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Terminal weevils of lodgepole pine and their parasitoid complex in British Columbia Kovacs, Ervin 1988

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TERMINAL WEEVILS OF LODGEPOLE PINE AND THEIR PARASITOID COMPLEX IN BRITISH COLUMBIA by ERVIN KOVACS B . S c . ( A g r . ) , U n i v e r s i t y of K i e v , 1983 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 F o r e s t r y , F o r e s t Entomology) We a c c e p t t h i s t h e s i s as c o n f o r m i n g t o the r e q u i r e d s t a n d a r d (3.0 u n i t s ) THE UNIVERSITY OF BRITISH COLUMBIA A p r i l 1988 © E r v i n Kovacs, 1988 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, 1 agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of Forest Sc.lfmf.eB The University of British Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date A p r i l 21 r1988  DE-6(3/8-n i i A b s t r a c t A study has been conducted w i t h the o b j e c t i v e s of (1) i d e n t i f y i n g w e e v i l s and t h e i r p a r a s i t o i d s emerging from i n f e s t e d l o d g e p o l e p i n e l e a d e r s , (2) d e t e r m i n i n g emergence p a t t e r n s of h o s t s and t h e i r p a r a s i t o i d s , and (3) o b t a i n i n g f u r t h e r i n f o r m a t i o n on the b i o l o g i e s of the t e r m i n a l w e e v i l s and t h e i r n a t u r a l enemies i n B r i t i s h C olumbia. The major e x p e r i m e n t s and b i o l o g i c a l o b s e r v a t i o n s were c a r r i e d out i n young spaced l o d g e p o l e p i n e , ( P i n u s c o n t o r t a D o u g l . v a r . l a t i f o l i a Engelm.), s t a n d s a t E l l i s c r e e k , near P e n t i c t o n , B.C. A t o t a l of 1046 i n f e s t e d l e a d e r s were c o l l e c t e d . O n e - t h i r d of the t e r m i n a l s were d i s s e c t e d and the numbers of w e e v i l s and p a r a s i t o i d s a t d e v e l o p m e n t a l s t a g e s were r e c o r d e d . The remainder of the l e a d e r s were s e t up f o r i n d i v i d u a l r e a r i n g . O b s e r v a t i o n s were a l s o made on the f e e d i n g and o v i p o s i t i o n a l b e h a v i o r of the w e e v i l s . F e e d i n g h a b i t s of t h e p a r a s i t o i d s were a l s o s t u d i e d . D i s s e c t i o n s showed t h a t a few a d u l t w e e v i l s emerge i n the f a l l of the y e a r of a t t a c k . The m a j o r i t y of a d u l t s o v e r w i n t e r as l a r v a e but p u p a t i o n a l s o may occur p r i o r t o w i n t e r . In a d d i t i o n , d i s s e c t i o n s i n d i c a t e d t h a t p a r a s i t i s m p l a y s an i m p o r t a n t r o l e i n l a r v a l m o r t a l i t y of w e e v i l s . i i i W e e v i l s which emerged i n the l a b o r a t o r y were i d e n t i f i e d as b e i n g of the f o l l o w i n g s p e c i e s : P i s s o d e s t e r m i n a l i s Hopping, M a g d a l i s g e n t i l i s LeC. and C y l i n d r o c o p t u r u s sp. (COLEOPTERA: C u r c u l i o n i d a e ) . M. g e n t i l i s i s the f i r s t w e e v i l s p e c i e s t o emerge, i n l a t e May. T h i s emergence i s f o l l o w e d by t h a t of P. t e r m i n a l i s from e a r l y June t h r o u g h m i d - J u l y , w h i l e C y l i n d r o c o p t u r u s sp. emerges from e a r l y June through m i d - J u l y . P. t e r m i n a l i s a t t a c k s the c u r r e n t y e a r ' s l e a d e r s , whereas a d u l t M. g e n t i l i s and C y l i n d r o c o p t u r u s sp. f e e d on f o l i a g e . A l l t h r e e w e e v i l s p e c i e s u t i l i z e l o d g e p o l e p i n e t e r m i n a l s h o o t s f o r b r e e d i n g . L a r v a l f e e d i n g under the bark almost always r e s u l t s i n the d e a t h of t h e t e r m i n a l . The t e r m i n a l w e e v i l s have a complex of n a t u r a l enemies i n B r i t i s h C olumbia. P a r a s i t o i d s b e l o n g t o s i x f a m i l i e s of the o r d e r Hymenoptera. The p t e r o m a l i d R h o p a l i c h u s  p u l c h r i p e n n i s C r a w f o r d i s the most w i d e l y d i s t r i b u t e d p a r a s i t o i d s p e c i e s i n the p r o v i n c e . Two s p e c i e s of Eurytoma (Eurytomidae) ranked second i n abundance. Emergence p a t t e r n s of a d u l t p a r a s i t o i d s a r e c l o s e l y s y n c h r o n i z e d w i t h t h a t of t h e i r h o s t s . i v P a r a s i t o i d s were obser v e d f e e d i n g on p o l l e n of f l o w e r i n g weeds i n the f i e l d . T h i s o b s e r v a t i o n s u g g e s t s t h a t n a t u r a l p a r a s i t o i d p o p u l a t i o n s c o u l d be enhanced by c u l t i v a t i n g l u p i n , L u p i n u s sp., i n l o d g e p o l e p i n e s t a n d s . I t was c o n c l u d e d t h a t e v e r y e f f o r t s h o u l d be made t o m i n i m i z e w e e v i l numbers i n o r d e r t o p r e v e n t f o r m a t i o n of c r o o k s , f o r k s and st a g - h e a d s . E a r l y emergence of M. g e n t i l i s s u g g e s t s t h a t l e a d e r c l i p p i n g p r o j e c t s s h o u l d be c a r r i e d out by e a r l y s p r i n g . F u r t h e r r e s e a r c h i s recommended t o ensure c o r r e c t a s s o c i a t i o n between p a r a s i t o i d s and hos t w e e v i l s p e c i e s and t o d e v e l o p or e s t a b l i s h methods f o r p r e s e r v a t i o n of p a r a s i t o i d s f o r c l i p p e d l e a d e r s f o r r e l e a s e i n the f o r e s t . V TABLE OF CONTENTS PAGE TITLE PAGE i ABSTRACT i i TABLE OF CONTENTS V LIST OF TABLES v i LIST OF FIGURES v i i i LIST OF APPENDIX x i ACKNOWLEDGEMENT x i i 1. INTRODUCTION 1.1 C u r r e n t Knowledge 1 1.1.1 The I n s e c t s 1 1.1.2 The Host 1 2 1.1.3 B i o l o g y of P i s s o d e s t e r m i n a l i s 17 1.1.4. B i o l o g y of M a q d a l i s g e n t i l i s 29 1.1.5. B i o l o g y of C y l i n d r o c o p t u r u s sp. 30 1.2 O b j e c t i v e s of the Study 34 2. METHODS 2.1 L o c a t i o n of the Study 35 2.2 D i s s e c t i o n of I n f e s t e d L e a d e r s 35 2.2.1. Sampling Techniques 39 2.3 I n c u b a t i o n of I n f e s t e d L e a d e r s 40 2.4 D i r e c t O b s e r v a t i o n s and Measurements 43 2.5 Data A n a l y s i s 45 3. RESULTS AND DISCUSSION 3.1 Summer C o l l e c t i o n s i n 1986 46 3.2 F e b r u a r y Leader C o l l e c t i o n i n 1987 46 3.3 Summer A c t i v i t i e s i n P e n t i c t o n i n 1987 54 3.3.1 D i s s e c t i o n of A t t a c k e d L e a d e r s 54 3.3.2 I n d i v i d u a l I n c u b a t i o n of I n f e s t e d L e a d e r s 62 3.3.3 O b s e r v a t i o n s on the B i o l o g i e s of P . t e r m i n a l i s 66 3.3.3.1. L o n g e v i t y and F e c u n d i t y of P i s s o d e s  t e r m i n a l i s 66 3.3.3.2. D u r a t i o n of Egg and P u p a l Stages of 69 3.3 . 4 . O b s e r v a t i o n s on the B i o l o g y of M. g e n t i l i s and C y l i n d r o c o p t u r u s sp. 71 3.3.5 P a r a s i t o i d s A s s o c i a t e d w i t h the W e e v i l s 81 3.3.6 A d u l t P a r a s i t o i d Food P l a n t 84 3.3.7 W e e v i l A t t a c k and A t t a c k H i s t o r y Survey 87 3.4 Management I m p l i c a t i o n s 91 3.5 Other S t r a t e g i e s f o r W e e v i l C o n t r o l 92 4. GENERAL DISCUSSION AND CONCLUSIONS 93 5. RECOMMENDATIONS 94 BIBLIOGRAPHY 95 APPENDIX 100 v i LIST OF TABLES TABLE PAGE I A r e a i n f e s t e d and l o s s e s caused by the Mountain P i n e B e e t l e , Dendroctonus ponderosae Hopkins i n f e s t a t i o n s i n B r i t i s h Columbia i n 1983-1987 2 I I T o t a l number of t r e e s of major s p e c i e s p l a n t e d 15 on Crown Land i n B r i t i s h Columbia i n 1984-1986 I I I I n s e c t s r e c o r d e d as b e i n g a s s o c i a t e d w i t h l e a d e r s of young l o d g e p o l e p i n e i n western N o r t h America 18 IV P a r a s i t i c Hymenoptera of P i s s o d e s t e r m i n a l i s 28 V P a r a s i t i c Hymenoptera of C y l i n d r o c o p t u r u s spp. 33 VI Summary of t e r m i n a l w e e v i l s and p a r a s i t o i d s r e a r e d from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d t h r o u g h o u t B r i t i s h Columbia i n June,1986 47 V I I Summary of the i n s e c t l i f e s t a g e s found i n i n f e s t e d l e a d e r s of l o d g e p o l e p i n e c o l l e c t e d near W i l l i a m s Lake i n February,1987 50 V I I I Numbers of t e r m i n a l w e e v i l s and p a r a s i t o i d s r e a r e d from i n f e s t e d l e a d e r s c o l l e c t e d a t R i s k e Creek, near W i l l i a m s Lake between F e b r u a r y 25 and A p r i l 2,1987 52 IX Summary t a b l e of l e a d e r s c o l l e c t e d f o r d i s s e c t i o n and i n c u b a t i o n i n s o u t h e r n B r i t i s h Columbia i n May,1987 55 X I n s e c t l i f e s t a g e s found d u r i n g d i s s e c t i o n of damaged l o d g e p o l e p i n e l e a d e r s c o l l e c t e d i n s o u t h e r n B r i t i s h Columbia i n May,1987 56 XI Summary t a b l e of the r e s u l t s of weekly d i s s e c t i o n of l o d g e p o l e p i n e t e r m i n a l s a t t a c k e d d u r i n g 1987 from E l l i s Creek 58 X I I Summary t a b l e of i n s e c t s r e a r e d from w e e v i l i n f e s t e d l o d g e p o l e p i n e t e r m i n a l s c o l l e c t e d a t E l l i s Creek and a l o n g the B i g White Road i n May,1987 63 X I I I L o n g e v i t y and o v i p o s i t i o n c h a r a c t e r i s t i c s of ten female P i s s o d e s t e r m i n a l i s r e a r e d a t 20+2°C a t UBC 83 TABLE XIV Hymenopterans found a s s o c i a t e d w i t h t e r m i n a l w e e v i l s i n B r i t i s h C o lumbia, 1986-1987 XV R e s u l t s of the w e e v i l a t t a c k i n c i d e n c e and a t t a c k h i s t o r y survey c onducted on a 33ha young spaced l o d g e p o l e p i n e s t a n d a t E l l i s Creek i n J u l y , 1 9 8 7 (n=!088) v i i i LIST OF FIGURES FIGURE PAGE Crook i n the main stem of l o d g e p o l e p i n e 4 2 Lodgepole p i n e e x h i b i t i n g f o r k i n the stem 5 3 R e p l a c i n g the dead t e r m i n a l by t h r e e or more l a t e r a l s r e s u l t s i n f o r m a t i o n of a stag-head 6 4 G e n e r a l i z e d r e l a t i o n s h i p of growth r a t e t o t e m p e r a t u r e 11 5 D i s t r i b u t i o n of P. t e r m i n a l i s and i t s h o s t s i n N o r t h America 1 3 6 A d u l t P i s s o d e s t e r m i n a l i s on e l o n g a t i n g l o d g e p o l e p i n e l e a d e r 21 7 D o r s a l view of the l a s t abdominal segment of the l o d g e p o l e t e r m i n a l w e e v i l 22 8 G e n e r a l i z e d l i f e c y c l e of P. t e r m i n a l i s on l o d g e p o l e p i n e 24 9 Second i n s t a r P. t e r m i n a l i s l a r v a m i n i n g beneath the e p i d e r m i s of the c u r r e n t y e a r ' s t e r m i n a l shoot 26 10 L a r v a (A) and pupa (B) of genus C y l i n d r o c o p t u r u s 31 11 Map showing the l o c a t i o n s of c o l l e c t i o n s of i n f e s t e d l e a d e r s i n s o u t h - c e n t r a l B r i t i s h Columbia i n the summer of 1986 36 12 L o c a t i o n s of the c o l l e c t i o n p l o t s i n the s o u t h e r n i n t e r i o r of B r i t i s h Columbia i n the summer of 1987 37 13 L o c a t i o n s of the s t u d y and s u r v e y p l o t s a t E l l i s Creek, near P e n t i c t o n , B.C. 38 14 Mass r e a r i n g of i n f e s t e d l o d g e p o l e p i n e l e a d e r s i n m o d i f i e d s e e d l i n g boxes. Note emergence j a r s e t i n t o the s i d e of the box a t t o p l e f t 41 15 I n d i v i d u a l r e a r i n g of i n f e s t e d l o d g e p o l e p i n e l e a d e r s i n m a i l i n g t u b e s . Note the v i a l e n t e r e d i n t o the lower h a l f of the cap 42 i x FIGURE PAGE 16 Two t y p e s of symptoms of i n f e s t e d l o d g e p o l e p i n e l e a d e r s by t e r m i n a l w e e v i l s . The t e r m i n a l s t i l l has i t s a p i c a l dominance but i t s n e e d l e s have a l r e a d y changed c o l o r ( A ) . The t e r m i n a l i s s t i l l green but has l o s t i t s a p i c a l dominance and l a t e r a l s have caught up w i t h the i t (B) 48 17 Emergence of l e a d e r w e e v i l s and t h e i r p a r a s i t o i d s from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d a t R i s k e Creek, near W i l l i a m s Lake, B.C. i n February,1987 53 18 Frequency d i s t r i b u t i o n of h e a d c a p s u l e w i d t h s of dead w e e v i l l a r v a e r e s u l t e d from weekly d i s s e c t i o n s of i n f e s t e d t e r m i n a l shoots 60 19 Frequency d i s t r i b u t i o n o f h e a d c a p s u l e w i d t h s of l i v e w e e v i l l a r v a e r e s u l t e d from weekly d i s s e c t i o n s of i n f e s t e d t e r m i n a l s h o o t s 61 20 Temporal emergence of M a g d a l i s g e n t i l i s and C y l i n d r o c o p t u r u s sp. from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d