c h moved a c r o s s t h e f o r e s t b a r r i e r i n t o A r e a s 4-7 had been c a p t u r e d o n l y once i n A r e a 2 b e f o r e m o ving, w h i c h s u g g e s t s t h a t t h e r e may be an i n h e r e n t d i f f e r e n c e between l o n g - d i s t a n c e d i s p e r s e r s and n o n - d i s p e r s e r s , and (b) t h e r e was no s i g n i f i c a n t e f f e c t of b a s a l p a t c h b l a c k n e s s on ' t h e p r o b a b i l i t y of an i n d i v i d u a l b e l o n g i n g t o t h e ' o n e - t i m e -o n l y ' c a p t u r e g r o u p , w h i c h i n d i c a t e s t h a t t h e p r o p o r t i o n of t i m e 138 s p e n t i n f l i g h t has no e f f e c t on t h e p r o b a b i l i t y of l e a v i n g t h e p o p u l a t i o n . L a r g e r f e m a l e s t e n d t o d i s p e r s e more t h a n s m a l l e r o n e s . S i n c e s i z e d i d n o t a f f e c t t h e p r o p o r t i o n o f t i m e males spend i n f l i g h t , t h e r e i s no r e a s o n t o s u p p o s e t h a t l a r g e r f e m a l e s s p e n d more time i n f l i g h t . I t i s u n l i k e l y t h e r e f o r e t h a t t h e g r e a t e r t e n d e n c y f o r l a r g e f e m a l e s t o d i s p e r s e i s due t o i n c r e a s e d f l i g h t a c t i v i t y . The c o r r e l a t i o n between s i z e and f e m a l e d i s p e r s a l may be b e c a u s e t h e e n e r g e t i c and r e p r o d u c t i v e c o s t o f d i s p e r s a l d e c r e a s e s w i t h i n c r e a s e d s i z e , as i n D r o s o p h i l a ( D i p t e r a ) ( R o f f , 1977). Adapt i v e S i g n i f i c a n c e o f V a r i a t i o n i n M e l a n i sm and S i z e S i n c e d a r k e r m ales ( b a s a l p a t c h b l a c k n e s s ) move l o n g e r d i s t a n c e s , and most p o p u l a t i o n s do n o t have a c o n t i n u o u s f o r e s t b o u ndary, t h e y w i l l t e n d t o be l o s t from t h e p o p u l a t i o n . However, w h i l e t h e y a r e w i t h i n a p o p u l a t i o n t h e y w i l l s p end a g r e a t e r p r o p o r t i o n of t i m e i n f l i g h t , and t h e r e f o r e w i l l have a g r e a t e r p r o b a b i l i t y of f i n d i n g a v i r g i n f e m a l e . At h i g h s o l a r r a d i a t i o n and a i r t e m p e r a t u r e t h e r e i s no d e pendence o f t i m e s p e n t i n f l i g h t on p h e n o t y p e , and l i g h t e r m a l e s have a l o w e r p r o b a b i l i t y o f l o s s t h r o u g h d i s p e r s a l . D a r k e r m a l e s w i l l have an a d v a n t a g e i n f i n d i n g f e m a l e s a t low l e v e l s of s o l a r r a d i a t i o n and a i r t e m p e r a t u r e b e c a u s e t h e added t i m e i n f l i g h t w i l l p r o b a b l y o u t w e i g h the added p r o b a b i l i t y of d i s p e r s a l . I n c r e a s e d t i m e i n f l i g h t i n t h e e a r l y m o r n i n g (low s o l a r r a d i a t i o n and low a i r t e m p e r a t u r e ) , s h o u l d r e s u l t i n a s u b s t a n t i a l i n c r e a s e i n t h e 1 39 p r o b a b i l i t y of a male f i n d i n g a mate be c a u s e most f e m a l e s emerge and mate a t t h a t t i m e ( S c o t t , 1973a). L a r g e r f e m a l e s w i l l have a r e p r o d u c t i v e a d v a n t a g e b e c a u s e egg p r o d u c t i o n i s p o s i t i v e l y c o r r e l a t e d w i t h s i z e i n b u t t e r f l i e s ( B a k e r , 1968; J o n e s e t a l . , 1982). T h i s a d v a n t a g e i n t o t a l egg p r o d u c t i o n w i l l t e n d t o be o f f s e t by t h e i n c r e a s e d d i s p e r s a l r a t e of l a r g e f e m a l e s . Most of my e v i d e n c e f o r t h e a d a p t i v e s i g n i f i c a n c e o f v a r i a t i o n i n m e l a n i s m i s f o r m a l e s , b e c a u s e f e m a l e s a r e much more d i f f i c u l t t o s t u d y . A p p a r e n t l y males spend a l l , or n e a r l y a l l , t h e i r t i m e e i t h e r i n f l i g h t or f e e d i n g i f t h e i r body t e m p e r a t u r e i s h i g h enough. T h e i r f l i g h t p a t t e r n i s a slow, smooth ' z i g - z a g ' p a t r o l w i t h i n one meter o f t h e t o p of t h e v e g e t a t i o n . M a l e s a r e u s u a l l y d o r s a l b a s k i n g when seen on t h e g r o u n d or f e e d i n g . F e m a l e s , i n c o n t r a s t , a p p a r e n t l y spend as l i t t l e t i m e i n f l i g h t a s p o s s i b l e , and when i n f l i g h t t r a v e l r a p i d l y , low t o t h e g r o u n d , and i n a s t r a i g h t l i n e . F e m a l e s s e e n on t h e g r o u n d or f e e d i n g a r e u s u a l l y v e r t i c a l b a s k i n g or body b a s k i n g , e x c e p t u n d e r c o n d i t i o n s o f low s o l a r r a d i a t i o n and a i r t e m p e r a t u r e when t h e y d o r s a l b a s k . From t h e s e b e h a v i o r a l d i f f e r e n c e s I p r e d i c t t h a t t h e t i m e s p e n t i n f l i g h t by f e m a l e s w i l l n o t be a f f e c t e d by wing c o l o r , b e c a u s e once above t h e minimum f l i g h t t h r e s h o l d t h e i r f l i g h t i s a p p a r e n t l y n o t d e p e n d e n t on body t e m p e r a t u r e . Wing c o l o r w i l l have a t h e r m o r e g u l a t o r y e f f e c t i n f e m a l e s o n l y i f t h e r e i s an e f f e c t o f wing c o l o r on body t e m p e r a t u r e w h i l e b a s k i n g on a s u b s t r a t e , b e c a u s e t h e t i m e s p e n t i n f l i g h t i s t o o low f o r h e a t 1 40 a b s o r p t i o n w h i l e f l y i n g t o be an i m p o r t a n t f a c t o r . I t i s p o s s i b l e t h a t t h e c o r r e l a t i o n between s e x e s f o r b a s a l p a t c h b l a c k n e s s ( C h a p t e r l ) i s t h e r e s u l t of t h e same genes b e i n g e x p r e s s e d i n f e m a l e s as i n m a l e s , r a t h e r t h a n b a s a l p a t c h b l a c k n e s s h a v i n g an a d a p t i v e s i g n i f i c a n c e i n b o t h s e x e s . The a d a p t i v e s i g n i f i c a n c e of v a r i a t i o n i n f e m a l e wing m e l a n i s m , e s p e c i a l l y t h e h i g h l y v a r i a b l e d i s t a l a r e a , w i l l be an i n t e r e s t i n g a r e a of f u t u r e i n v e s t i g a t i o n . V a r i a t i o n i n d i s t a l m e l a n i s m has no e f f e c t on body t e m p e r a t u r e or a c t i v i t y , and i t s a d a p t i v e s i g n i f i c a n c e i s a n o t h e r a r e a f o r f u t u r e i n v e s t i g a t i o n . 141 GENERAL DISCUSSION I have d e s c r i b e d t h e p a t t e r n s of v a r i a t i o n i n s i x p h e n o t y p i c c h a r a c t e r s w i t h r e s p e c t t o t h e i r c o r r e l a t i o n s w i t h e l e v a t i o n , l a t i t u d e , mean a i r t e m p e r a t u r e , and e a c h other.. From t h e s e o b s e r v a t i o n s I f o r m u l a t e d h y p o t h e s e s as t o t h e a d a p t i v e s i g n i f i c a n c e of v a r i a t i o n i n e a c h c h a r a c t e r . I t h e n s e l e c t e d t h o s e h y p o t h e s e s r e l a t e d t o t h e t h e r m o r e g u l a t o r y s i g n i f i c a n c e of v a r i a t i o n i n e a c h c h a r a c t e r and p r e d i c t e d t h e e f f e c t o f e a c h on body t e m p e r a t u r e and a c t i v i t y . The r e s u l t s a r e summarized i n T a b l e X I I . I c o u l d not t e s t a l l t h e p r e d i c t i o n s f u l l y , but s e v e r a l c o n c l u s i o n s can be drawn: (1) V a r i a t i o n i n b a s a l p a t c h b l a c k n e s s has a t h e r m o r e g u l a t o r y e f f e c t w h i c h has b o t h an a d a p t i v e a d v a n t a g e ( i n c r e a s e d f l i g h t a c t i v i t y ) and a l o c a l d i s a d v a n t a g e ( i n c r e a s e d d i s p e r s a l ) ; (2) V a r i a t i o n i n b a s a l p a t c h w i d t h has a t h e r m o r e g u l a t o r y e f f e c t w h i c h has an a d a p t i v e a d v a n t a g e ( i n c r e a s e d f l i g h t a c t i v i t y ) , b u t no d i s a d v a n t a g e was d e t e c t e d . (3) V a r i a t i o n i n b a s a l t r a n s p a r e n c y has o n l y a s m a l l e f f e c t on body t e m p e r a t u r e , w h i c h has no e f f e c t on a c t i v i t y . (4) V a r i a t i o n i n s i z e has an e f f e c t on s u r v i v a l and d i s p e r s a l . I t must have an a d a p t i v e s i g n i f i c a n c e o t h e r t h a n t h a t o f a f f e c t i n g t h e r m o r e g u l a t i o n b e c a u s e i t does n o t a f f e c t f l i g h t a c t i v i t y . T h e r e f o r e a l t h o u g h s i z e i s wea k l y c o r r e l a t e d w i t h b a s a l p a t c h w i d t h and' b l a c k n e s s , t h e y have q u i t e d i f f e r e n t a d a p t i v e s i g n i f i c a n c e s . (5) V a r i a t i o n i n d i s t a l wing b l a c k n e s s and t r a n s p a r e n c y has no 142 T a b l e X I I . Summary of v a r i a t i o n i n wing p h e n o t y p e and i t s c o r r e l a t i o n s w i t h a c t i v i t y . C h a r a c t e r Sex E l e v a t i o n & L a t i t u d e Body Temp. Time i n F l i g h t S u r -v i v a l D i s -p e r s a l G e n e t i c : E n v i r . * * BASAL M + po s . p o s . ns ns g e