i n the s o u t h e r n i n t e r i o r of B r i t i s h Columbia i n May,1987. R e a r i n g s t a r t e d on May 7,1987 64 21 Temporal emergence of P i s s o d e s t e r m i n a l i s and p a r a s i t o i d s from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d i n t h e s o u t h e r n i n t e r i o r of B r i t i s h Columbia i n May,1987. R e a r i n g s t a r t e d on May 7,1987 65 22 Schematic diagram of t e r m i n a l w e e v i l emergence and o v i p o s i t i o n i n r e l a t i o n t o l e a d e r e l o n g a t i o n of P i n u s c o n t o r t a 67 23 S u r v i v a l s h i p and f e c u n d i t y of t e n P i s s o d e s t e r m i n a l i s females 70 24 M a q d a l i s g e n t i l i s a d u l t on e l o n g a t i n g l o d g e p o l e p i n e t e r m i n a l shoot 72 25 One h e a l t h y l o d g e p o l e p i n e n e e d l e and 11 n e e d l e s a f f e c t e d by the f e e d i n g of a d u l t M. g e n t i l i s . The a p i c a l p a r t of the n e e d l e s have d e s s i c a t e d , d i s c o l o r e d and c u r l e d up a l o n g the l o n g i t u d i n a l a x i s . A p i c a l p o r t i o n of the l a s t t h r e e n e e d l e s had been broken e i t h e r by wind or r a i n 73 26 O v i p o s i t o n s i t e made by M. g e n t i l i s female i n the c u r r e n t y e a r ' s t e r m i n a l s h o o t . O v i p o s i t i o n was a t the v e r y base of the n e edle 74 PAGE 27 O v i p o s i t i o n s i t e made by P. t e r m i n a l i s female i n t o the e l o n g a t i n g t e r m i n a l 28 C y l i n d r o c o p t u r u s sp. a d u l t 29 F e e d i n g p u n c t u r e s on l o d g e p o l e p i n e n e e d l e s caused by C y l i n d r o c o p t u r u s sp. a d u l t s 30 Lodgepole p i n e t e r m i n a l damaged by M a g d a l i s  q e n t i l i s l a r v a e 31 Lodgepole p i n e t e r m i n a l damaged by l a r v a e of C y l i n d r o c o p t u r u s sp. 32 F i r e w e e d , E p i l o b i u m a n q u s t i f o l i u m L. i n young l o d g e p o l e p i n e s t a n d 33 L u p i n , L u p i n u s sp. growing i n young l o d g e p o l e p i n e s t a n d 34 1985 and 30-year monthly minimum and maximum average t e m p e r a t u r e s a t the P e n t i c t o n a i r p o r t 35 1986 and 30-year monthly minimum and maximum average t e m p e r a t u r e s a t the P e n t i c t o n a i r p o r t LIST OF APPENDIX APPENDIX I Form used f o r r e c o r d i n g i n s e c t l i f e s t a g e s found d u r i n g d i s s e c t i o n I I Form used f o r r e c o r d i n g emerged specimens from r e a r i n g boxes I I I P r o j e c t e d f i e l d emergence of w e e v i l s c a l c u l a t e d w i t h t h r e e methods of degree-day c a l c u l a t i o n f o r d i f f e r e n t d e v e l o p m e n t a l minimum t e m p e r a t u r e s (TMIN) when the maximum d e v e l o p m e n t a l t e m p e r a t u r e (TMAX) was unknown and s e t a t 30°C IV Weekly measurements of 25 e l o n g a t i n g l o d g e p o l e p i n e l e a d e r s ACKNOWLEDGEMENT I would l i k e t o acknowledge the h e l p and c r i t i c i s m r e c e i v e d from my s u p e r v i s o r , Dr. J.A. McLean, committee members, D r s . M.B. Isman, and J.W.E. H a r r i s ; and Dr. M.A. Hulme d u r i n g the r e s e a r c h . I a l s o would l i k e t o thank the F o r e s t R e s e a r c h Development Agreement f o r p r o v i d i n g the g r a n t f o r t h i s r e s e a r c h . I would l i k e t o e x p r e s s my a p p r e c i a t i o n t o the s c i e n t i s t s of the B i o s y s t e m a t i c s R e search I n s t i t u t e i n Ottawa f o r the i d e n t i f i c a t i o n of the specimens. I d e d i c a t e t h i s t h e s i s t o my p a r e n t s who encouraged me d u r i n g the s t u d y ; t o my Aunt and U n c l e , Mrs. and Mr. M. Kovacs who p r o v i d e d f i n a n c i a l s u p port f o r my expenses d u r i n g my U n i v e r s i t y y e a r s . I a l s o would l i k e t o thank my f r i e n d , A l d o I n t r i e r i f o r h i s encouragement I r e c e i v e d w h i l e a t U n i v e r s i t y . 1 1. INTRODUCTION 1.1 C u r r e n t knowledge 1.1.1 The i n s e c t s Lodgepole p i n e , P i n u s c o n t o r t a D o u g l . v a r . l a t i f o l i a Engelm. has become an i m p o r t a n t component of the a n n u a l a l l o w a b l e c u t i n B r i t i s h Columbia and i n 1986 i t a c c o u n t e d f o r 23% of a l l t i m b e r h a r v e s t e d (MOF 1986). Much of the t i m b e r was h a r v e s t e d as a r e s u l t of mountain p i n e b e e t l e , Dendroctonus ponderosae Hopk. (COLEOPTERA: S c o l y t i d a e ) , i n f e s t a t i o n s . Mature l o d g e p o l e p i n e s t a n d s have s u f f e r e d enormous l o s s e s from a t t a c k s by t h i s i n s e c t p e s t throughout the p r o v i n c e f o r the p a s t f i v e y e a r s ( T a b l e I ) . In o u t b r e a k s i t u a t i o n s s a n i t a t i o n l o g g i n g i s recommended ( S a f r a n y i k e t a l . 1974). The i n t e n s i v e s a l v a g e o p e r a t i o n s have r e s u l t e d i n l a r g e c l e a r c u t s which a r e now b e i n g r e g e n e r a t e d . Young s t a n d s of l o d g e p o l e p i n e a r e v u l n e r a b l e t o a t t a c k by many i n s e c t p e s t s (Amman and S a f r a n y i k 1984). Among the numerous i n s e c t s a t t a c k i n g open-grown r e g e n e r a t i o n , t w i g w e e v i l s , b e l o n g i n g t o the f a m i l y C u r c u l i o n i d a e ( o r d e r C o l e o p t e r a ) , o f t e n cause s e r i o u s damage t o the t e r m i n a l s of young c o n i f e r o u s t r e e s (Keen 1952). In B r i t i s h Columbia, one of the most i m p o r t a n t t w i g w e e v i l s i s the l o d g e p o l e t e r m i n a l w e e v i l , P i s s o d e s t e r m i n a l i s Hopping but o t h e r w e e v i l s such as M a q d a l i s g e n t i l i s LeConte ( F u r n i s s and C a r o l i n 1977; Wood et a_l. 1987) and C y l i n d r o c o p t u r u s spp. 2 T a b l e 1: Area i n f e s t e d and l o s s e s caused by Mountain p i n e b e e t l e , Dendroctonus ponderosae Hopkins i n f e s t a t i o n s i n B r i t i s h Columbia i n 1983-1987. F o r e s t 1983-1987  Region a r e a # of t r e e s volume k i l l e d i n f e s t e d k i l l e d ( m i l l i o n m 3) (ha) ( m i l l i o n s ) C a r i b o o 1,063,000 72. 1 21.4 Kamloops 214,410 21.9 11.8 N e l s o n 107,300 7.5 2.7 P r i n c e Rupert 73,400 4.2 3.6 P r i n c e George 10,225 7.4 0.25 T o t a l 1,468,335 113.1 39.75 1 1983 d a t a are not a v a i l a b l e 3 ( F u r n i s s and C a r o l i n 1977) are a l s o known as damaging a g e n t s . T e r m i n a l s a t t a c k e d by P. t e r m i n a l i s change c o l o r from a h e a l t h y green t o a d u l l brown. D y i n g and dead l e a d e r s a r e c l e a r l y r e c o g n i z a b l e by e a r l y August ( D r o u i n e_t a l . 1963). The l o s s of the t e r m i n a l l e a d e r r e s u l t s i n c o m p e t i t i o n between the l a t e r a l s i n the w h o r l below the dead l e a d e r . I f o n l y a s i n g l e l a t e r a l assumes a p i c a l dominance the .stem d e v e l o p s a permanent c r o o k ( F i g . 1 ) . Sometimes two l a t e r a l s compete f o r dominance which r e s u l t s i n a f o r k i n the stem ( F i g . 2 ) . I f w e e v i l a t t a c k r e o c c u r s on the same t r e e f o r t h r e e or more c o n s e c u t i v e y e a r s or as a r e s u l t of a s i n g l e y e a r ' s a t t a c k , t h r e e or more l a t e r a l s can assume a p i c a l dominance, and the t r e e becomes m u l t i - l e a d e r e d or stag-headed ( F i g . 3 ) . Maher (1982) e s t i m a t e d t h a t e v e r y time t h e t e r m i n a l i s a t t a c k e d by P. t e r m i n a l i s and the dead l e a d e r i s r e p l a c e d by one of the l a t e r a l s , the t r e e s u f f e r s a 42% h e i g h t growth l o s s . Salman (1935) s t a t e d t h a t the impact of P i s s o d e s  t e r m i n a l i s on l o d g e p o l e p i n e i s s i m i l a r , i n some r e s p e c t s , t o t h a t caused by the S i t k a s pruce w e e v i l , P i s s o d e s s t r o b i Peck. ( C o l e o p t e r a : C u r c u l i o n i d a e ) i n e a s t e r n w h i t e p i n e , P i n u s s t r o b u s L. Marty (1959) i n d i c a t e d two t y p e s of l o s s e s due t o P. s t r o b i a t t a c k . The f i r s t t y p e of l o s s i s r e d u c t i o n i n r e c o v e r a b l e volume which i s due t o f o r m a t i o n of F i g u r e 1 Crook i n the main stem of l o d g e p o l e p i n e . 5 Figure 2 Lodgepole pine exhibiting fork in the stem. F i g u r e 3 R e p l a c i n g the dead t e r m i n a l by t h r e e or more l a t e r a l s r e s u l t s i n f o r m a t i o n of a s t a g - h e a d . 7 c r o o k s , f o r k s and sta g - h e a d s . The second type of l o s s i s lumber degrade i n the r e m a i n i n g volume. M. g e n t i l i s c a uses damage from C a l i f o r n i a t o B r i t i s h Columbia and i n Montana (Keen 1952; F u r n i s s and C a r o l i n 1977). Damage caused by M. g e n t i l i s i s not r e s t r i c t e d t o any p a r t i c u l a r p o r t i o n of the crown ( F e l l i n and Schmidt 1966) and F e l l i n (1973) does not c o n s i d e r i t t o be s e r i o u s . A d u l t w e e v i l s p u n c t u r e the n e e d l e s w i t h t h e i r m a n d i b l e s and f e e d on the p a l i s a d e and o t h e r t i s s u e s ( F e l l i n 1973). Sometimes the f e e d i n g i s so e x t e n s i v e t h a t a h o l e i s p u n c t u r e d t h r o u g h the n e e d l e . D i s t a l p o r t i o n s of the n e e d l e s , i n r e l a t i o n t o the f e e d i n g s i t e , d e s s i c a t e and d i s c o l o r ( F e l l i n and Schmidt 1966), and a r e broken o f f by wind, r a i n or snow (Plumb 1950; F e l l i n 1973). Such i n j u r y resembles t h a t caused by the f e e d i n g of l a r v a e of L e p i d o p t e r a or T e n t h r i d i d s a w f l i e s (Plumb 1950). F e l l i n (1973) s t a t e d t h a t d e f o l i a t i o n was t h e o n l y t y p e of damage he had ob s e r v e d and " t h e r e was no i n d i c a t i o n t h a t a d u l t s o v i p o s i t nor t h a t l a r v a e f e e d i n or on the s h o o t s or any o t h e r p o r t i o n s of the s t a n d i n g t r e e " . A p p r o x i m a t e l y 35 w e e v i l s p e c i e s d e s c r i b e d from N o r t h America belong t o genus C y l i n d r o c o p t u r u s but o n l y a few s p e c i e s have been r e p o r t e d t o be of economic importance (Eaton 1942). S e v e r a l s p e c i e s i n t h i s genus a t t a c k the t w i g s and b o l e s of young c o n i f e r o u s t r e e s , such as p i n e , P i n u s spp., l a r c h , L a r i x spp., and t r u e f i r , A b i e s spp. i n 8 N o r t h America (Buchanan 1940). W e e v i l s cause damage i n both the a d u l t and l a r v a l s t a g e s (Eaton 1942). E x c e l l e n t a c c o u n t s of the b i o l o g y and f e e d i n g h a b i t s of t h e s e w e e v i l s are g i v e n by Eaton (1942) and Buchanan (1940). A d u l t w e e v i l s p u n c t u r e the e p i d e r m i s of the needle and eat the m e s o p h y l l t i s s u e . In e x c e p t i o n a l c a s e s , d e f o l i a t i o n i s so e x t e n s i v e t h a t i t l e a d s t o the d e a t h of the whole t r e e . A d u l t s a l s o feed on the t i s s u e s of the c u r r e n t y e a r ' s growth. Eaton (1942) p o i n t e d out t h a t l a r v a l f e e d i n g i n the stem and t w i g s would be more s e r i o u s than f e e d i n g by a d u l t s on the f o l i a g e , because the former was u s u a l l y f o l l o w e d by the d e a t h of the t r e e . A l l t he p r e v i o u s l y mentioned w e e v i l s p e c i e s p r e f e r young, open-grown t r e e s . The impact of the s e w e e v i l s on young s t a n d s urges us t o make e v e r y e f f o r t t o reduce w e e v i l p o p u l a t i o n s . The o n l y c o n t r o l method c u r r e n t l y used a g a i n s t the l o d g e p o l e t e r m i n a l w e e v i l i s l e a d e r c l i p p i n g (MOF 1987) f o l l o w e d by immediate b u r n i n g of i n f e s t e d t e r m i n a l s (McLean, J.A. 1 p e r s o n a l c o m m u n i c a t i o n ) . However, c l i p p i n g i s an e x p e n s i v e , time consuming o p e r a t i o n and i t does not g i v e 100% c o n t r o l . In c o n t r a s t , l o s s e s caused by M. g e n t i l i s and C y l i n d r o c o p t u r u s sp. a r e not c o n s i d e r e d i m p o r t a n t ( F u r n i s s 1942; F e l l i n 1973; F u r n i s s and C a r o l i n 1977) and t h e r e f o r e no p r a c t i c a l measures have been dev e l o p e d f o r the c o n t r o l of th e s e w e e v i l s . Eaton (1942) and F u r n i s s (1942) suggested 1 P r o f e s s o r of F o r e s t Entomology, 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 , Vancouver, B.C. 9 some