n e t i c WIDTH F ns ns BASAL M + + + p o s . p o s . ns p o s . g e n e t i c ? BLACKNESS F + + ns e n v i r o n . BASAL M + neg. ns ns ns g e n e t i c TRANSPAR. F + ns e n v i r o n . DISTAL M l o c a l + + none* ns ns ns g e n e t i c ? BLACKNESS F l o c a l + + none* ns DISTAL M l o c a l + + none* ns ns ns g e n e t i c TRANSPAR. F l o c a l + + none* ns WING M y e s * ns p o s . ns g e n e t i c LENGTH F y e s * p o s . e n v i r o n . * R e s u l t s from D o u g l a s (1978) f o r b u t t e r f l i e s i n g e n e r a l . * * C o n c l u s i o n s on whether v a r i a t i o n i n a c h a r a c t e r i s due t o g e n e t i c d i f f e r e n c e s o r d e v e l o p m e n t a l e n v i r o n m e n t , t h e y a r e a p p l i c a b l e t o b o t h s e x e s . B l a n k s = not t e s t e d . ns = no s i g n i f i c a n t c o r r e l a t i o n f o u n d ; + = v e r y weak p o s i t i v e c o r r e l a t i o n ; ++ = weak p o s i t i v e c o r r e l a t i o n ; +++ = s t r o n g p o s i t i v e c o r r e l a t i o n ; = s t r o n g n e g a t i v e c o r r e l a t i o n ; p o s . = p o s i t i v e c o r r e l a t i o n ; neg. = n e g a t i v e c o r r e l a t i o n . t h e r m o r e g u l a t o r y e f f e c t on a c t i v i t y . ( 6 ) V a r i a t i o n i n most, and p o s s i b l y a l l , of t h e c h a r a c t e r s i s due t o b o t h g e n e t i c d i f f e r e n c e s and d i f f e r e n c e s i n d e v e l o p m e n t a l e n v i r o n m e n t . The i n f o r m a t i o n a v a i l a b l e f o r t h e e f f e c t s of b a s a l p a t c h b l a c k n e s s on a c t i v i t y i s s u f f i c i e n t t o p r o p o s e an e x p l a n a t i o n of how n a t u r a l s e l e c t i o n c o u l d p r o d u c e t h e o b s e r v e d p a t t e r n of v a r i a t i o n i n male b a s a l p a t c h b l a c k n e s s w i t h i n and between p o p u l a t i o n s . The p r e s e n c e o f b o t h an a d a p t i v e a d v a n t a g e and d i s a d v a n t a g e 1 43 f o r b a s a l p a t c h b l a c k n e s s w i l l t e n d t o r e s u l t i n e a c h p o p u l a t i o n s t a b i l i z i n g a t t h e ' o p t i m a l ' p h e n o t y p e f o r e a c h e n v i r o n m e n t . The ' o p t i m a l ' p h e n o t y p e w i l l be t h e one w h i c h r e s u l t s i n t h e minimum r e d u c t i o n , t h r o u g h d i s p e r s a l , i n t h e p r o b a b i l i t y of m a t i n g r e l a t i v e t o t h e g a i n f r o m i n c r e a s e d t i m e i n f l i g h t . I f t h e e n v i r o n m e n t were s t a b l e and v a r i a t i o n s t r o n g l y h e r i t a b l e , t h a t optimum p h e n o t y p e would be a s p e c i f i c l e v e l of m e l a n i s m and w i t h i n - p o p u l a t i o n v a r i a t i o n would be low. However, v a r i a t i o n i n b a s a l b l a c k n e s s i s a t b e s t o n l y m o d e r a t e l y h e r i t a b l e , and e n v i r o n m e n t a l v a r i a b i l i t y w i t h i n a p o p u l a t i o n i s a c t u a l l y v e r y l a r g e due t o f l u c t u a t i o n s i n s o l a r r a d i a t i o n and a i r t e m p e r a t u r e w i t h i n d a y s , between d a y s , w i t h i n a f l i g h t s e a s o n , and between f l i g h t s e a s o n s . D u r i n g a s i n g l e day, males w i t h h i g h b a s a l p a t c h b l a c k n e s s w i l l have an s e l e c t i v e a d v a n t a g e i n t h e e a r l y m o r n i n g b e c a u s e t h e y can spend a g r e a t e r p r o p o r t i o n of t i m e i n f l i g h t . S i n c e f e m a l e s t e n d t o emerge i n t h e m o r n i n g , f r e q u e n t l y mate i m m e d i a t e l y a f t e r emergence b e f o r e t h e i r wings have expanded, and o n l y mate once ( S c o t t , 1973a), i n c r e a s e d f l i g h t i n t h e m o r n i n g w i l l g r e a t l y i n c r e a s e t h e p r o b a b i l i t y of m a t i n g . L a t e r i n t h e day, d a r k e r m a l e s w i l l n o t be i n f l i g h t anymore t h a n l i g h t o n e s , but d a r k males w i l l be d i s p e r s i n g from t h e p o p u l a t i o n more b e c a u s e of t h e p o s i t i v e c o r r e l a t i o n between f l i g h t s p e e d and body t e m p e r a t u r e i n b u t t e r f l i e s (Rammer, 1970). The a m p l i t u d e of t h e d i u r n a l c y c l e s i n a i r t e m p e r a t u r e and s o l a r r a d i a t i o n change f r o m day t o day, and t h e r e f o r e t h e r e l a t i v e ' d a i l y ' f i t n e s s o f m a l e s w i t h d i f f e r e n t l e v e l s o f m e l a n i s m w i l l 144 v a r y . D i f f e r e n t p a r t s of t h e f l i g h t s e a s o n , and d i f f e r e n t y e a r s , w i l l have d i f f e r e n t mean a i r t e m p e r a t u r e s and s o l a r r a d i a t i o n as d i f f e r e n t weather s y s t e m s move t h r o u g h an a r e a . S i n c e b o t h e n v i r o n m e n t a l v a r i a b l e s and b a s a l p a t c h b l a c k n e s s a r e c o n t i n u o u s v a r i a b l e s , most l e v e l s o f b a s a l b l a c k n e s s w i l l be ' o p t i m a l ' a t some t i m e . As a r e s u l t a v e r y b r o a d range of p h e n o t y p e s w i l l have a r o u g h l y e q u a l r e l a t i v e f i t n e s s o v e r s e v e r a l g e n e r a t i o n s . The i n d i v i d u a l s w i t h t h e most extreme p h e n o t y p e s w i l l have t h e l o w e s t p r o b a b i l i t y of e n c o u n t e r i n g t h e e n v i r o n m e n t a l c o n d i t i o n s under w h i c h t h e i r r e l a t i v e f i t n e s s i s h i g h e s t , and t h e r e f o r e t h e y w i l l be s e l e c t e d a g a i n s t most of t h e t i m e . As a r e s u l t p h e n o t y p i c v a r i a t i o n w i l l t e n d t o r e m a i n w i t h i n a l i m i t e d r a n g e . When a new p o p u l a t i o n i s f o u n d e d a t a new mean a i r t e m p e r a t u r e and s o l a r r a d i a t i o n , t h e p h e n o t y p e s whose ' o p t i m a l ' e n v i r o n m e n t a l c o n d i t i o n s a r e f u r t h e s t from t h e new ones w i l l be s e l e c t e d a g a i n s t most f r e q u e n t l y , and t h e p o p u l a t i o n mean w i l l s h i f t t o w a r d s a new a v e r a g e optimum p h e n o t y p e . An ' o p t i m a l ' p h e n o t y p e i s t h e r e f o r e o n l y d e f i n a b l e i n t e r m s of a s p e c i f i c s e t of e n v i r o n m e n t a l c o n d i t i o n s , and no s i n g l e p h e n o t y p e w i l l have t h e h i g h e s t r e l a t i v e f i t n e s s more t h a n a s m a l l p r o p o r t i o n of t h e t ime. I have assumed i n t h i s d i s c u s s i o n t h a t t h e ' p o p u l a t i o n s ' a r e g e n e t i c a l l y i s o l a t e d , and t h e r e f o r e can d i v e r g e from e a c h o t h e r . However, the a c t of f o u n d i n g a new p o p u l a t i o n i m p l i e s t h a t d i s p e r s a l c a n , and p r o b a b l y w i l l , o c c u r between the two p o p u l a t i o n s . S i n c e p o p u l a t i o n s o f E u p h y d r y a s b u t t e r f l i e s can 145 g e n e t i c a l l y d i v e r g e on a s i n g l e r i d g e w i t h exchange r a t e s of 3% pe r y e a r ( E h r l i c h e t a l . , 1975), low r a t e s of i m m i g r a t i o n w i l l n ot p r e v e n t g e n e t i c d i v e r g e n c e i f s e l e c t i o n i s a c t i n g , . The o f f s p r i n g o f i m m i g r a n t s from a d i f f e r e n t e n v i r o n m e n t w i l l t e n d t o f a l l i n t o t h e 'extreme p h e n o t y p e ' g r o u p s , and t h e r e f o r e w i l l be r e p r o d u c t i v e l y l e s s s u c c e s s f u l t h a n t h e n a t i v e i n d i v i d u a l s . . The a c t u a l gene f l o w w i l l t h e r e f o r e t e n d t o be much l e s s t h a n t h e i m m i g r a t i o n r a t e , even a s s u m i n g t h a t i m m i g r a n t s have an e q u a l i n i t i a l r e p r o d u c t i v e s u c c e s s r e l a t i v e t o t h e n a t i v e p o p u l a t i o n , w h i c h i s u n l i k e l y ( E h r l i c h e_t a_l. , 1975). The p a t t e r n s of v a r i a t i o n i n b a s a l p a t c h b l a c k n e s s b o t h w i t h i n and between p o p u l a t i o n s c an t h e r e f o r e be e x p l a i n e d ( a t l e a s t i n p a r t ) i n terms of e n v i r o n m e n t a l h e t e r o g e n e i t y o v e r s p a c e and t i m e , w i t h o r w i t h o u t low r a t e s of i m m i g r a t i o n . B o t h t h e e f f e c t s of s e l e c t i o n a c t i n g t o change t h e mean p o p u l a t i o n g e n o t y p e , and t h e e f f e c t o f d i f f e r e n t d e v e l o p m e n t a l e n v i r o n m e n t s , w i l l c o n t r i b u t e t o t h e v a r i a t i o n i n b a s a l p a t c h b l a c k n e s s w i t h i n and between p o p u l a t i o n s . I have d e v e l o p e d a new h y p o t h e s i s s a y i n g t h a t t h e e f f e c t s o f t h e w i n g s on t h e r m o r e g u l a t i o n i n d o r s a l b a s k i n g b u t t e r f l i e s i s more i m p o r t a n t w h i l e i n f l i g h t t h a n w h i l e b a s k i n g on a s u b s t r a t e , and d e s c r i b e d f o r t h e p u r p o s e o f f u t u r e r e s e a r c h t h e t e s t s f o