c o n t r o l measures t o c o n t r o l C y l i n d r o c o p t u r u s s p e c i e s which i n c l u d e d d e s t r u c t i o n of the immature s t a g e s of w e e v i l s by removing and b u r n i n g i n f e s t e d t r e e s , c o l l e c t i n g and d e s t r o y i n g a d u l t w e e v i l s and u s i n g c o n c e n t r a t e d s p r a y s ( l e a d a r s e n a t e and c r y o l i t e ) t o p r e v e n t a t t a c k . In the absence of e f f e c t i v e c o n t r o l methods, i t seems t h a t n a t u r a l c o n t r o l w i l l p l a y the most im p o r t a n t r o l e i n r e d u c t i o n of numbers of thes e p e s t s . N a t u r a l c o n t r o l i n c l u d e s a l l p h y s i c a l and b i o l o g i c a l f a c t o r s which have an e f f e c t on p e s t p o p u l a t i o n d e n s i t i e s (DeBach 1964). P a r a s i t i s m i s one of the most imp o r t a n t n a t u r a l m o r t a l i t y f a c t o r s . Enhancement of n a t u r a l p a r a s i t o i d p o p u l a t i o n s a l r e a d y p r e s e n t i n the environment c o u l d l e a d t o r e d u c t i o n i n p e s t numbers. G r e a t h e a d (1986) d i s t i n g u i s h e s m a n i p u l a t i o n s i n t e n d e d t o enhance the impact of n a t u r a l enemies from the term " c l a s s i c a l b i o l o g i c a l c o n t r o l " which i n c l u d e s i n t r o d u c t i o n and e s t a b l i s h m e n t of e x o t i c s p e c i e s t o s u p p r e s s p e s t p o p u l a t i o n s . Perhaps the knowledge of emergence p a t t e r n s of h o s t s and p a r a s i t o i d s and an u n d e r s t a n d i n g of the p h e n o l o g i e s of h o s t s and t h e i r p a r a s i t o i d s c o u l d r e s u l t i n the development of method(s) f o r enhancing n a t u r a l enemy p o p u l a t i o n s . However, emergence p a t t e r n s of i n s e c t s r e a r e d a r t i f i c i a l l y i n l a b o r a t o r i e s o f t e n do not match th o s e i n the f i e l d because development of most organisms i s t e m p e r a t u r e -dependent ( A l l e n 1976) and the h i g h e r t e m p e r a t u r e s i n 10 c o n t r o l l e d c o n d i t i o n s may cause development of organisms t o be u n n a t u r a l l y a c c e l e r a t e d . In 1735, Reaumur a t t e m p t e d t o i n t e g r a t e t emperature and time i n t o d e s c r i b i n g p o i k i l o t h e r m y of i n s e c t s and development of p l a n t s f o r the f i r s t time ( A l l e n 1976; H i g l e y e t a l . 1986). A l t h o u g h t h i s approach has been d e v e l o p i n g ever s i n c e , the ass u m p t i o n s of c a l c u l a t i n g degree-days have not changed ( H i g l e y e t a l . 1986). A l l e n (1976) s t a t e d t h a t "the most g e n e r a l assumption i s t h a t the r a t e of development i s some f u n c t i o n of t e m p e r a t u r e " . A c c o r d i n g t o H i g l e y e t a l . (1986) the r e l a t i o n s h i p of tem p e r a t u r e t o growth i s complex as a r e s u l t of v a r i o u s temperature-dependent b i o c h e m i c a l r e a c t i o n s i n or g a n i s m s , such as the a v a i l a b i l i t y of enzymes and t h e i r s u b s t r a t e s . F l u c t u a t i o n s i n tem p e r a t u r e can g r e a t l y a f f e c t the r a t e of l a r v a l development. U s u a l l y , the a c t u a l c u r v i l i n e a r r e l a t i o n s h i p between te m p e r a t u r e and growth r a t e ( F i g . 4) i s c a l c u l a t e d as a l i n e a r r e l a t i o n s h i p . T h i s approach i s a c c e p t a b l e i f the c o n s i d e r e d t e m p e r a t u r e s a r e i n the l i n e a r s e c t i o n of the f u n c t i o n ( H i g l e y e t a l . 1986). The type of a p p r o x i m a t i o n becomes i m p o r t a n t when we t a k e the d e v e l o p m e n t a l minimum and maximum te m p e r a t u r e s i n t o c o n s i d e r a t i o n . Techniques f o r c a l c u l a t i n g d e v e l o p m e n t a l maxima a r e not p r e c i s e and most o f t e n , they a r e not d e t e r m i n e d . C a l c u l a t i o n s of the minimum and maximum Temperature FIGURE 4. G e n e r a l i z e d r e l a t i o n s h i p of growth r a t e t o t e m p e r a t u r e (from H i g l e y e t a_l. 1986). 12 d e v e l o p m e n t a l t h r e s h o l d s a r e not p r e c i s e and may make degree-day c a l c u l a t i o n s l e s s a c c u r a t e . A common problem i s when t o b e g i n c a l c u l a t i o n s . U s u a l l y , c a l c u l a t i o n s s t a r t when t e m p e r a t u r e s exceed the lower t h r e s h o l d . However, development of the organism may not occur a t t h i s time ( H i g l e y et a l . 1986). Degree-day e s t i m a t e s i n c l u d e v a r i o u s e r r o r s , such as d i f f e r e n c e s between ambient and m i c r o h a b i t a t t e m p e r a t u r e s , or d i f f e r e n c e s between measurements of d i f f e r e n t weather s t a t i o n s . There a r e s e v e r a l approaches f o r c a l c u l a t i n g d e gree-days. The r e c t a n g l e method i s the most f r e q u e n t l y used method which c a l c u l a t e s the a r e a under the t i m e -t e m p e r a t u r e c u r v e as a r e c t a n g l e ( A r n o l d 1959), whereas the m o d i f i e d t r i a n g l e method and s i n e wave methods c a l c u l a t e the a r e a under the c u r v e on a h a l f - d a y b a s i s ( H i g l e y e t a l . 1986). 1.1.2 The host Lodgepole p i n e i s the most w i d e l y d i s t r i b u t e d c o n i f e r i n w estern N o r t h America ( F i g . 5) where i t spans 33 degrees of l a t i t u d e (Wheeler and C r i t c h f i e l d 1985) from s o u t h e a s t e r n A l a s k a and the i n t e r i o r of Yukon T e r r i t o r y t o the B a j a p e n i n s u l a i n C a l i f o r n i a (Maher 1982). I t s l o n g i t u d i n a l range spans from the N o r t h Coast t o the B l a c k H i l l s i n South Dakota (Maher 1982) over 3900m of e l e v a t i o n (Wheeler and C r i t c h f i e l d 1985). FIGURE 5. D i s t r i b u t i o n of P. t e r m i n a l i s and i t s h o s t s i n N o r t h America (from Maher 1982). 14 The n a t u r a l range of l o d g e p o l e p i n e i s s u b d i v i d e d i n t o t h r e e p r i n c i p a l r e g i o n s : (1) the P a c i f i c c o a s t , (2) the mountain c h a i n c o m p r i s i n g the Cascade Range, S i e r r a Nevada and the h i g h mountains of s o u t h e r n C a l i f o r n i a , (3) the I n t e r m o u n t a i n r e g i o n and Rocky mountain system from the Yukon t o s o u t h e r n C o l o r a d o (Wheeler and C r i t c h f i e l d 1985). P a t t e r n s of v a r i a t i o n i n m o r p h o l o g i c a l , p h y s i o l o g i c a l and b i o c h e m i c a l t r a i t s c o i n c i d e w i t h t h e s e major g e o g r a p h i c s u b d i v i s i o n s (Wheeler and C r i t c h f i e l d 1985). P i n u s c o n t o r t a D o u g l . v a r . l a t i f o l i a Engelm. w i l l be r e f e r r e d t o as l o d g e p o l e p i n e throughout t h i s s t u d y . In Canada, l o d g e p o l e p i n e grows i n t h r e e r e g i o n s : B r i t i s h C olumbia, A l b e r t a and the Yukon (Kennedy 1985), and i n B.C. i t a c c o u n t s f o r 15% of the the t o t a l s t a n d i n g volume of mature t i m b e r (McDougal 1973; Kennedy 1985). Lodgepole p i n e ranked second among t r e e s p e c i e s p l a n t e d i n B r i t i s h Columbia i n 1984-1986 and was exceeded o n l y by s p r u c e s ( T a b l e I I ) . Lodgepole p i n e e x h i b i t s a l a r g e e c o l o g i c a l a m p l i t u d e w i t h r e s p e c t t o c l i m a t e as i s seen from i t s wide growing range. T h i s t r e e s p e c i e s i s q u i t e v a r i a b l e i n i t s response t o c l i m a t i c f a c t o r s such as wind, h u m i d i t y and day l e n g t h . However, w i t h r e g a r d t o l i g h t , i t s r e q u i r e m e n t s a r e q u i t e c o n s t a n t ( S a t t e r l a n d 1973). Weetman e t a l . (1985) s t a t e d t h a t l o d g e p o l e p i n e i s not a n u t r i e n t demanding t r e e s p e c i e s 15 T a b l e I I . T o t a l number of t r e e s of major s p e c i e s p l a n t e d on Crown Land i n B r i t i s h Columbia i n 1984-1986 Spec i e s T o t a l number of t r e e s p l a n t e d ( thousands) 1 984- 1985 1985-1986 Engelmann, White Spruce 64, 052 57,832 Lodgepole P i n e 25, 289 29,981 D o u g l a s - F i r 15, 840 13,800 Am a b a l i s F i r 2, 635 1 ,381 Western Red Cedar 2, 244 2,385 S i t k a Spruce 2, 144 2,082 Source: M i n i s t r y of F o r e s t r y , Annual R e p o r t s 1984-1985, 1985-1986. 16 and i t grows w e l l on poor s i t e s but a l s o responds q u i c k l y t o n i t r o g e n f e r t i l i z a t i o n . F i r e p l a y s an i m p o r t a n t r o l e i n the e s t a b l i s h m e n t and s t r u c t u r e of l o d g e p o l e p i n e s t a n d s . Brown (1973) s t a t e d t h a t i n s t a n d s dominated by l o d g e p o l e p i n e f i r e u s u a l l y l e a d s t o the e s t a b l i s h m e n t of pure l o d g e p o l e p i n e s t a n d s due t o i t s s e r o t i n y . Lodgepole p i n e r e a d i l y e s t a b l i s h e s i t s e l f on c u t o v e r and burned-over s i t e s (Brown 1973) but i t o f t e n r e p r o d u c e s o v e r a b u n d a n t l y ( A l e x a n d e r 1960). As many as 100,000 s e e d l i n g s may e s t a b l i s h per a c r e (Woodhead 1934) and T a c k l e (1959) r e p o r t e d 2.5 m i l l i o n s e e d l i n g s per h e c t a r e . R a p i d r a t e s of j u v e n i l e growth h e l p l o d g e p o l e p i n e t o o v e r t o p most of competing t r e e s p e c i e s f o r the f i r s t two-t h r e e decades of i t s growth (Schmidt and A l e x a n d e r 1985). I n t e n s i v e i n t r a s p e c i f i c c o m p e t i t i o n f o r n u t r i e n t s , water, l i g h t and l i v i n g space slows the r a t e of growth i n o v e r s t o c k e d s t a n d s (Woodhead 1934; A l e x a n d e r 1960). Woodhead (1934) p o i n t e d out t h a t l a t e r n a t u r a l t h i n n i n g t a k e s p l a c e which r e s u l t s i n the d e a t h of l e s s v i g o r o u s t r e e s . However, t h i s n a t u r a l t h i n n i n g p r o c e s s i s not e f f i c i e n t i n terms of s i l v i c u l t u r e ( A l e x a n d e r 1960). O v e r s t o c k e d stands r e s u l t i n r e d u c t i o n i n d i a m e t e r and h e i g h t growth ( A l e x a n d e r 1960), t o t a l and mer c h a n t a b l e volume, i n i n c r e a s e d l e n g t h of r o t a t i o n and h i g h c o s t of h a r v e s t i n g (Johnstone 1985). E f f e c t s of o v e r s t o c k e d s t a n d s 17 can be reduced by j u v e n i l e s p a c i n g . Precommercial t h i n n i n g ( s p a c i n g ) i s a c c o m p l i s h e d through d i f f e r e n t o p e r a t i o n a l p r a c t i c e s which i n c l u d e s c h e m i c a l , m e c h a n i c a l and s e l e c t i v e t h i n n i n g . Johnstone (1985) i n d i c a t e d t h a t t h i n n i n g has a d i r e c t e f f e c t on dia m e t e r and h e i g h t growth of young t r e e s . Thus s p a c i n g i s i n c r e a s i n g l y r e c o g n i z e d as i m p o r t a n t as a s i l v i c u l t u r a l t e c h n i q u e t o improve the growth and merc h a n t a b l e y i e l d of dense l o d g e p o l e p i n e s t a n d s . B e s i d e t w i g w e e v i l s , e l o n g a t i n g l o d g e p o l e p i n e t e r m i n a l s a r e s u s c e p t i b l e t o a t t a c k by d i f f e r e n t i n s e c t s . The degree of impact v a r i e s from agent t o agent. T a b l e I I I summarizes phytophagous i n s e c t s damaging young l e a d e r s . 