r t h i s h y p o t h e s i s . F u r t h e r r e s e a r c h s i m i l a r t o t h a t w h i c h I have done h e r e w i l l p r o b a b l y d e t e c t an a d a p t i v e d i s a d v a n t a g e t o b a s a l p a t c h w i d t h , w h i c h I b e l i e v e w i l l be a weak c o r r e l a t i o n between p a t c h w i d t h and p r o b a b i l i t y of d i s p e r s a l . A p r o f i t a b l e a r e a o f r e s e a r c h on t h e a d a p t i v e 1 46 s i g n i f i c a n c e of v a r i a t i o n i n s i z e w i l l p r o b a b l y be t o i n v e s t i g a t e t h e e f f e c t s of d e c r e a s i n g l e n g t h of g r o w i n g s e a s o n and p o s s i b l y a d e c r e a s e i n f o o d q u a l i t y w i t h i n c r e a s e d e l e v a t i o n . The p a t t e r n s o f v a r i a t i o n i n d i s t a l b l a c k n e s s and t r a n s p a r e n c y d e f i n i t e l y c a n n o t be a c c o u n t e d f o r by t h e r m o r e g u l a t o r y e f f e c t s , and t h e r e f o r e f u t u r e r e s e a r c h s h o u l d f o c u s on p r e d a t i o n and s e x u a l i n t e r a c t i o n s . . More i n f o r m a t i o n on t h e h e r i t a b i l i t y of p h e n o t y p i c v a r i a t i o n , w i t h i n and between s e x e s , i s b a d l y needed. The summit of W h i t e Rocks Mt. n e a r Kelowna, B.C. c o u l d be u s e d t o measure t h e r a t e of a d a p t a t i o n t o a new e n v i r o n m e n t by e s t a b l i s h i n g a new c o l o n y u s i n g low e l e v a t i o n s t o c k . 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V a r i a t i o n i n s p e c t r a l r e f l e c t a n c e of a l p i n e and a r c t i c C o l i a s ( L e p i d o p t e r a : P i e r i d a e ) . Can. J . Z o o l . 5 6 ( 6 ) : 1447-1453. R o l a n d , J e n s . 1982. M e l a n i s m and d i e l a c t i v i t y of a l p i n e C o l i a s ( L e p i d o p t e r a : P i e r i d a e ) . O e c o l o g i a 53: 214-221. S c o t t , J.A. 1973a. P o p u l a t i o n b i o l o g y and a d u l t b e h a v i o r of t h e c i r c u m p o l a r b u t t e r f l y , P a r n a s s i u s phoebus F. ( P a p i l i o n i d a e ) . E n t . S c a n d . 4: 161-168. 1 52 S c o t t , J.A. 1973b. L i f e s p a n of b u t t e r f l i e s . J . Res. L e p i d , 1 2 ( 4 ) : 225-230. S e b e r , G.A.F. 1973. The e s t i m a t i o n o f a n i m a l a b u n d a n c e . C h a r l e s G r i f f i n and Co. L t d . , London. 506pp. S h a p i r o , A.M. 1970. The r o l e of s e x u a l b e h a v i o r i n d e n s i t y -r e l a t e d d i s p e r s a l o f P i e r i d b u t t e r f l i e s . Amer. N a t . 104: 367-372. Southwood, T.R.E. 1962. M i g r a t i o n o f t e r r e s t r i a l a r t h r o p o d s i n r e l a t i o n t o h a b i t a t . B i o l . Rev. 37: 171-214. S t i l e s , E.W. 1979. E v o l u t i o n of c o l o r p a t t e r n and p u b e s c e n c e c h a r a c t e r i s t i c s i n male bumblebees: A u t o m i m i c r y vs t h e r m o r e g u l a t i o n . E v o l u t i o n 33: 941-957. W a s s e r t h a l , L.T. 1975. The r o l e of b u t t e r f l y w i n gs i n r e g u l a t i o n of body t e m p e r a t u r e . J . I n s e c t P h y s i o l . 21: 1921-1930. Watt, W.B. 1968. A d a p t i v e s i g n i f i c a n c e o f p i g m e n t p o l y m o r p h i s m s i n C o l i a s b u t t e r f l i e s . I . V a r i a t i o n o f m e l a n i n p i g m e n t i n r e l a t i o n t o t h e r m o r e g u l a t i o n . E v o l u t i o n 22: 437-458. Watt, W.B. 1969. A d a p t i v e s i g n i f i c a n c e of p i g m e n t p o l y m o r p h i s m s i n C o l i a s b u t t e r f l i e s , I I . T h e r m o r e g u l a t i o n and p h o t o p e r i o d i c a l l y c o n t r o l l e d m e l a n i s m v a r i a t i o n ^ i n C o l i a s  e u r y t h e m e . P r o c . N a t . A c a d . S c i . 63: 767-774. Z a r , J.H. 1974. B i o s t a t i s t i c a l a n a l y s i s . P r e n t i c e - H a l l I n c . , E n g l e w o o d C l i f f s , N . J . 620pp. 153 APPENDIX 1. POPULATIONS AND SUBPOPULATIONS OF P. PHOEBUS E l e v a t i o n ' m i d p o i n t ' i s t h e m i d p o i n t of t h e p o p u l a t i o n , r a t h e r t h a n t h e m i d p o i n t of t h e e l e v a t i o n r a n g e ; l a t i t u d e and l o n g i t u d e a r e i n d e g r e e s ( d e c i m a l n o t a t i o n ) ; 'U.T.M. R e f . ' i s t h e U n i v e r s a l T r a n s v e r s e M e r c a t o r g r i d c o o r d i n a t e s (where known); n = t h e number of m a l e s (M) and f e m a l e s (F) i n t h e sample; t h e f l i g h t p e r i o d i s e s t i m a t e d as d e s c r i b e d i n C h a p t e r 1; t h e ' C a p t u r e D a t e s ' a r e g i v e n i n t h e form 'm.d' where m=month and d=day ( y e a r o m i t t e d ) . A l l o f t h e s p e c i m e n s c a p t u r e d i n ' a r e a s 4-6' on V e r n o n H i l l were r e l e a s e d w i t h o u t p h e n o t y p e s c o r i n g , r e s u l t i n g i n a sample s i z e o f z e r o . L o c a t i o n E l e v a t i o n L a t . U.T.M. n F l i g h t P e r i o d ( M i d p o i n t ) L o n g . R e f . M 0F C a p t u r e Date B.C. Mt. Apex, 2190-2247m 49. 36 Zone 11U 42 Aug. 1 -Sep.15 Pe n t i c t o n summi t (221Om) 1 19. 91 KE888713 4 07. 30; 08. 1 3 08. 21 ; 08. 23 B.C. Mt. Apex, 1980-2040m 49. 36 Zone 11U 35 J u l . 1 5-Aug. 31 P e n t i c t o n s .e. s i d e (201Om) 1 19. 90 KE894710 . 1 0 08. 13; 08. 21 B.C. Mt. Apex, 1920-1980m 49. 36 Zone 11U 4 J u l . 1 5-Aug. 31 P e n t i c t o n s . e . s i d e (1950m) 1 19. 89 KE899712 1 08. 13; 08. 21 B.C. Mt. Apex 1800-1860m 49. 36 Zone 11U 1 1 J u l . 1 5-Aug. 31 P e n t i c t o n s . e . s i d e (1830m) 1 19. 89 KE899709 3 07. 22; 08. 1 3 B.C. Mt. Apex, 1700-1800m 49. 35 Zone 11U 20 J u l . 1 -Aug. 1 5 Pent i c t o n s . e . s i d e (1750m) 1 19. 89 KE905702 6 08. 1 3 B.C. G r e e n Mt. 1740-l800m 49. 39 Zone 11U 9 J u l . 1 -Aug. 1 5 Pe n t i c t o n K l o h t e l t Ck (1770m) 1 19. 87 KE919745 1 08. 09 B.C. G r e e n Mt. 1800-I860m 49. 39 Zone 11U 5 J u l . 1 -Aug. 1 5 P e n t i c t o n K l o h t e l t Ck (1830m) 1 19. 87 KE917746 0 08. 09 B.C. G r e e n Mt. 1800-1890m 49. 39 Zone 11U 33 J u l . 1 5-Aug. 31 P e n t i c t o n K l o h t e l t Ck (1850m) 119. 87 KE928753 5 08. 1 4 B.C. G r e e n Mt. 1860-1950m 49. 39 Zone 11U 1 5 J u l . 1 5-Aug. 31 P e n t i c t o n K l o h t e l t Ck (1905m) 1 19. 87 KE915747 5 08. 09 B.C. G r e e n Mt. 1890-1980m 49. 40 Zone 11U 23 J u l . 1 5-Aug. 31 Pent i c t o n K l o h t e l t Ck (1950m) 1 19. 87 KE920753 6 08. 1 4 B.C. Gree n Mt. 1 800m 49. 39 Zone 11U 2 J u l . 1 -Aug. 1 5 Pent i c t o n K l o h t e l t Ck 1 1*9. 90 KE921741 1 08. 1 4 B.C. G r e e n Mt. 1980-2010m 49. 40 Zone 11U 4 Aug. 1 -Sep. 1 5 Pent i c t o n K l o h t e l t Ck (1995m) 1 19. 86 KE927757 2 08. 1 4 154 B.C. Gr e e n Mt. 1825-I890m P e n t i c t o n Keremeos Ck (1860m) B.C. G r e e n Mt. 1 525m Pent i c t o n O l d Apex Rd B.C. Gr e e n Mt. 1580-1650m Pent i c t o n O l d Apex Rd ( 1 615m) B.C. Gr e e n Mt. 1 220m Pent i c t o n O l d Apex Rd B.C. G r e e n Mt. 1800-1950m Pent i c t o n New Apex Rd (1860m) B.C. D i v i d e n d 1580-1675m Pent i c t o n Mt., e a s t (1630m) r i d g e B.C. D i v i d e n d 1675-1825m Pent i c t o n Mt., e a s t (1750m) r i d g e B.C. D i v i d e n d 1860-1920m Pent i c t o n Mt., e a s t (1890m) r i d g e B.C. D i v i d e n d 1920-2040m Pent i c t o n Mt.summi t (1980m) B.C. T r o u t Lk 790-820m Pent i c t o n Hwy 3A (805m) B.C. Red Mt. 2225-2315m Keremeos e a s t s i d e (2270m) B.C. Red Mt. 231 5m Keremeos s. r i d g e B.C. Q u i n i s c o e 2225-2315m Keremeos Mt., e a s t (2270m) r i d g e B.C. Q u i n i s c o e 2285m Keremeos M t . , s o u t h s i d e 49. 40 Zone 11U 20 J u l . 15- Aug .31 1 1 9 . 85 KE914742 8 08. 09; 08 .14 49. 38 Zone 11U 2 J u l . 1 -Aug . 15 1 19. 88 KE913728 1 06. 18; 07 .09 07. 29 49. 38 Zone 11U 86 J u l . 1 5-Aug .31 1 19. 88 KE913737 38 07. 19 07. 30- 08. 1 4 49. 38 Zone 11U 0 J u n . 1 -J u l . . 15 119. 88 KE913737 1 06. 18 49. 40 Zone 11U 81 J u l . 15- Aug .31 1 19. 90 KE895760 43 08. 08; 08 .09 08. 17; 08 . 18 49. 35 Zone 1 1U 1 4 J u l . 1-Aug .15 1 19. 85 KE932705 1 1 07. 29- 08. 05 49. 36 Zone 11U 38 J u l . 1 5-Aug .31 1 19. 86 KE926709 4 07. 29- 08. 04 49. 36 Zone 11U 6 J u l . 15-•Aug .31 119. 86 KE923709 . 1 07. 29 49. 35 Zone 11U 26 J u l . 1 5-•Aug ...31 1 19. 91 KE915716 4 08. 01-•08. 07 49. 34 Zone 11U 4 May 1 5-Jun .30 1 19. 72 KE028690 0 06. 1 1 49. 08 Zone 10U 6 Aug. 1 -Sep.. 15 120. 20 GK047401 6 08. 22 49. 08 Zone 10U 1 1 Aug. 1-Sep . 15 1 20. 20 GK044395 2 08. 22 49. 07 Zone 10U 1 4 Aug. 1 -Sep .15 1 20. 20 GK046389 8 08. 22 49. 07 Zone 10U 2 Aug. . 1 -Sep .15 120. 22 GK034385 2 08. 22 B.C. Keremeos C r a t e r M t . , s o u t h s i d e 2l30-2225m 49.08 Zone 10U (2190m) 120.18 GK129510 25 Aug. 1-Sep.15 5 08.17; 08.18 08.25; 08.31 155 B.C. 1675m 49. Zone 10U 1 J u l . 1-Aug. 15 H e d l e y 120. 0 08.04 B.C. Manning Pk 1525-1675m 49.07 Zone 10U 7 Jul.15-Aug.31 P r i n c e t o n G i b s o n P a s s (1580m) 120.89 FK540375 4 07.24; 07.27 B.C. Manning Pk 1370m 49.07 Zone 10U 0 J u l . 