1.1.3 B i o l o g y of P i s s o d e s t e r m i n a l i s Hopping (1920) gave a d e s c r i p t i o n of P i s s o d e s  t e r m i n a l i s f o r the f i r s t time i n 1907, when he ob s e r v e d the i n s e c t and i t s damage i n young l o d g e p o l e p i n e s t a n d s i n Kern County, C a l i f o r n i a . Only t h r e e s h o r t a r t i c l e s c o n c e r n i n g the w e e v i l ' s d i s t r i b u t i o n , b i o l o g y and damage (Keen 1928; Salman 1935; Keen 1952) were p u b l i s h e d u n t i l t he e a r l y 1960's. S t u d i e s on P. t e r m i n a l i s commenced i n the e a r l y 1960's when D r o u i n e_t a l . (1963) r e p o r t e d the o c c u r r e n c e and l i f e h i s t o r y of the w e e v i l i n young st a n d s of j a c k p i n e , P i n u s b a n k s i a n a Lamb., from the p r a i r i e p r o v i n c e s of Canada. B e s i d e the p r a i r i e p r o v i n c e s , damage caused by the l o d g e p o l e t e r m i n a l w e e v i l was ob s e r v e d i n o t h e r p a r t s of Canada, 18 Table III. Insects recorded as being associated with leaders of young lodgepole ptne in western North America Order Fam i 1y COLEOPTERA Curculionidae Scolyt i dae LEPIDOPTERA 01ethreut1dae Genus/specles P i ssodes radlatae Hopk. P. term 1 na 1 1 s Hopp. Wagda11s h1spo1des LeC. M. gent i l l s LeC. M. 1econtei Horn. P i tyophthorus opimus Blm. Reference Furniss and Carol In ( 1977) Stark and Wood (1964) Stevens and Knopf (1974) Furniss and Carol in ( 1977) Undgren (1980) Evans (1982). B e l l a (1985) Keen (1952) Furniss and Carol1n (1977) F e l U n and Schmidt ( 1966) F e l l in ( 1973) Keen ( 1952), Fel11n ( 1973) Stevens (1973) Stevens and Knopf (1974) Eucosma sonomana Kft. Grant (1958) Petrova alblcapitana Busck.Furniss and Carol in ( 1977) P. metal 1 lea Busck. Furniss and C a r o l m ( 1977) Rhyacionia buoliana S c l f f . Furniss and Carol in ( 1977) Undgren (1980) 19 i n c l u d i n g A l b e r t a , B r i t i s h Columbia and the Northwest T e r r i t o r i e s (Stevenson and P e t t y 1968; B e l l a 1985a, 1985b), and i n the USA, i n c l u d i n g Oregon and Washington ( S t a r k and Wood 1964), C o l o r a d o and Idaho (Stevens and Knopf 1974) and C a l i f o r n i a , Wyoming and South Dakota ( F u r n i s s and C a r o l i n 1977). F i g u r e 5 shows the d i s t r i b u t i o n of P. t e r m i n a l i s i n N o r t h A m e r i c a . W i t h i n B r i t i s h Columbia the a n n u a l F o r e s t I n s e c t and D i s e a s e Survey (FIDS) has r e p o r t e d w e e v i l damage from d i f f e r e n t p a r t s of the p r o v i n c e . Stevens and Knopf (1974) d e s c r i b e d the egg of P. t e r m i n a l i s as b e i n g s u b - g l o b o s e , h a v i n g t h i n , t r a n s l u c e n t c h o r i o n and measuring 0.5x0.8 mm i n s i z e . In 1911, Hopkins gave a f u l l g e n e r i c d e s c r i p t i o n of the l a r v a l form of the genus P i s s o d e s , but no a c t u a l g e n e r i c d i a g n o s i s . He s t a t e d t h a t " a l l of these c h a r a c t e r s ( i . e . p r e s ence or absence of e y e s p o t s , the form and p r o p o r t i o n of s e v e r a l a n a t o m i c a l p a r t s , e t c ) have not been s u f f i c i e n t l y s t u d i e d t o p r e s e n t them i n t a b u l a r form f o r the i d e n t i f i c a t i o n of t h e s e s p e c i e s " . D i a g n o s t i c f e a t u r e s f o r P. t e r m i n a l i s l a r v a e have not been p u b l i s h e d and i t i s d i f f i c u l t t o d i s t i n g u i s h them from o t h e r w e e v i l l a r v a e such as t h o s e of M a q d a l i s . In g e n e r a l , l a r v a e of P. t e r m i n a l i s a r e creamy w h i t e i n c o l o r and have t a n head c a p s u l e s and l a c k abdominal p r o l e g s (Maher 1982). F u l l y d e v e l o p e d l a r v a e t e n d t o be l o n g e r than a d u l t s by a p p r o x i m a t e l y 1-3 mm (Stevens and Knopf 1974). 20 Pupae a r e creamy w h i t e (Stevens and Knopf 1974), or w h i t e (Duncan 1986), are a p p r o x i m a t e l y the same s i z e as the a d u l t , and bear a prominent s n o u t . A d u l t w e e v i l s a r e e a s i l y r e c o g n i z a b l e from t h e i r prominent r o s t r u m w i t h a p a i r of c l a v a t e antennae. Length of a d u l t s ranges from 5.5 mm t o 6.3 mm (Hopping 1920). The w e e v i l s a r e m o t t l e d y e l l o w i s h - b r o w n (Stevens and Knopf 1974) ( F i g . 6 ) . The pronotum i s g r a d u a l l y r e s t r i c t e d a n t e r i o r l y . The a n t e r i o r p a r t of the e l y t r a i s c o v e r e d w i t h y e l l o w s c a l e s , whereas t h e r e i s a more or l e s s f u s e d p o s t e r i o r band of w h i t e and y e l l o w s c a l e s near the v e r t e x of the d e c l e v i t y . These w h i t e bands a l s o e x t e n d a c r o s s the median p o r t i o n of the femora of both the mid and l e g s (Hopping 1920). A d u l t males and females of genus P i s s o d e s can be s e p a r a t e d by d i f f e r e n t c h a r a c t e r i s t i c s (Hopkins 1911). The most c o n v e n i e n t and most p r e c i s e way t o s e p a r a t e the sexes i s by e x a m i n a t i o n of the s e v e n t h and e i g h t h abdominal t e r g i t e s a f t e r removing the e l y t r a and membranous h i n d w i n g s . In males t h e r e a r e e i g h t v i s i b l e abdominal t e r g i t e s ; the s e v e n t h i s d i s t i n g u i s h e d by the b r o a d l y r e t u s e p o s t e r i o r m a r g i n , w h i l e the e i g h t h tergum i s prominent w i t h the apex rounded ( F i g . 7 ) . In females t h e r e a r e o n l y seven v i s i b l e abdominal t e r g i t e s , -the e i g h t h b e i n g c o m p l e t e l y c o v e r e d by the s e v e n t h ( F i g . 7) (Hopkins 1911). Hopkins (1911) mentioned o t h e r d i s t i n g u i s h i n g c h a r a c t e r i s t i c s between males and females as w e l l . A c c o r d i n g t o him, the r o s t r u m of the male i s s t o u t , s h o r t e r , l e s s s h i n i n g and more d i s t i n c t l y F i g u r e 6 A d u l t P i s s o d e s t e r m i n a l i s on e l o n g a t i n g l o d g e p o l e p i n e l e a d e r . 9 A A ) FIGURE 7: D o r s a l view of the l a s t abdominal .segment of the l o d g e p o l e t e r m i n a l w e e v i l . 23 p u n c t u r e d , whereas i n the female the beak i s l o n g e r , smoother and more s l e n d e r than i n males. The i n n e r a p i c a l t o o t h of the t i b i a e i s u s u a l l y more prominent. Chromosomal polymorphism a l s o e x i s t s between males and females of genus P i s s o d e s which was documented by Manna and Schmidt (1958) and then by Smith and Takenouchi (1969). S t a r k and Wood (1964) r e p o r t e d t h a t t h e r e a r e c o n s i d e r a b l e d i f f e r e n c e s i n the l i f e h i s t o r y of P. t e r m i n a l i s on j a c k p i n e i n the p r a i r i e p r o v i n c e s of Canada and on l o d g e p o l e p i n e i n C a l i f o r n i a . S i n c e the n a t u r a l range of j a c k p i n e d i d not i n c l u d e the study a r e a s the f o l l o w i n g d e s c r i p t i o n w i l l d e a l o n l y w i t h P. t e r m i n a l i s p o p u l a t i o n s a t t a c k i n g l o d g e p o l e p i n e . When the host i s l o d g e p o l e p i n e the l i f e c y c l e of the w e e v i l i s completed i n one y e a r . In the genus P i s s o d e s the l o d g e p o l e t e r m i n a l w e e v i l i s the o n l y s p e c i e s which " c o n s i s t e n t l y l a y s i t s eggs i n t o the d e v e l o p i n g t e r m i n a l " ( S t a r k and Wood 1964). C o n s e q u e n t l y , the l i f e of the w e e v i l must be s y n c h r o n i z e d w i t h the phenology of the t e r m i n a l s h o o t . A d u l t w e e v i l s emerge presumably from h i b e r n a t i o n s i t e s i n the ground ( S t a r k and Wood 1964) and i n f e s t e d l e a d e r s from l a t e A p r i l t o l a t e June ( F i g . 8 ) . A d u l t emergence i s f o l l o w e d by m a t u r a t i o n f e e d i n g d u r i n g which w e e v i l s f e e d on the phloem of e l o n g a t i n g t e r m i n a l s . A f t e r mating the female e x c a v a t e s a c y l i n d r i c a l o v i p o s i t i o n a l p u n c t u r e i n t o which i t d e p o s i t s up t o t h r e e 24 FIGURE 8. G e n e r a l i z e d l i f e c y c l e of P. t e r m i n a l i s on l o d g e p o l e p i n e (from P r i c e 1980) 25 eggs (D r o u i n et a l . 1963) and then s e a l s the h o l e w i t h f o e c a l p e l l e t ( S t a r k and Wood 1964). The t i s s u e s u r r o u n d i n g the p u n c t u r e i s c h a r a c t e r i z e d by purple-brown d i s c o l o r a t i o n . O v i p o s i t i o n o c c u r s on the main stem of the c u r r e n t y e a r ' s t e r m i n a l growth which makes P. t e r m i n a l i s unique i n the genus i n terms of o v i p o s i t i o n . Eggs h a t c h w i t h i n two weeks (Stevens and Knopf 1974) and f i r s t i n s t a r l a r v a e b e g i n random m i n i n g under the e p i d e r m i s . Second i n s t a r l a r v a e a r e c h a r a c t e r i z e d by n e g a t i v e g e o t r o p i s m ( D r o u i n e t a_l. 1963) as they mine toward the a p i c a l bud ( F i g . 9 ) . As a r e s u l t of r u p t u r i n g r e s i n d u c t s l a r v a l f e e d i n g t u n n e l s f i l l w i t h r e s i n and t h e s e a r e a s become p u r p l i s h ( S t evens and Knopf 1974). The s p i r a l f a s h i o n of f e e d i n g i n the cambium r e g i o n k i l l s the c u r r e n t y e a r ' s l e a d e r . L a t e r i n s t a r s move i n t o the p i t h and c o n t i n u e t u n n e l i n g . L a r v a e r e a c h m a t u r i t y d u r i n g the f o u r t h i n s t a r . F a l l t e m p e r a t u r e s a f f e c t l a r v a l development and t hey might pupate and a d u l t s even emerge and o v e r w i n t e r p r o b a b l y i n the d u f f . However, the m a j o r i t y of the w e e v i l s o v e r w i n t e r i n the p i t h as l a r v a e and complete t h e i r development and pupate i n the f o l l o w i n g s p r i n g ( S t evens and Knopf 1974) ( F i g . 8 ) . D i f f e r e n t f a c t o r s c o n t r i b u t e t o w e e v i l m o r t a l i t y t h roughout the l i f e c y c l e of the w e e v i l . High m o r t a l i t y from drowning i n r e s i n o c c u r s among e a r l y l a r v a l s t a g e s when l a r v a e r u p t u r e r e s i n d u c t s of the e l o n g a t i n g l e a d e r w h i l e 26 F i g u r e 9 Second i n s t a r P. t e r m i n a l i s l a r v a m i n i n g beneath the e p i d e r m i s of the c u r r e n t y e a r ' s t e r m i n a l shoot . 27 a t t e m p t i n g t o e s t a b l i s h f e e d i n g s i t e s ( D r o u i n et a_l. 1963). Another f a c t o r which c o n t r i b u t e s t o h i g h l a r v a l m o r t a l i t y i s p a r a s i t i s m , e s p e c i a l l y among e a r l y i n s t a r l a r v a e ( S t a r k and Wood 1964). Maher (1982) s t a t e d t h a t w e e v i l m o r t a l i t y i s g r e a t e s t d u r i n g the l a r v a l and pupal s t a g e s . When l a t e r i n s t a r l a r v a e move i n t o the p i t h , the r o l e of p a r a s i t i s m d e c r e a s e s and s t a r v a t i o n p l a y s an i m p o r t a n t r o l e i n r e d u c i n g l a r v a l s u r v i v a l . Sometimes 5-6 l a r v a e e n t e r the p i t h and t h e r e may not be s u f f i c i e n t food f o r a l l l a r v a e t o complete t h e i r development. T h i s r e s u l t s i n s t a r v a t i o n and d e a t h . A l t h o u g h Finnegan (1958) r e p o r t e d 90% p r e d a t i o n of P i s s o d e s  a p p r o x i m a t u s Hopkins ( C o l e o p t e r a : C u r c u l i o n i d a e ) by the downy woodpecker, Dendrocopus pubescens medianus (S w a i n s o n ) , the r o l e of b i r d p r e d a t i o n i n r e g u l a t i n g P. t e r m i n a l i s p o p u l a t i o n s has not y e t been d e t e r m i n e d . Some p a r a s i t i s m a l s o o c c u r s d u r i n g the p u p a l p e r i o d . A l i s t of p a r a s i t o i d s a s s o c i a t e d w i t h the l o d g e p o l e t e r m i n a l w e e v i l i s p r e s e n t e d i n Ta b l e IV. There a r e i n s e c t s , o t h e r than p a r a s i t o i d s found i n a s s o c i a t i o n w i t h P. t e r m i n a l i s . S t e v e n s (1973) found a s m a l l s c o l y t i d b e e t l e , P i t y o p h t h o r u s opimus Blackman (COLEOPTERA: S c o l y t i d a e ) l i v i n g commensally w i t h the w e e v i l . P. opimus u t i l i z e s p o r t i o n s of the phloem and xylem of the t e r m i n a l a f t e r i t s t a r t s t o d e s s i c a t e as a r e s u l t of w e e v i l a c t i v i t y . Stevens (1973) c o l l e c t e d t h r e e s p e c i e s of p a r a s i t i c Hymenoptera from P. opimus g a l l e r i e s : A c e r o c e p h a l a a t r o v i o l a c e a Cwfd. ( P t e r o m a l i d a e ) , Eurytoma T a b l e IV. P a r a s i t i c Hymenoptera 1 of P i s s o d e s t e r m i n a l i s O r d e r / F a m i l y Spec i e s HYMENOPTERA BETHYLIDAE BRACONIDAE EULOPHIDAE EURYTOMIDAE ICHNEUMONIDAE PTEROMALIDAE Cephalonomia (?) h y a l i n i p e n n i s B r a c h i s t e s sp. Bracon p i n i Mues. Eub a d i z o n s t r i g i t e r g u m (Cushman) Tr i a s p i s p i ssodes V i e r e c k T e t r a s t i c h u s s p. Eurytoma p i c e a e Bugbee Eurytoma p i ssodes G i r a u l t Eurytoma sp. n r . c l e r i D o l i c h o m i t u s t e r e b r a n s n u b i l i p e n n i s V i e r H e l c o s t i z u s s u b r e c t u s Townes H a b r o c y t u s sp. Mesopolobus sp. R h o p a l i c h u s p u l c h r i p e n n i s (Cwfd.) 1 From S t a r k and Wood (1964), Stevenson and P e t t y ( 1 968), Stevens and Knopf (1974). 29 t o m i c i Ashmed (Eurytomidae) and R h o p a l i c h u s p u l c h r i p e n n i s Cwfd. ( P t e r o m a l i d a e ) . The l a t t e r two s p e c i e s a r e a l s o p a r a s i t o i d s of P. t e r m i n a l i s (Table I V ) . 1.1.4. B i o l o g y of M a g d a l i s g e n t i l i s The genus M a g d a l i s i n c l u d e s 25 s p e c i e s i n Europe but o n l y f r a g m e n t a r y i n f o r m a t i o n i s a v a i l a b l e about t h e i r b i o l o g y and economic importance i n the l i t e r a t u r e (Bukzeeva 1965). A l t h o u g h a few c o l l e c t i o n s of M a g d a l i s spp. had been made p r i o r t o 1963, t h e r e were no r e c o r d s t h a t any s p e c i e s of t h i s genus had caused any s i g n i f i c a n t amount of damage i n the N o r t h e r n Rocky Mountains ( F e l l i n 1973). The f i r s t s i g n i f i c a n t damage caused by M. q e n t i l i s LeC. was r e c o r d e d i n the L e w i s and C l a r k N a t i o n a l F o r e s t i n w e s t - c e n t r a l Montana i n 1965 ( F e l l i n and Schmidt 1966). T h i s w e e v i l o c c u r s from C a l i f o r n i a t o B r i t i s h Columbia and Montana ( F u r n i s s and C a r o l i n 1977). H o s t s of the w e e v i l i n c l u d e ponderosa p i n e , P i n u s ponderosae D o u g l . , J e f f r e y p i n e , P i n u s  j e f f r e y i G r e v i l l e and B a l f o u r ( F u r n i s s and C a r o l i n 1977), and l o d g e p o l e p i n e ( F e l l i n and Schmidt 1966). B e s i d e i t s damage, M. q e n t i l i s has not been s t u d i e d i n d e t a i l and t h e r e i s no a v a i l a b l e i n f o r m a t i o n on i t s b i o l o g y or l i f e c y c l e i n the l i t e r a t u r e . G e n e r i c d i a g n o s t i c f e a t u r e s of a d u l t s and l a r v a e a l s o have not y e t been d e s c r i b e d . The l i t e r a t u r e a l s o l a c k s i n f o r m a t i o n on i n s e c t s , such as p a r a s i t o i d s , a s s o c i a t e d w i t h t h i s w e e v i l . 30 1.1.5. B i o l o g y of C y l i n d r o c o p t u r u s spp. The genus C y l i n d r o c o p t u r u s i s w i d e l y d i s t r i b u t e d i n western N o r t h A m e r i c a . D i f f e r e n t r e p r e s e n t a t i v e s of the genus can be found from A l b e r t a and B r i t i s h Columbia t o Washington and C a l i f o r n i a and from Oregon t o Montana ( F u r n i s s and C a r o l i n 1977). Hosts i n c l u d e many s p e c i e s of p i n e s but w e e v i l s a l s o a t t a c k D o u g l a s - f i r , Pseudotsuga  m e n z i e s i i ( M i r b . ) F r a n c o ( F u r n i s s 1942; Eaton 1942; F u r n i s s and C a r o l i n 1977). Eggs a r e o v o i d ( F u r n i s s 1942) o r pear-shaped (Eaton 1942) and 0.48mm l o n g and 0.30mm wide ( F u r n i s s 1942; Eaton 1942). F i n e d i a g n o s t i c f e a t u r e s of l a r v a e of C y l i n d r o c o p t u r u s were g i v e n by Anderson (1941). He d e s c r i b e d the l a r v a ( F i g . 10) as s t o u t , c u r v e d and c y l i n d r i c a l . A p p r o x i m a t e l y o n e - h a l f of the head i s r e t r a c t e d i n t o the p r o t h o r a x and i t s a n t e r i o r p a r t c o n t a i n s a few s e t a e . The p o s t e r i o r h a l f of the p r o t h o r a x c o n t a i n s many a s p e r i t i e s which make t h i s p a r t c o n s p i c u o u s . The c o l o r of the body ranges from w h i t e t o c r e a m - c o l o r e d depending upon the age of the l a r v a (Eaton 1942), and the body l e n g t h v a r i e s between 3.0-3.5mm ( F u r n i s s 1942). The pupa ( F i g . 10), i s c r e a m - c o l o r e d as are mature l a r v a e but s l i g h t l y l o n g e r than the a d u l t . A d u l t s a r e c o v e r e d w i t h l i g h t and dark s c a l e s which makes them appear gray i n c o l o r . L e n g t h of males and 31 FIGURE 10. L a r v a (A) and pupa (B) of genus C y l i n d r o c o p t u r u s (from Anderson 1941). 32 females ranges from 2.5mm t o 2.6mm (Eaton 1942; F u r n i s s 1942). Eaton (1942) gave some e x t e r n a l d i s t i n g u i s h i n g c h a r a c t e r s which s e p a r a t e males and fe m a l e s : "the most r e l i a b l e c h a r a c t e r i s the median i m p r e s s i o n of the f i r s t v i s i b l e abdominal s t e r n i t e " . T h i s s t e r n i t e i s s t r o n g l y i m p r essed i n the c e n t r e i n the males w h i l e i t i s "convex and o n l y s l i g h t l y f l a t t e n e d i n the f e m a l e s " . D i f f e r e n t s p e c i e s of the genus emerge a t d i f f e r e n t t i m e s from May through August ( F u r n i s s 1942; Eaton 1942; F u r n i s s and C a r o l i n 1977). A f t e r some m a t u r a t i o n f e e d i n g t h e female o v i p o s i t s a s i n g l e egg i n t o each o v i p o s i t i o n a l p u n c t u r e and s e a l s each w i t h a f a e c a l p e l l e t . O v i p o s i t i o n o c c u r s i n stems and bran c h e s . When e c l o s i o n o c c u r s l a r v a e s t a r t f e e d i n g which r e s u l t s i n g i r d l i n g of the b r a n c h e s . As l a r v a e age, f e e d i n g t u n n e l s run t o g e t h e r and the e n t i r e phloem-cambium r e g i o n i s d e s t r o y e d (Eaton 1942). Most of the l a r v a e o v e r w i n t e r i n the wood and p u p a t i o n t a k e s p l a c e t h e f o l l o w i n g s p r i n g . P a r a s i t i s m of C y l i n d r o c o p t u r u s spp. o c c u r s t h r o u g h o u t t h e i r l i f e c y c l e . A l i s t of i n s e c t s a s s o c i a t e d w i t h t h e s e w e e v i l s i s p r e s e n t e d i n T a b l e V. 33 T a b l e V. P a r a s i t i c Hymenoptera 1 of C y l i n d r o c o p t u r u s spp. O r d e r / F a m i l y Spec i e s HYMENOPTERA BRACONIDAE EULOPHIDAE EUPELMIDAE EURYTOMIDAE ICHNEUMONIDAE PTEROMALIDAE SCELIONIDAE TRICHOGRAMMATIDAE Den d r o s o t e r scaber Mues. M i c r o b r a c o n p i n i Mues. M i c r o t o n u s n. sp. U r o s i g a l p h u s p i n i Cush. Euderus a r g y r e s t h i a e (Cwfd.) Euderus subopaca (Gahan) T e t r a s t i c h u s sp. E u p h e l m e l l a v a s i c u l a r i s ( R e t z . ) Eurytoma t o m i c i Ashm. C a l l i e p h i a l t e s c o m s t o c k i i ( C r e s s . ) Amblymerus near v e r d i t o r ( N o r t . ) C a e n a c i s sp. C e c i d o s t i b a d e n d r o c t o n i Ashm. R h o p a l i c h u s p u l c h r i p e n n i s Cwfd. Telenomus sp. Z a g e l l a sp. 1 From Eaton (1942) and F u r n i s s (1942). 34 1.2 The major o b j e c t i v e s of the p r e s e n t study were: 1. To determine the i n s e c t s i n f e s t i n g l o d g e p o l e p i n e t e r m i n a l s . 2. To i d e n t i f y emerging p a r a s i t o i d s from i n f e s t e d l o d g e p o l e p i n e l e a d e r s i n B r i t i s h C olumbia. 3. To d e t e r m i n e w e e v i l and p a r a s i t o i d emergence p a t t e r n s from i n f e s t e d l o d g e p o l e p i n e l e a d e r s i n o r d e r t o e v a l u a t e l e a d e r c l i p p i n g s t r a t e g i e s t o m i n i m i z e w e e v i l numbers and enhance n a t u r a l p a r a s i t o i d p o p u l a t i o n s . 35 2. METHODS AND MATERIALS 2.1. L o c a t i o n of the st u d y In 1986, l o c a t i o n s of the p r e l i m i n a r y s t u d i e s were chosen on the b a s i s of a n n u a l r e p o r t s of tho s e F o r e s t R egions where l o d g e p o l e t e r m i n a l w e e v i l a t t a c k s had been r e p o r t e d by the Canadian F o r e s t r y S e r v i c e ' s F o r e s t I n s e c t and D i s e a s e Survey ( F I D S ) . Samples of i n f e s t e d l o d g e p o l e p i n e t e r m i n a l s were c o l l e c t e d from s i x l o c a t i o n s i n s o u t h c e n t r a l B r i t i s h Columbia ( F i g . 11). One t h i r d of the s e i n f e s t e d l e a d e r s were d i s s e c t e d and the remainder were s e t up f o r mass r e a r i n g i n waxed s e e d l i n g boxes a t UBC. Based on the r e s u l t s of the r e a r i n g e x p e r i m e n t , s a m p l i n g p l o t s were chosen a t R i s k e Creek, 80km west of W i l l i a m s Lake i n February,1987. The 1987 summer r e s e a r c h was c a r r i e d out i n the s o u t h e r n i n t e r i o r of B r i t i s h Columbia ( F i g . 12-13). T h i s study was s u p p o r t e d by the a v a i l a b i l i t y of the f a c i l i t i e s of the A g r i c u l t u r e Canada Re s e a r c h S t a t i o n i n Summerland ( F i g . 12). 2.2. D i s s e c t i o n s of i n f e s t e d l e a d e r s One t h i r d of the c o l l e c t e d l e a d e r s from each s i t e were d i s s e c t e d i n the l a b o r a t o r y u s i n g a d i s s e c t i n g m i c r o s c o p e . The 1987 w i n t e r d i s s e c t i o n was c a r r i e d out t o : - de t e r m i n e o v e r w i n t e r i n g s t a g e s of the w e e v i l s and p a r a s i t o i d s . F i g u r e 11. Map showing the l o c a t i o n s of c o l l e c t i o n s of i n f e s t e d l e a d e r s i n s o u t h - c e n t r a l B r i t i s h Columbia i n the summer of 1986. u> FIGURE 12. L o c a t i o n s of t h e c o l l e c t i o n p l o t s i n t h e s o u t h e r n i n t e r i o r of B r i t i s h Columbia- i n the sur.rr.er cf 19B7. 38 ) 4 u \ .'X418 \ •' { i r i vv 428 _ —. ( - < 3 ^ 209 ! ) / "slash i I -.T. r , r ^ road / / / F i g u r e 13. L o c a t i o n s of t h e . s t u d y and s u r v e y p l o t s a t E l l i s c r e e k , near P e n t i c t o n , B.C. 39 The 1987 summer d i s s e c t i o n s aimed t o : - d etermine the o v e r w i n t e r i n g s u c c e s s of w e e v i l s and p a r a s i t o i d s , and - m o n i t o r the development of h o s t w e e v i l s and t h e i r p a r a s i t o i d s . 2.2.1. Sampling t e c h n i q u e s The 1987 summer d i s s e c t i o n i n c l u d e d weekly c o l l e c t i o n s and d i s s e c t i o n s of c u r r e n t y e a r ' s t e r m i n a l s showing symptoms of w e e v i l a t t a c k . S i x randomly chosen i n f e s t e d l e a d e r s were c o l l e c t e d weekly from June 5 t o June 30,1987 f o l l o w e d by 10 l e a d e r s weekly between J u l y 8 and Sept. 30,1987. There were i n s u f f i c i e n t i n f e s t e d t e r m i n a l s a t the b e g i n n i n g of the study t o c o l l e c t 10 l e a d e r s e v e r y week. As emergence of a d u l t w e e v i l s c o n t i n u e d , the number of a t t a c k e d t e r m i n a l s i n c r e a s e d which a l l o w e d the c o l l e c t i o n of 10 i n f e s t e d t e r m i n a l s / w e e k . I n f e s t e d l e a d e r s were c l i p p e d w i t h a hand pruner and taken back t o the l a b o r a t o r y where they were d i s s e c t e d . A s p e c i a l d a t a form was c r e a t e d f o r t h i s d i s s e c t i o n on which l e a d e r l e n g t h , b a s a l and a p i c a l l e a d e r d i a m e t e r , number of w e e v i l and p a r a s i t o i d emergence h o l e s , and a l l host and p a r a s i t o i d d e v e l o p m e n t a l s t a g e s were r e c o r d e d (Appendix I ) . A l l d a t a were grouped a c c o r d i n g t o the type of d i s s e c t i o n (May d i s s e c t i o n or weekly d i s s e c t i o n of 40 c u r r e n t y e a r ' s a t t a c k s ) . T o t a l s were summed f o r p r e s e n t a t i o n i n T a b l e s . A 33ha young, spaced l o d g e p o l e p i n e s t a n d ( s i t e 209, F i g . 13) was s u r v e y e d f o r w e e v i l a t t a c k i n c i d e n c e and a t t a c k h i s t o r y . A s y s t e m a t i c s a m p l i n g system was used i n the s u r v e y . The d i s t a n c e between the t r a n s e c t l i n e s was 15m and a t r e e was sampled a t 10m i n t e r v a l s a l o n g the t r a n s e c t l i n e . I f t h e r e was no t r e e a t the 10m p o i n t , the c l o s e s t t r e e w i t h i n a 5m r a d i u s was chosen f o r s a m p l i n g . I f two t r e e s were e q u a l l y f a r from the 10m s i g n a c o i n t o s s was used t o d e c i d e which t r e e would be sampled. 2.3. I n c u b a t i o n of i n f e s t e d l e a d e r s Two t h i r d s of the c o l l e c t e d a t t a c k e d t e r m i n a l s from each s i t e were s e t up f o r e i t h e r mass or i n d i v i d u a l r e a r i n g s i n m o d i f i e d waxed s e e d l i n g boxes ( F i g . 14) and paper m a i l i n g t u b e s ( F i g . 15), r e s p e c t i v e l y . A s m a l l j a r or v i a l was i n s e r t e d i n t o the upper h a l f of each box ( F i g . 14) or i n t o the lower h a l f of one of the caps of t h e m a i l i n g tubes ( F i g . 15). Boxes and tubes were kept at room temperature and emerging specimens were c o l l e c t e d and r e c o r d e d d a i l y . Emerged specimens were kept e i t h e r i n 70% a l c o h o l or were p i n n e d . P i n n e d specimens were sent t o the B i o s y s t e m a t i c R e s e a r c h I n s t i t u t e i n Ottawa f o r i d e n t i f i c a t i o n . A c o l l e c t i o n of p i n n e d specimens i s m a i n t a i n e d i n the F o r e s t Entomology l a b o r a t o r y of the F a c u l t y of F o r e s t r y a t UBC. 41 F i g u r e 14 Mass r e a r i n g of i n f e s t e d l o d g e p o l e p i n e l e a d e r s i n m o d i f i e d s e e d l i n g boxes. Note emergence j a r s e t i n t o the s i d e of t h e box a t t o p l e f t . 