1-Aug.15 P r i n c e t o n G i b s o n P a s s 120.89 FK538366 1 07.24 B.C. Mt. B a l d y , 1825-I890m 49.16 Zone 11U 2 Jul.15-Aug.31 Osoyoos s. r i d g e (1860m) 119.26 LE352458 0 08.20 B.C. Mt. Kobau, 1800-I870m 49.11 Zone 11U 40 J u l . 15-Aug.. 31 Osoyoos summit (1840m) 119.67 LE044430 23 08.18 B.C. Mt. Kobau, 1825-I860m 49.12 Zone 1 1U 3 J u l . 15-Aug .'31 Osoyoos west r i d g e (1840m) 119.68 LE045437 1 08.18 B.C. Mt. Kobau, 1220-1430m 49.18 Zone 11U 17 J u n . 1- J u l . V 5 Cawston n.e. s i d e (1340m) 119.69 LE040512 8 07.21 B.C. Mt. Kobau, 1065m 49.23 Zone 11U 0 J u n . 1 5 - J u l . 3 0 O l i v e r n. s i d e 119.65 LE070562 1 07.21 B.C. S i l v e r • 1705-1860m. 50.37 Zone 11U 81 Jul.15-Aug.31 V e r n o n S t a r Mt., (1800m) 119.06 LF534812 28 07.26 summit 07.31-08.17 B.C. V e r n o n 1l00-1200m 50.26 Zone 11U 346 J u n . 1 - J u l . . 15 V e r n o n H i l l (1150m) 119.18 LF448694 147 06.01-07.18 s. s i d e " a r e a 2" B.C. V e r n o n 1050-1 150m 50.25 Zone 11U 0 J u n . 1-Jul.'1-5 V e r n o n H i l l (1100m) 119.18 LF443690 0 s. s i d e " a r e a 4" B.C. V e r n o n 1075-1125m 50.25 Zone 11U 0 J u n . 1 - J u l - 1 5 V e r n o n H i l l (1100m) 119.18 LF443688 0 s. s i d e " a r e a 5" B.C. V e r n o n 1075-1125m 50.25 Zone 11U 0 J u n . 1 - J u l . T 5 V e r n o n H i l l (1100m) 119.17 LF449687 0 s. s i d e " a r e a 6" B.C. V e r n o n 1200-1250m 50.25 Zone 11U 14 J u n . 1 - J u l . 1 5 V e r n o n H i l l (1225m) 119.17 LF451685 7 07.18 s. s i d e " a r e a 7" B.C. V e r n o n 1l90-1235m 50.25 Zone 11U 53 J u n . 1 - J u l . l 5 V e r n o n H i l l (1210m) 119.17 LF455686 2 06.15 s. s i d e 1 56 B.C. V e r n o n 820-915m 50. 26 Zone 11U 2 J u n . 1-J u l . 15 V e r n o n H i l l (870m) 119. 21 LF428700 0 06. 1 2 s.w. s i d e B.C. V e r n o n 1 1 60m 50. 27 Zone 11U 1 J u n . 1 -J u l . 15 V e r n o n H i l l 1 19. 19 LF428700 0 06. 05 s.w. s i d e B.C. Swan Lake 610-670m 50. 33 Zone 11U 3 J u n . 1 -J u l . 15 V e r n o n e a s t s i d e (640m) 1 19. 23 LF415771 0 06. 09 B.C. T e r r a c e 1860-1900m 50. 10 Zone 1 1U 41 J u l . 15- Sep. 1 V e r n o n Mt.summit (1890m) 119. 62 LF124525 7 08. 1 7 B.C. Q u i l p i r u k 945m 50. 07 Zone 11U 3 J u n . 1 5-J u l . 30 V e r n o n Ck.1.5 km 1 19. 52 LF197490 0 07. 04 s o u t h B.C. B r a s h Ck 610-760m 50. 57 Zone 11U 30 May 1 5-J u n . 30 E n d e r b y west s i d e (670m) 119. 07 LG545040 74 06. 04; 06. 1 3 06. 1 6 B.C. Hwy 97A 610-670m 50. 67 Zone 11U 40 May 1 5-J u n . 30 SalmonArm r o c k r i d g e (640m) 1 19. 21 LG442154 33 06. 12; 06. 1 6 B.C. 27 km e. 885-1000m 49. 84 Zone 11U 25 J u n . 1 -J u l . 1 5 Kelowna on Hwy 33 (945m) 1 19. 1 1 LF485238 4 0.6. 08; 06. 1 1 06. 13; 07. 17 B.C. 27 km e. 1160-1280m 49. 86 Zone 11U 38 J u n . 1 5-J u l . 30 Kelowna on Hwy 33 (1220m) 1 19. 1 0 LF490243 1 9 06. 13; • 07. 1 7 B.C. L a s t Mt. 960m 49. 85 Zone 11U 5 Jun . 1 -J u l . 1 5 Westbank r o a d 119. 71 LF049257 4 06. 30 B.C. L a s t Mt. 1 020m 49. 86 Zone 11U 4 J u n . 1 -J u l . 1 5 Westbank r o a d 119. 71 LF043266 0 06. 30 B.C. L a s t Mt. 1 220m 49. 89 Zone 11U 24 J u n . 1 5-J u l . 30 Westbank r o a d 1 19. 70 LF061297 25 06. 30 B.C. L a s t Mt. 1 21 5m 49. 89 Zone 11U 1 4 J u n . 1 5-J u l . 30 Westbank r o a d 119. 70 LF061294 0 06. 30 B.C. Lambly Ck 850m 50. 00 Zone 11U 1 8 J u n . 1 -J u l . 1 5 Westbank & T e r r a c e 119. 61 LF129415 1 3 06. 30 Ck. J e t . B.C. Bear Ck & 1 220m 50. 00 Zone 1 1U 2 J u l . 1 -Aug. 1 5 Westbank WhiteRock 1 19. 68 LF081424 0 06. 30 M a i n s J e t B.C. 9 km n. on 7 1 6m 49. 86 Zone 11U 0 May 1 5-J u n . 30 Westbank S m i t h Ck Rd 1 19. 67 LF082261 1 06. 30 1 57 B.C. M c D o u g a l l 91 5-975m 49. 91 Zone 11U 22 J u n . 1 5 - J u l . .30 Westbank C k 0 C a r r o t (945m) 1 19. 61 LF125313 6 06. 30 Ck Rd. B.C. L i t t l e 1 31 Om 49. 1 3 Zone 11U 1 1 J u n . 1 - J u l .15 Rock Ck F i s h L k. 119. 15 LE431440 1 5 07. 15; 07 .17 2 km w. B.C. L i t t l e 1 280m 49. 12 Zone 11U 7 J u n . 1 - J u l .15 Rock Ck F i s h Lk. .1 19. 1 2 LE458434 3 07. 05; 07 . 15 0.8 km e. B.C. 8 km e a s t 1 035m 50. 09 Zone 10U 4 J u n . 1 5 - J u l .30 M e r r i t t on Hwy 5 1 20. 68 FL656511 1 07. 05; 07 .08 B.C. B o t a n i e 760-915m 50. 30 Zone 10U 28 J u n . 1 5 - J u l .30 L y t t o n M t . s . s i d e (840m) 121. 57 FL017726 1 2 07. 05; 07 .07 07. 1 3 B.C. B o t a n i e 1370-1525m 50. 39 Zone 10U 37 J u l . 15-Aug .31 L y t t o n L k . n . s i d e (1430m) 121. 56 FL024830 30 07. 28; 08 . 16 B.C. B o t a n i e 1490-1555m 50. 42 Zone 10U 23 J u l . 15-Aug .31 L y t t o n L k . n . s i d e (1525m) 121. 58 FL008864 1 5 08. 16 B.C. M i s s i o n 1830-I900m 50. 