42 F i g u r e 15 I n d i v i d u a l r e a r i n g of i n f e s t e d l o d g e p o l e p i n e l e a d e r s i n m a i l i n g t u b e s . Note the v i a l e n t e r e d i n t o t h e lower h a l f of t h e cap. 43 R e a r i n g s and l e a d e r e l o n g a t i o n measurements aimed t o compare the r e l a t i o n s h i p between w e e v i l emergence p a t t e r n s and l e a d e r phenology. T w e n t y - f i v e l o d g e p o l e p i n e t r e e s were randomly chosen on s i t e and marked w i t h r e d p l a s t i c r i b b o n s . A number was a s s i g n e d t o each r i b b o n so t h a t measurements c o u l d be taken from the same t r e e s r e p e a t e d l y . Leader e l o n g a t i o n was measured and r e c o r d e d e v e r y 7 days. R e a r i n g s a l s o h e l p e d t o : - d e s c r i b e emergence p a t t e r n s of t h e w e e v i l s and t h e i r a s s o c i a t e d p a r a s i t o i d s from i n f e s t e d l o d g e p o l e p i n e l e a d e r s , - p r o v i d e specimens f o r i d e n t i f i c a t i o n , - d e s c r i b e the r e l a t i o n s h i p between host and p a r a s i t o i d p h e n o l o g i e s . 2.4. D i r e c t o b s e r v a t i o n s and measurements D i r e c t o b s e r v a t i o n s were c a r r i e d out i n the f i e l d and i n the l a b o r a t o r y t o : - o b t a i n a b e t t e r u n d e r s t a n d i n g of the b i o l o g y of the w e e v i l s by o b s e r v i n g f e e d i n g h a b i t s , o v i p o s i t i o n and damage caused by d i f f e r e n t w e e v i l s p e c i e s , - o b t a i n i n f o r m a t i o n on the f e e d i n g h a b i t s of the p a r a s i t o i d s , and - measure the l o n g e v i t y and f e c u n d i t y of P. t e r m i n a l i s . 44 Ten p a i r s of P. t e r m i n a l i s a d u l t s were p l a c e d i n 0.5 L j a r s c o v e r e d w i t h cheese c l o t h and kept a t room temperature (20+2°C). Each day a 10-cm-long s e c t i o n of l o d g e p o l e p i n e t e r m i n a l was p l a c e d i n each j a r . The l e a d e r s were kept i n p l a s t i c bags i n a r e f r i g e r a t o r u n t i l used. D i a m e t e r s of the removed t e r m i n a l s were measured a t m i d - p o i n t and the number of n e e d l e f a s c i c l e s c o u n t e d . Numbers of f e e d i n g p u n c t u r e s and o v i p o s i t i o n s i t e s were cou n t e d w i t h the a i d of a d i s s e c t i n g m i c r o s c o p e . O v i p o s i t i o n s i t e s were opened and the number of eggs per s i t e c o u n t e d and r e c o r d e d . D i r e c t o b s e r v a t i o n s a l s o i n c l u d e d measuring the d u r a t i o n of the egg stage of P. t e r m i n a l i s . I n c u b a t i o n was d e t e r m i n e d o n l y f o r those eggs which were kept under l a b o r a t o r y c o n d i t i o n s . S e c t i o n s of c u r r e n t y e a r ' s t e r m i n a l growth w i t h o v i p o s i t i o n s i t e s were o b t a i n e d from the l o n g e v i t y and f e c u n d i t y e xperiment d e s c r i b e d e a r l i e r . These l e a d e r s e c t i o n s were p l a c e d on m o i s t (but not wet) paper t o w e l s i n paper boxes which were c l o s e d and kept a t room t e m p e r a t u r e . D e s s i c a t i o n of l e a d e r s was p r e v e n t e d by m o i s t e n i n g the paper t o w e l s i n the bottom of the boxes d a i l y . Eggs were checked d a i l y under a d i s s e c t i n g m i c r o s c -ope i n o r d e r t o r e c o r d h a t c h i n g of the eggs. A f t e r e x a m i n a t i o n , o v i p o s i t i o n s i t e s were c l o s e d t o p r e v e n t d e h y d r a t i o n of eggs. Date of o v i p o s i t i o n and date of h a t c h i n g were r e c o r d e d . Widths of head c a p s u l e s of newly 45 h a t c h e d l a r v a e were measured u s i n g an o c u l a r micrometer i n a d i s s e c t i n g m i c r o s c o p e . O b s e r v a t i o n s were a l s o c a r r i e d out t o measure the p u p a l p e r i o d of P. t e r m i n a l i s . Twenty w e e v i l l a r v a e were o b t a i n e d from the May d i s s e c t i o n . Each l a r v a was kept i n a s e p a r a t e numbered p e t r i d i s h . Dates of p u p a t i o n and a d u l t emergence were r e c o r d e d f o r a l l 20 l a r v a e . 2.5. Data a n a l y s i s I n c u b a t i o n of i n f e s t e d l e a d e r s i n the l a b o r a t o r y a c c e l e r a t e d the development and emergence of w e e v i l s and p a r a s i t o i d s . To c a l c u l a t e the number of accumulated degree-d a y s , a degree-day program, (DEGDAY) ( H i g l e y e t a l . 1986) was used. The program c a l c u l a t e d the number of h e a t i n g degree-days u s i n g the r e c t a n g l e , t r i a n g l e and sine-wave methods. Minimum d e v e l o p m e n t a l t h r e s h o l d t e m p e r a t u r e s , (TMIN) f o r the w e e v i l s a r e not known, so h e a t i n g degree-days were c a l c u l a t e d f o r a r b i t r a r i l y chosen TMIN v a l u e s (TMIN=4,5,6,7 and 8°C). The maximum de v e l o p m e n t a l t h r e s h o l d t e m p e r a t u r e , (TMAX) was unknown and when TMAX was a l s o unknown and DEGDAY a l l o w e d f o r an "unknown" s e t t i n g and i t was a l s o run w i t h TMAX=30°C. The program was run on an IBM-AT p e r s o n a l computer a t UBC. 46 3. RESULTS AND DISCUSSION 3.1. Summer c o l l e c t i o n s i n 1986 A t o t a l of 1100 i n f e s t e d l e a d e r s were c o l l e c t e d t h r o u g h o u t the p r o v i n c e i n June,1986. I n c u b a t i o n of i n f e s t e d t e r m i n a l s showed t h a t b e s i d e the l o d g e p o l e t e r m i n a l w e e v i l , a n o t h e r w e e v i l , M a g d a l i s g e n t i l i s , i s a l s o p r e s e n t i n the P r i n c e George and W i l l i a m s Lake a r e a s ( T a b l e V I ) . 3.2. F e b r u a r y l e a d e r c o l l e c t i o n i n 1987 As a r e s u l t of h i g h w e e v i l a t t a c k i n c i d e n c e and easy a v a i l a b i l i t y , i n f e s t e d t e r m i n a l s (335) were c o l l e c t e d a t R i s k e C reek, near W i l l i a m s Lake i n F e b r u a r y , 1 9 8 7 . Two t y p e s of symptoms of w e e v i l a t t a c k were o b s e r v e d d u r i n g c o l l e c t i n g . F i g u r e 16A shows an i n f e s t e d t e r m i n a l which s t i l l has i t s a p i c a l dominance but i t s n e e d l e s are dead and had changed c o l o r t o d u l l brown. In F i g u r e 16B the t e r m i n a l i s s t i l l green but i t has l o s t i t s a p i c a l dominance. T h i s o b s e r v a t i o n s u g g e s t s t h a t t h i s t e r m i n a l grew s l o w l y a f t e r the a t t a c k and the l a t e r a l s had a chance t o c a t c h up w i t h the t e r m i n a l . I f the a t t a c k i s not heavy or l a r v a e had been s u c c e s s f u l l y p i t c h e d out the t e r m i n a l manages t o m a i n t a i n i t s m o i s t u r e c o n t e n t and i t remains green but i t s growth i s slowed down. T h i s r a i s e s the q u e s t i o n , "Does each symptom s p e c i f i c a l l y r e s u l t from one of the w e e v i l s p e c i e s ? " To 47 T a b l e VI. Summary of t e r m i n a l w e e v i l s and p a r a s i t o i d s r e a r e d from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d throughout B r i t i s h Columbia i n June ,1986 L o c a t i o n P. t e r m i n a l i s M. g e n t i l i s P a r a s i t e s P r i n c e George 6 1 56 P e n t i c t o n 418 19 — 3 P e n t i c t o n 421 4 - 2 Kelowna 14 - 31 Kamloops 1 3 - 13 Kamloops 2 1 — 18 W i l l i a m s Lake 1 4 1 15 W i l l i a m s Lake 2 - - 1 1 M e r r i t t — — 19 T o t a l 51 2 168 < 48 A B F i g u r e 16 Two t y p e s of symptoms of i n f e s t e d l o d g e p o l e p i n e l e a d e r s by t e r m i n a l w e e v i l s . The t e r m i n a l s t i l l has i t s a p i c a l dominance but i t s n e e d l e s have a l r e a d y changed c o l o r . ( A ) . The t e r m i n a l i s s t i l l g reen but has l o s t i t s a p i c a l dominance and l a t e r a l s have caught up w i t h the i t ( B ) . 49 t e s t t h i s h y p o t h e s i s , i n f e s t e d l e a d e r s e x h i b i t i n g e i t h e r of t h e s e symptoms would need t o be r e a r e d i n d i v i d u a l l y . D r o u i n et a l . (1963) r e p o r t e d t h a t n e e d l e s of t e r m i n a l s h o o t s a t t a c k e d by P. t e r m i n a l i s changed c o l o r and were c l e a r l y r e c o g n i z a b l e by e a r l y August. In the c u r r e n t study i t was o b s e r v e d t h a t n e e d l e s of a t t a c k e d l e a d e r s t u r n e d brown ( w i t h a few e x c e p t i o n s ) o n l y i n J a n u a r y - F e b r u a r y of the f o l l o w i n g y e a r . In August, i d e n t i f i c a t i o n of d y i n g t e r m i n a l s was p o s s i b l e o n l y by c l o s e e x a m i n a t i o n of the t e r m i n a l s . R e s u l t s of d i s s e c t i o n s a r e summarized i n T a b l e V I I . I t i s not known which w e e v i l s p e c i e s c o m p l e t e d i t s development and emerged i n the p r e v i o u s f a l l l e a v i n g the f o u r emergence h o l e s . W e e v i l l a r v a l m o r t a l i t y ranged from 54% t o 60% on the t h r e e s i t e s . As d i a g n o s t i c f e a t u r e s of the l a r v a e of P. t e r m i n a l i s and M. g e n t i l i s were not a v a i l a b l e i t was not p o s s i b l e t o d e t e r m i n e the r a t i o of the l i v e and dead l a r v a e of the two s p e c i e s . Advanced decay of the two dead pupae i n d i c a t e d t h a t these pupae must have d i e d i n the p r e v i o u s f a l l . The m a j o r i t y of t h e w e e v i l s o v e r w i n t e r e d as l a r v a e r a t h e r than as pupae. I n c u b a t i o n of i n f e s t e d l o d g e p o l e p i n e t e r m i n a l s was v e r y s u c c e s s f u l . Only one dead P. t e r m i n a l i s a d u l t was found i n one of the boxes a f t e r the r e a r i n g had f i n i s h e d . 50 Table VII. Summary of the Insect l i f e stages dissected from Infested leaders of lodgepole pine c o l l e c t e d from stands on the Palmer Lake Road, Riskle Creek, near Will1ams Lake In February 1987 . LocatIon Number of leaders Emergence h o l e s 1 Weevil l i f e 5 stages Paras 1 to Id l i f e stages dissected W P adult pupae L D L D 1arvae L D adult pupae L D L D 1arvae L D 15km l e f t 36 1 2 30 40 - -15km r i g h t 27 - - 24 29 - -21km r i g h t 50 4 1 3 30 42 4 1 - 1 -1 W = weevil 2 L = 11ve, , P = p a r a s i t o i d D = dead 51 F i f t y t h r e e M. gent i l i s and o n l y two P. t e r m i n a l i s a d u l t s emerged from the boxes (T a b l e V I I I ) . The f i r s t w e e v i l s p e c i e s t o emerge was M. g e n t i 1 i s ( F i g . 17). Emergence s t a r t e d on March 16,1987 (20 days a f t e r r e a r i n g was i n i t i a t e d on F e b r u a r y 25,1987) and f i n i s h e d on March 23,1987. The two P. t e r m i n a l i s a d u l t s emerged 23 days a f t e r M. g e n t i l i s had c o m p l e t e d i t s emergence ( F i g . 17). As a r e s u l t of i n c u b a t i o n of i n f e s t e d l e a d e r s a t room temperature emergence of the w e e v i l s was a c c e l e r a t e d compared t o t h a t i n the f i e l d but the number of a c c u m u l a t e d h e a t i n g days has not been c a l c u l a t e d . Many p a r a s i t o i d s were a l s o r e a r e d (Table V I I I ) . Samples were dominated by the e c t o p a r a s i t o i d R h o p a l i c h u s  p u l c h r i p e n n i s C r a w f o r d (HYMENOPTERA: P t e r o m a l i d a e ) and t h e e n d o p a r a s i t o i d A l l o d o r u s sp. (HYMENOPTERA: B r a c o n i d a e ) , and t o a l e s s e r e x t e n t by two s p e c i e s of e u r y t o m i d s . F i g u r e 17 shows the emergence p a t t e r n s of the p a r a s i t o i d s . A l l s p e c i e s , e xcept the e u r y t o m i d s , s t a r t e d emerging on March 17,1987, a f t e r 21 days a t 20°C. A l l o d o r u s sp. f i n i s h e d i t s emergence w i t h i n 8 days but R. p u l c h r i p e n n i s and the e u r y t o m i d s emerged u n t i l A p r i l 10,1987. D e l o m e r i s t a  j a p o n i c a Cushman (HYMENOPTERA: Ichneumonidae) was a l s o p r e s e n t on the s i t e s . 52 Table VIII. Numbers of terminal weevils and parasitoids reared from leaders of lodgepole pine c o l l e c t e d from stands on the Palmer Lake Road. Rlske Creek, near Williams Lake, between February 25 end Ap r i l 2,1987. Locat ton Number of leaders reared Number P. terminal is of specimens reared M. q e n t i l i s Parasitoids 15km l e f t 52 1+10 2 1 6 2 15km r i g h t 50 1*10 2 - 33 3 21km r i g h t 90 1-M0 2 1 14 11 1 number of leaders set up for mass rearing 2 number of leaders reared i n d i v i d u a l l y F i g u r e 17 Emergence of l e a d e r w e e v i l s and t h e i r p a r a s i t o i d s from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d a t R i s k e c r e e k , near W i l l i a m s L a k e , B.C. i n February,1987. 54 3.3. Summer a c t i v i t i e s i n P e n t i c t o n i n 1987 3.3.1. D i s s e c t i o n s of a t t a c k e d t e r m i n a l s The r e s e a r c h was c o n t i n u e d i n P e n t i c t o n between May 4,1987 and Sept. 30,1987. D u r i n g t h i s study a t o t a l of 1046 i n f e s t e d t e r m i n a l s showing symptoms of w e e v i l a t t a c k were c o l l e c t e d on 12 s i t e s a t E l l i s c r e e k ( F i g . 13) and on two s i t e s a l o n g the B i g White Road. Of the 1046 l e a d e r s c o l l e c t e d , 309 were d i s s e c t e d and 737 were s e t up f o r i n d i v i d u a l r e a r i n g (Table I X ) . R e s u l t s of the d i s s e c t i o n s a r e summarized i n Ta b l e X. These d i s s e c t i o n s showed t h a t M. g e n t i l i s i s p r e s e n t a l s o i n the s o u t h e r n I n t e r i o r of B r i t i s h C o l u m b i a . I t was not known which w e e v i l s p e c i e s l e f t the f o u r emergence h o l e s , t h e r e f o r e a d u l t m o r t a l i t y c o u l d not be d e t e r m i n e d . However, i t was noted t h a t one t e r m i n a l c o l l e c t e d on s i t e 120 near Kelowna c o n t a i n e d seven l i v e and one dead a d u l t and one l i v e l a r v a of M. g e n t i l i s . T h i s i n d i c a t e s t h a t h i g h numbers of M. g e n t i l i s a r e a b l e t o s u r v i v e i n a s i n g l e t e r m i n a l which can l e a d t o h i g h w e e v i l a t t a c k i n c i d e n c e . When i n d i v i d u a l i n c u b a t i o n of i n f e s t e d t e r m i n a l s was comp l e t e d a t the end of J u l y , r e a r e d l e a d e r s were d i s s e c t e d t o check the s u c c e s s of r e a r i n g . In the 737 d i s s e c t e d l e a d e r s 11 dead P. t e r m i n a l i s a d u l t s , 17 l i v e 55 T a b l e IX. Summary t a b l e of l e a d e r s c o l l e c t e d f o r d i s s e c t i o n and r e a r i n g i n P e n t i c t o n and Kelowna i n May,1987 Number of l e a d e r s c o l l e c t e d S i t e f o r d i s s e c t i o n f o r r e a r i n g 6 5 -10 - 15 32 24 51 34 10 17 36 30 61 71 40 82 414 29 90 418 n o r t h 40 62 418 s o u t h 22 47 421 1 1 23 428 46 1 03 114 (Kelowna) 19 47 120 (Kelowna) 15 49 road 18 70 s l a s h - 20 T o t a l 309 737 Table X. Insect l i r e stages found during d i s s e c t i o n of damaged lodqepole pine leaders c o l l e c t e d In Penticton and Kelowna in May,1987 Weevil l i f e stages P a r a s i t o i d Emergonce S i t e adult pupae 1arvae mortality adult pupae larvae _ holes L D L D L D 1 t W Pa 6 - - - - - 6 75. 0 - - 1 2 -32 - - - - 10 18 64, 3 - - 3 - -34 - - - - 4 4 44 . 4 - - 1 1 -3S - - - - 16 19 54, 3 - 1 2 - -71 - - - - 18 15 45. .5 - - - -4 14 - IP - - 5 16 66. .7 - 1 - 2 -4 18 north - - - - 8 18 60. .0 - 3 1 4 -4 18 south - 3P - - 3 11 50. 0 - 2 - 5 2 421 - - - - 5 6 46. 1 - - - 2 -428 - IP - - 9 35 77. 7 - 3 12 - -114 Kelowna - 2P - 1 8 24 68. 6 2 1 - - -120 Kelowna 1M t - 3 12 48. 0 - 1 - 1 1 road - - - 1 2 9 75. 0 - 2 - 1 2 Total 7M 7P 1 2 91 195 60 .5 2 15 22 IB 5 1M 1 L • 11 vs, 0 • dead. P " P- terminal 13, M - M. a e n t I l l s 2 w « weevil. Pa • p a r a s i t o i d 57 C y l i n d r o c o p t u r u s sp. l a r v a e , 768 dead w e e v i l l a r v a e , 4 dead w e e v i l pupae, 17 dead p a r a s i t o i d a d u l t s and 23 l i v e and 17 dead p a r a s i t o i d pupae were found. W e e v i l l a r v a l m o r t a l i t y was 75% which i s much h i g h e r than t h a t found i n the May,1987 d i s s e c t i o n ( T a b l e X ) . T h i s h i g h l a r v a l m o r t a l i t y might be the r e s u l t of h i g h t e m p e r a t u r e s and low a i r h u m i d i t y i n the l a b o r a t o r y which l e d t o r a p i d d e s s i c a t i o n of the l e a d e r s . The room t e m p e r a t u r e a v e r a g e d 23^C d u r i n g the r e a r i n g . H u m i d i t y of the a i r was not measured. I t was i n t e r e s t i n g t o note t h a t 23 l i v e p a r a s i t o i d pupae were s t i l l i n the l e a d e r s . T h i s r a i s e s the f o l l o w i n g q u e s t i o n s , "Do t h e s e p a r a s i t o i d s o v e r w i n t e r f o r a second year or do they emerge l a t e r i n the c u r r e n t year? I f they emerge l a t e r what a r e they g o i n g t o f e e d on?". These p a r a s i t o i d pupae were kept i n p e t r i d i s h e s a t room temp e r a t u r e but by t h e end of September o n l y t h r e e R. p u l c h r i p e n n i s , one Mesopolobus sp. and two D. j a p o n i c a emerged. The r e s t of the pupae d e h y d r a t e d and d i e d . Weekly d i s s e c t i o n s of c u r r e n t y e a r ' s a t t a c k s s t a r t e d on June 5,1987 which showed the development of the w e e v i l s d u r i n g the summer. T a b l e XI shows the r e s u l t s of t h e s e d i s s e c t i o n s . As a r e s u l t of the w e e v i l complex the r e l a t i v e numbers of l a r v a e of the d i f f e r e n t w e e v i l s p e c i e s i s unknown but t h i s d a t a s t i l l p r o v i d e s some i n s i g h t i n t o the 58 Table XI. Summary table of the result s of weekly d i s s e c t i o n of lodgepole pine leaders attacked at E l l i s creek during the summer of 1987 Number of Number of Eggs* Larvae Pupae Adults Date feeding o v i p o s i t i o n punctures s 1 tes L D P L D P L D P L D June 5 6 44 27 c c c 10 6 90 48 43 - 7 - - - - -16 6 84 76 55 6 14 - - - - -23 6 90 48 19 10 17 2 30 6 46 39 3 11 19 6 July 8 10 129 81 33 18 24 6 14 10 ? d 7 d 1 4 30 7 22 10 10 47 10 29 10 12 18 22 2 Aug. 5 10 7 45 6 1 12 10 21 9 1 19 10 17 2 26 10 16 2 Sept.2 10 19 3 3 9 10 17 2 5 17 10 22 6 3 1 - -23 10 26 2 4 1 - -30 10 17 3 8 - - 1 a number of leaders c o l l e c t e d b L=live. D=dead, P=parasitized c number of eggs was not counted d i t was not possible to i d e n t i f y the feeding punctures after July 8., 1'987. 59 development of the w e e v i l s . E x t e n s i v e r e s i n f l o w and l a r v a l m i n i n g made c o u n t i n g the number of f e e d i n g p u n c t u r e s and o v i p o s i t i o n s i t e s i m p o s s i b l e a f t e r J u l y 8,1987. The l a s t l i v e egg was found on J u l y 14,1987. I t was not p o s s i b l e t o determine how many l i v e eggs were a t t a c k e d by egg p a r a s i t o i d s . An egg was c o n s i d e r e d dead i f i t appeared g r e y i s h or the c h o r i o n was w r i n k l e d . The mean number of l a r v a e per l e a d e r (160 l e a d e r s d i s s e c t e d ) was 3.4 (S.D.=2.79, range=1-17). S t a r k and Wood (1964) d i s s e c t e d l e a d e r s (n=305) c o l l e c t e d i n l o d g e p o l e p i n e stands i n f e s t e d by P. t e r m i n a l i s o n l y . They found 4.2 l a r v a e per l e a d e r s (range=1 - 19) . Headcapsule w i d t h s of dead and l i v e l a r v a e (n=115 and 376, r e s p e c t i v e l y ) o b t a i n e d from t h e d i s s e c t i o n s were measured. The frequency d i s t r i b u t i o n s a r e p r e s e n t e d i n F i g s . 18-19. S t a r k and Wood (1964) measured the h e a d c a p s u l e w i d t h s of the fo u r i n s t a r s of P. t e r m i n a l i s and the arrows ( F i g s . 18-19) d e s i g n a t e the mean h e a d c a p s u l e w i d t h of each i n s t a r measured by them. The s l i g h t d i f f e r e n c e s i n the measurements i n the two e xperiments a l s o i n d i c a t e the pr e s e n c e of the t e r m i n a l w e e v i l complex i n P e n t i c t o n . 60 12 16 20 24 28 32 36 40 44 46 52 micrometer divisions F i g u r e 18 Frequency d i s t r i b u t i o n of h e a d c a p s u l e w i d t h s of dead w e e v i l l a r v a e r e s u l t e d from weekly d i s s e c t i o n s of i n f e s t e d t e r m i n a l s h o o t s ( 1 m i c r o m e t e r d i v i s i o n = 0.025mm). 61 12 16 20 24 28 32 36 40 44 48 52 micrometer divisions F i g u r e 19 Frequency d i s t r i b u t i o n of h e a d c a p s u l e w i d t h s of l i v e w e e v i l l a r v a e r e s u l t e d from weekly d i s s e c t i o n s of i n f e s t e d t e r m i n a l s h o o t s ( 1 mi c r o m e t e r d i v i s i o n - 0.025mm). 62 3.3.2. I n d i v i d u a l i n c u b a t i o n of i n f e s t e d t e r m i n a l s . Numbers of a c c u m u l a t e d degree-days were c a l c u l a t e d (Appendix I I I ) . S i n c e the lower d e v e l o p m e n t a l t h r e s h o l d s of the w e e v i l s are unknown the e x a c t number of a c c u m u l a t e d degree-days cannot be d e t e r m i n e d . T h e r e f o r e a l l emergence d a t e s t h r o u g h o u t t h i s s t u d y r e p r e s e n t emergence d a t e s i n the l a b o r a t o r y . I n d i v i d u a l r e a r i n g , which s t a r t e d between May 7-11,1987 , r e v e a l e d t h a t , b e s i d e P. t e r m i n a l i s and M. g e n t i l i s , a t h i r d t e r m i n a l w e e v i l s p e c i e s from th e genus C y l i n d r o c o p t u r u s i s a l s o p r e s e n t i n the P e n t i c t o n a r e a ( T a b l e X I I ) . The f i r s t w e e v i l s p e c i e s t o emerge was M. q e n t i l i s which s t a r t e d emerging on May 12,1987 and f i n i s h e d seven days l a t e r , on May 19,1987 ( F i g . 2 0 ) . The r a p i d emergence of M. g e n t i l i s was s i m i l a r t o t h a t r e c o r d e d from th e e a r l i e r W i l l i a m s Lake r e a r i n g s ( F i g . 17). C y l i n d r o c o p t u r u s sp. emerged between June 9,1987 and J u l y 9,1987 ( F i g . 2 0 ) . A l t h o u g h the p a t t e r n of the emergence was s c a t t e r e d i t s u g g e s t s t h a t the m a j o r i t y of the a d u l t s emerge i n the f i r s t two weeks of J u l y or l a t e r a l l o w i n g f o r e l e v a t e d r e a r i n g t e m p e r a t u r e s . On June 3,1987 (30 days i n r e a r i n g ) P. t e r m i n a l i s s t a r t e d t o emerge and reached i t s peak on June 7,1987 (34 days i n r e a r i n g ) ( F i g . 2 1 ) . Emergence c o n t i n u e d u n t i l J u l y 7,1987. 63 Table XII. Summary table of insects reared frore weevil infested lodgepole pine leaders c o l l e c t e d on Cartni Road at E l l i s Creek and on the Big Wnite Road in May,1987 Number of Si t e leaders P. terminal is M. gent 111s Cylindrocopturus sp. Parasitoids reared 10 15 - 1 - 2 32 51 6 5 6 9 34 17 - 3 - 4 36 61 8 - - 12 71 82 4 1 3 6 4 14 90 8 2 - 13 418 north 62 3 5 - 13 418 south 47 - - - 13 421 23 - 5 - 8 428 103 6 2 - 14 1 14 47 10 - - 10 120 49 7 7 - 15 road 70 7 - - 7 slash 20 - - 3 1 Total 737 59 24 12 127 10 9 -8 -7 -6 -5 -4 -3 -1 -' I i »» • I i i i » I i i i i I i i i i'|'i i i i | i i i i | i i i DAYS IN REARING AT 20*C CYUNDROCOPTURUS SP I ' ' 1 ' 4 9 14 19 24 29 34 39 44 49 54 59 64 69 V~7\ MAGDAUS GENTILIS FIGURE 20: Temporal emergence of M a g d a l i s q e n t i l i s and C y l i n d r o c o p t u r u s sp. from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d i n the s o u t h e r n i n t e r i o r of B r i t i s h Columbia i n May,1987. R e a r i n g s t a r t e d on May 7,1987. o V) C 0 2 15 10 0 ™ ™ H J ^ ^ t e r m i n a l i s R. p u l c h r i p e n n i s D. t e r e b r a n s E u r y t o m a s p p . D D n - n / A l l o d o r u s s p . U_U_D D LMesopolobus s p . D. j a p o n i c a 10 15 1 i 1 1 1 1 i 1 1 1 1 i 1 1 1 1 i 1 1 1 1 i 1 1 1 1 i * 1 1 1 i • 1 • 1 i 2 0 2 5 3 0 3 5 4 0 4 5 5 0 5 5 H y s s o p u s s p . D a y s i n R e a r i n g a t 2 0 °C F i g u r e 21 Temporal emergence of P i s s o d e s t e r m i n a l i s and p a r a s i t o i d s from i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d i n the s o u t h e r n i n t e r i o r of B r i t i s h Columbia i n May,1987. R e a r i n g s t a r t e d on May 7,1987. 66 S i n c e females o v i p o s i t i n t o the e l o n g a t i n g t e r m i n a l s h o o t , l e a d e r e l o n g a t i o n i s c r i t i c a l f o r s u c c e s s f u l o v i p o s i t i o n . Measurements were t a k e n from 25 e l o n g a t i n g l o d g e p o l e p i n e l e a d e r s . E l o n g a t i o n began a t the b e g i n n i n g of May,1987. A l t h o u g h a s l i g h t i n c r e a s e i n l e a d e r l e n g t h o c c u r r e d i n mid-August, t e r m i n a l growth was p r a c t i c a l l y completed by e a r l y August,1987 ( F i g . 22) (Appendix I V ) . The p e r i o d of r a p i d l e a d e r e l o n g a t i o n i s the p e r i o d when the e p i d e r m i s i s the most v u l n e r a b l e t o i n s e c t f e e d i n g . F i g u r e 20 shows the emergence of the t e r m i n a l w e e v i l s i n r e l a t i o n t o t i p e l o n g a t i o n . I t i s c l e a r t h a t w e e v i l emergence c o i n c i d e s w i t h l e a d e r e l o n g a t i o n . 3.3.3. O b s e r v a t i o n s on the b i o l o g i e s of P. t e r m i n a l i s 3.3.3.1. L o n g e v i t y and f e c u n d i t y of P i ssodes  t e r m i n a l i s L o n g e v i t y and f e c u n d i t y of 10 female P. t e r m i n a l i s were d e t e r m i n e d . Average l o n g e v i t y of females was 112.8 days; one female l i v e d f o r 226 days ( T a b l e X I I I ) . I t was observed t h a t female P. t e r m i n a l i s a r e a b l e t o l a y eggs as soon as 2 days f o l l o w i n g emergence and t h e r e f o r e do not r e q u i r e l o n g e r m a t u r a t i o n f e e d i n g . In t h i s s t u d y , the p r e o v i p o s i t i o n p e r i o d a veraged 10.1 days (S.D.