77 Zone 10U 1 1 J u l . 15-Aug .31 L i l l o o e t R i d g e , (1865m) 1 22. 20 EM568238 8 08. 01; 08 .07 west end 08. 09 B.C. Mi s s i o n 2130-2285m 50. 73 Zone 10U 0 J u l . 30-Sep . 1 5 L i l l o o e t R i d g e (2200m) 1 22 . 07 EM655205 2 08. 01 Mt.McLean B.C. 29 km. 457m 58 . 04 Zone 9V 1 J u n . 1 5 - J u l .30 T e l e g r a p h Ck. n.w. 1 30. 88 UQ885342 0 07 . 1 7 Wash. H a r t s 1800m? 48. 72 Zone 10U 1 J u l . 15-Aug .31 OK.County P a s s 1 20. 68 0 08. 04 B.C. S k i h i s t 400m 50. 25 Zone 10U 0 J u n . 1 - J u l . 1 5 L y t t o n Campgrnd 121 . 50 FL063671 1 06. 10 B.C.,Rev- 38 km.s.e. 67 0m 50. 77 Zone 11U 0 J u l . 1-Aug. 15 e l s t o k e on A k o l - 117. 98 MG3025 1 07 .1 9 k o l e x Rd. B.C.,Rev- Mt . C a r t i e r 1 61 5m 50 . 78 Zone 11U 3 J u l . 30-Sep .15 v e l s t o k e west s i d e 118. 07 MG2440 0 08. 04 A l a s k a Nome 47 km.w. 45-75m 64.74 ( S i n u k R) (60m) 165.94 3 Aug. I-Sep.15 5 08.14 1 58 Wa./Port H u r r i c a n e 1 830m 48 5 Aug. 1-Sep. 1 5 A n g e l e s R i d g e 1 23 • 6 08. 29 B.C. Mt. Arrow- 1525-1675m 49 .26 Zone 10U 9 Aug. 1-Sep. 15 A l b e r n i s m i t h (1585m) 1 24 .59 CK843549 1 08. 12; 08. 13 B.C. Inkawatha 1800-I860m 49 .61 Zone 10U 2 J u l . 1 5-Aug. 31 Y a l e Lake (1830m) 121 .53 FK060965 3 08. 19 B.C. Chipmunk 1 770m 50 .59 Zone 10U 10 J u l . 1 5-Aug. 31 Pemberton Mt. 123 .00 DM994042 7 08. 03; 08. 07 B.C. D u f f y Lk 2200-2255m 50 .42 Zone 10U 5 Aug. 1 -Sep. 1 5 L i l l o o e t n. s i d e (2225m) 1 22 .37 EL452845 5 08. 10 B . C . G o l d - E l d o r a d o 2135-2225m 50 .83 Zone 10U 1 0 J u l . 15- Aug. 31 b r i d g e Mt., s . o f (2165m) 122 .85 EM108473 9 08. 09 B . C . , B i g B i g Dog 2040m 51 .03 Zone 10U 5 J u l . 1 5-Aug. '31 Bar C r e e k Mt. 1 22 .53 EM328538 6 08. 02 B . C . , B i g Buck Mt. 1 860m 51 . 1 5 Zone 10U 5 J u l . 1 5-Aug. 31 Bar C r e e k 1 22 .63 EM261663 5 08. 01 B . C . , B i g P o i s o n 2195-2255m 51 . 1 5 Zone 10U 3 J u l . 1 5-Aug. .31 Bar C r e e k Mt. (2225m) 1 22 .60 EM280662 5 08. 01 B.C. , B i g P o i s o n 2135-2195m 51 . 1 5 Zone 10U 9 J u l . 1 5-Aug.. 31 Bar C r e e k Mt. (2165m) 1 22 .60 EM276667 9 08. 01 B . C . , B i g P o i son 1675-1705m 51 . 12 Zone 10U 5 J u l . 1 -Aug. T5 Bar C r e e k Mt. (1690m) 1 22 .58 EM295632 3 07. 28; 08.. 01 B . C . , B i g 9 - M i l e 1 525m 51 .08 Zone 10U 3 J u l . 1 -Aug. 15 Bar C r e e k R i d g e 1 22 .52 EM338587 6 07. 28 A l b e r t a Nanton 2040-2135m 50 .19 Zone 11U 7 J u l . 1 -Aug.. 15 Kananas- Rd.,5 km. (2090m) 1 1 4 .44 PF825636 3 07. 19; 07.. 20 k i s Rd. e a s t on A l b e r t a C a t C r e e k 1615-1675m 50 .40 Zone 11U 0 J u l . 1 -Aug. 15 Kananas- (1645m) 1 1 4 .68 PF630868 4 07. 18 k i s Rd. A l b e r t a P l a t e a u x 21 35m 50 .23 Zone 11U 2 J u l . 1 -Aug. 15 Kananas- Mt. 1 1 4 . 50 PF783668 3 07. 23; 07.. 21 k i s Rd. A l b e r t a R e g a l Ck 1525m? 49 • 2 J u l . 1 -•Aug. 1 5 Rananas- 1 1 4 • 2 07. 20 Montana Snow Bowl 1525m? 47 12 J u l . 1 -•Aug. '1 5 M i s s o u l a S k i a r e a 1 1 4 1 1 07. 21 159 Montana C a r b o n C o . B e a r t o o t h P a s s 31 40m 45. 1 09. 00 42 Wyoming Togwotee Pa s s 2865m 43. 110. 75 03 Montana B i t t e r -r o o t Mts Woods Ck. Rd.,Mi.5 2200m 45. 114. 55 47 Montana H o r s e Ck P a s s B e a v e r Ck . 1700-2000m (1850m) 45. 114. 52 33 IdahOo Montana B o r d e r Between Wood Ck & H o r s e Ck 2200-2400m (2300m) P a s s e s 45. 114. 52 42 Montana Lower Red Rock Lk 2045m 44. 111. 63 87 Wyoming G r a n i t e P a s s 1 980m 42. 1 07. 48' 1 7 Montana Babb C h i e f Mt. 1 770m 48. 113. 93 53 Wyoming A l b a n y C o P o l e Mtn. 2590m 42. 1 05. 58 87 C o l o r a d o Huer f a n o C o u n t y W.Span i sh Peak A p i s h i p a 3500m Pas s 37. 1 05. 28 00 C o l o r a d o G o l d e n L o v e l a n d P a s s Hwy 3660m 6 39. 1 05. 63 88 Yukon A t l i n Rd. S n a f u Ck Campgrnd 760-790m (775m) 60. 1 33. 1 3 80 U t a h , S a l t Lake C i t y Red Rock H i l l 2925m 40. 111. B . C . R i s k e C r e e k B a l d Mt. 1 220m 51 . 1 22. 88 67 B.C. Waneta 4 km. e a s t 67 0m 49. 117. 02 57 1 Jul.15-Aug.31 0 08.01 1 Jul.15-Aug.31 0 08.04 1 J u l . 1-Aug.15 1 07.30 1 J u l . I - A u g . 1 5 0 07.31 1 J u l . 1 5 - A u g . 3 0 2 07.30 4 J u l . 1 5 - A u g . 3 0 4 08.01 2 J u n . 1 5 - J u l . 3 0 0 07.06 1 J u l . 1-Aug.15 0 07.20 1 J u n . 1 5 - J u l . 3 0 1 06.27 1 J u l . 1-Aug.15 1 07.09 4 Aug. 1-Sept.1 1 08.08 44 J u n . 1 5 - J u l . 3 0 20 06.28; 07.22 1 J u l . 1 5 - A u g . 3 0 2 07.25 2 J u n . 1 5 - J u l , 3 0 0 06.29 1 May 15-Jun.30 0 05.28 

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