=+6.8, range=2-22 d a y s ) . O v i p o s i t i o n began on day 2 and t h e r e a f t e r i n c r e a s e d u n t i l day 21, d e c l i n e d t o about t w o - t h i r d i t s peak o v e r w i n t e r i n g LARVAE/pupae/adults d i s p e r s a l t o new t e r m i n a l s c a d u l t emergence 1 1 o v i p o s i t i o n I 1 o v e r w i n t e r i n g LARVAE/pupae/adults i O * l O O r -Cn c 5 3 B 50 * o 0 « Bud Leader E l o n g a t i o n of the t e r m i n a l of P. c o n t o r t a . Jan Feb Mar Apr May Jun J u l Aug Sep. Oct Nov, Dec F i g u r e 22: Schematic diagram of t e r m i n a l w e e v i l emergence and o v i p o s i t i o n i n r e l a t i o n t o t i p e l o n g a t i o n of P i n u s c o n t o r t a . Table x111 . Longevity and o v i p o s i t i o n c h a r a c t e r i s t i c s of 10 female P i ssodes terminal 1s reared at 20°C+2°C at UBC. C h a r a c t e r i s t i c s Mean S.D. Range Longevity (days) 112.8 74.9 32-226 Pre o v i p o s i t i o n period (days) 10.1 6.8 2-22 Total eggs per female 115.0 67.5 9-216 Eggs per o v i p o s i t i o n s i t e 0.94 - 1 0-2 Eggs per female per day 2 1.57 1.34 0.39-4.62 1 S.D. has not been c a l c u l a t e d 2 from f i r s t day of o v i p o s i t i o n to death of female 69 v a l u e on day 28, and then i n c r e a s e d u n t i l day 35 ( F i g . 2 3 ) . O v i p o s i t i o n s t a r t e d t o decrease s h a r p l y a f t e r day 49, a l t h o u g h o n l y t h r e e females had d i e d . F e c u n d i t y averaged 115 eggs per female (S.D.=+67.5, range=9-216). A t o t a l of 1246 o v i p o s i t i o n s i t e s were counted and 1150 eggs were found g i v i n g a mean number of eggs per o v i p o s i t i o n s i t e of 0.94. Females most o f t e n l a i d one egg per p i t (91.8% of p i t s ) , but sometimes they o v i p o s i t e d two eggs (0.5%) or none ( 7 . 7 % ) . F i v e eggs were l a i d d i r e c t l y on the bark s u r f a c e ( 0 . 4 % ) . These r e s u l t s a r e i n g e n e r a l agreement w i t h o b s e r v a t i o n s t h a t g e n e r a l l y o n l y one egg i s d e p o s i t e d i n each p i t ( D r o u i n e t a l . ; S t a r k and Wood 1964). Females l a i d 1.57 eggs per day on an average b a s i s (S.D.=1.34, range=0.39-4.62) ( T a b l e X I I I ) . 3.3.3.2. D u r a t i o n of egg stage and p u p a l s t a g e s of P. t e r m i n a l i s T e r m i n a l s w i t h o v i p o s i t i o n s i t e s from the l o n g e v i t y and f e c u n d i t y e x p e r i m e n t were e v a l u a t e d f o r the d u r a t i o n of the egg s t a g e of P. t e r m i n a l i s . O b s e r v a t i o n of 54 eggs r e v e a l e d t h a t t h e average d u r a t i o n of the egg s t a g e i s 8 days (S.D.=+0.98, r a n g e = 5 ~ l 8 ) . T h i s i s i n agreement w i t h the f i n d i n g s of St e v e n s and Knopf (1974) who r e p o r t e d t h a t eggs of P. t e r m i n a l i s hatched w i t h i n two weeks. Two eggs ha t c h e d 5 days a f t e r o v i p o s i t i o n and 9 took 18 days t o h a t c h . Head c a p s u l e s of 36 f i r s t i n s t a r l a r v a e were 7 21 35 49 63 77 91 105 119 133 147 161 175 189 203 217 231 days • egg/female/week + number of females F i g u r e 23: S u r v i v a l s h i p and f e c u n d i t y of t e n P i s s o d e s t e r m i n a l i s f e m a l e s . o 71 measured i n d i c a t i n g t h a t t he average head c a p s u l e w i d t h was 0.371mm (S.D.=+0.022, range=0.300-0.400). O b s e r v a t i o n s on the pu p a l s t a d i u m of 20 P. t e r m i n a l i s pupae showed t h a t on average i t t a k e s 15 days (S.D.=+2.5, range=11-20) from p u p a t i o n t o a d u l t emergence a t 20°C. 3. 3 . 4 . O b s e r v a t i o n s on the b i o l o g y of M. g e n t i l i s and C y l i n d r o c o p t u r u s sp. M. g e n t i 1 i s i s a b l a c k w e e v i l ( F i g . 2 4 ) . Measurements showed t h a t the average l e n g t h of 7 males was 3.94mm (S.D.=+0.13) whereas the average l e n g t h of 13 females was 4.92mm (S.D.=+0.23). A d u l t s a r e d e f o l i a t o r s ( F i g . 2 5 ) . F e l l i n (1973) s t a t e d t h a t " t h e r e was no i n d i c a t i o n t h a t a d u l t s of M. q e n t i l i s o v i p o s i t nor t h a t l a r v a e f e e d i n or on the s h o o t s or any o t h e r p o r t i o n s of the s t a n d i n g t r e e " . In the p r e s e n t s t u d y , however, M. g e n t i l i s a d u l t s were r e a r e d from c u r r e n t y e a r ' s i n f e s t e d l o d g e p o l e p i n e l e a d e r s ( T a b l e s V I , V I I I and X I I ) . F u r t h e r m o r e , o v i p o s i t i o n by M. g e n t i l i s was a l s o o b s e r v e d . O v i p o s i t i o n a l s i t e s were always a t the base of the n e e d l e s ( F i g . 2 6 ) . P. t e r m i n a l i s never o v i p o s i t e d a t the base of the n e e d l e s ( F i g . 2 7 ) . I t i s not known i f o v i p o s i t i o n i s r e s t r i c t e d t o the t e r m i n a l or a l s o o c c u r s on o t h e r p o r t i o n s of the crown. Plumb (1950) and Keen (1952) r e p o r t e d t h a t l a r v a e of c e r t a i n M a g d a l i s s p e c i e s a r e t w i g m i n e r s . F u r n i s s and C a r o l i n (1977) a l s o r e c o r d e d F i g u r e 24 M a q d a l i s g e n t i l i s a d u l t on e l o n g a t i n g l o d g e p o l e p i n e t e r m i n a l s h o o t . 73 F i g u r e 25 One h e a l t h y l o d g e p o l e p i n e n e e d l e and 11 n e e d l e s a f f e c t e d by the f e e d i n g of a d u l t M. g e n t i l i s . The a p i c a l p a r t of the n e e d l e s have d e s s i c a t e d , d i s c o l o r e d and c u r l e d up a l o n g the l o n g i t u d i n a l a x i s . A p i c a l p o r t i o n of t h e l a s t t h r e e n e e d l e s had been broken e i t h e r by wind or r a i n . 74 F i g u r e 26 . O v i p o s i t i o n s i t e made by M. q e n t i l i s i n t o the c u r r e n t y e a r ' s t e r m i n a l s h o o t . O v i p o s i t i o n was made a t the v e r y base of the n e e d l e . F i g u r e 27 O v i p o s i t i o n s i t e made by P. t e r m i n a l i s female i n t o the e l o n g a t i n g t e r m i n a l . 76 l a r v a e of M. g e n t i l i s m i n i n g branches of ponderosa p i n e and J e f f r e y p i n e . A l t h o u g h l a r v a e of P . t e r m i n a l i s and M. q e n t i l i s a r e i n d i s t i n g u i s h a b l e , l e a d e r s damaged by t h e two w e e v i l s p e c i e s can be s e p a r a t e d examining the f o l l o w i n g c h a r a c t e r i s t i c s . L a r v a l b o r i n g of M. q e n t i l i s i s powdery and f i n e - g r a i n e d , whereas P. t e r m i n a l i s l a r v a e produce shredded b o r i n g s . L a r v a e of M. g e n t i l i s t e n d t o work i n t o t h e wood deeper t h a n do P. t e r m i n a l i s l a r v a e . F u r t h e r m o r e , p u p a l chambers of M. g e n t i l i s a r e smooth, whereas th o s e of P. t e r m i n a l i s a r e l i n e d w i t h some shredded wood (Keen 1952; F u r n i s s and C a r o l i n 1977; Hulme, M.A.2^ p e r s o n n e l c o m m u n i c a t i o n ) . L i k e a d u l t s of M. g e n t i l i s , C y l i n d r o c o p t u r u s s p. a d u l t s ( F i g . 28) a l s o f e e d on n e e d l e s ( F i g . 2 9 ) . L a r v a e of b o t h M. g e n t i 1 i s and C y l i n d r o c o p t u r u s sp. mined beneath the bark of the t e r m i n a l f e e d i n g i n the phloem r e g i o n ( F i g s . 30-31), but they never e n t e r e d the p i t h . L a r v a e of t h e s e two w e e v i l s p e c i e s pupate between the bark and t h e wood of the t e r m i n a l . L a r v a e of n e i t h e r s p e c i e s c o n s t r u c t cocoons. B r e e d i n g by more than one of the t e r m i n a l w e e v i l s p e c i e s i n the same t e r m i n a l was never o b s e r v e d . 2 R e s e a r c h s c i e n t i s t , P a c i f i c F o r e s t r y C e n t r e , V i c t o r i a , B.C. 77 F i g u r e 28 C y l i n d r o c o p t u r u s sp. a d u l t s . F i g u r e 29 F e e d i n g p u n c t u r e s on l o d g e p o l e p i n e n e e d l e s caused by C y l i n d r o c o p t u r u s sp. a d u l t s . 79 F i g u r e 30 Lodgepole p i n e t e r m i n a l damaged by Magdal  q e n t i l i s l a r v a e . 80 F i g u r e 31 Lodgepole p i n e t e r m i n a l damaged by l a r v a e C y l i n d r o c o p t u r u s sp. 81 A s m a l l b l a c k b e e t l e , P i t y o p h t h o r u s prob. b o y c e i Sw. (COLEOPTERA: S c o l y t i d a e ) was a l s o found a s s o c i a t e d w i t h the t e r m i n a l w e e v i l s d u r i n g t h i s s t u d y . I t was o b s e r v e d t h a t the m a j o r i t y of P. p r o b . b o y c e i u t i l i z e d untouched p o r t i o n s of p r e v i o u s l y a t t a c k e d l e a d e r s . However, t h i s b e e t l e a l s o a t t a c k e d h e a l t h y t e r m i n a l s h o o t s . 3.3.5. P a r a s i t o i d s a s s o c i a t e d w i t h the w e e v i l s A l a r g e number of p a r a s i t o i d s was a l s o r e a r e d . A l l samples were dominated by the p t e r o m a l i d R. p u l c h r i p e n n i s . Two s p e c i e s of Eurytoma c o l l e c t i v e l y r a n ked second i n abundance. The f i r s t p a r a s i t o i d s t o emerge were two ichneumonids P o l i c h o m i t u s t e r e b r a n s n u b i l i p e n n i s ( V i e r e c k ) , and D e l o m e r i s t a j a p o n i c a Cushman and the b r a c o n i d A l l o d o r u s sp. ( F i g . 2 1 ) . Emergence p a t t e r n s show t h a t , a l t h o u g h the two s p e c i e s of Eurytoma s t a r t e d t o emerge among the f i r s t p a r a s i t o i d s , they had an extended emergence d u r i n g the r e a r i n g . A l l o t h e r e a r l y emerging s p e c i e s completed emergence w i t h i n 7-14 days and were f o l l o w e d by the p t e r o m a l i d s R. p u l c h r i p e n n i s and Mesopolobus sp. and the e u l o p h i d Hyssopus sp. R e a r i n g p a r a s i t o i d s i n the l a b o r a t o r y a c c e l e r a t e d emergence r e l a t i v e t o the f i e l d s i t u a t i o n , presumably owing t o the h i g h e r t e m p e r a t u r e s i n the l a b o r a t o r y . Minimum 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 of the p a r a s i t o i d s a r e unknown 82 and t h u s the number of accumulated degree-days c o u l d not be d e t e r m i n e d . Emergence p a t t e r n s of the p a r a s i t o i d s c o i n c i d e d w i t h those of P. t e r m i n a l i s and C y l i n d r o c o p t u r u s sp. ( F i g s . 20-21). Emergence of M. g e n t i l i s p receeded t h a t of t h e p a r a s i t o i d s but p r o b a b l y by the time newly emerged a d u l t w e e v i l s f i n i s h e d m a t u r a t i o n f e e d i n g and o v i p o s i t i o n , p a r a s i t o i d s were a l s o ready t o a t t a c k . F i g s . 20 and 21 show t h a t phenology of the w e e v i l s i s c l o s e l y f o l l o w e d by t h a t of the p a r a s i t o i d s . C l o s e r e l a t i o n s h i p s between h o s t and p a r a s i t o i d p h e n o l o g i e s i s one of the c r i t e r i a f o r s u c c e s s f u l b i o l o g i c a l c o n t r o l of the p e s t ( E h l e r and Andres 1983; Murdoch et a l . 1985). I n c u b a t i o n of i n f e s t e d l o d g e p o l e p i n e l e a d e r s c o l l e c t e d t h r o u g h o u t B r i t i s h Columbia f o r t h e l a s t two y e a r s has made i t p o s s i b l e t o summarize d a t a on p a r a s i t o i d s of the t e r m i n a l w e e v i l s (Table X I V ) . P a r a s i t o i d s b e l o n g t o 6 f a m i l i e s i n the o r d e r Hymenoptera. As a r e s u l t of the t e r m i n a l w e e v i l complex, c o r r e c t a s s o c i a t i o n s between h o s t s and p a r a s i t o i d s c annot be e n s u r e d . In the c o u r s e of t h i s s t u d y n e i t h e r p r e d a t o r s nor entomophagous f u n g i were found i n a s s o c i a t i o n w i t h any of the w e e v i l s . 83 Table XIV. Hymenoterans found associated with the terminal weevils in B r i t i s h Columbia, 19B6-1987. Fam i1y Genus/Spec i es Locat1 on Relat ive abundance Type of paras i to i d Eu1oph i dae Eurytom i dae Coeloides ruf ovar jepatus (Prov.) Hyssopus sp. Eurytoma spp. Braconidae A11odorus sp. Kelowna Pent icton Prince George Williams Lake Kelowna Kelowna Pent icton Kami oops Kelowna Merritt Penticton Prince George Will lams Lake Ichneumonidae P o l i chom i tus Penticton terebrans nubi1ipennis Williams Lake (Ratzeburg) Delomer i sta japonica Penticton Cushman Pteromalidae Rhopa1i chus Kamloops pu1chr ipenni s Cwfd. Kelowna Merritt Penticton Prince George Will lams Lake Kamloops Merr1tt PentIcton Kami oops Will lams Lake Mesopc1obus sp. Scellonidae Te1enomus sp. Endo Ecto Endo Ecto Ecto Ecto Ecto unknown Endo very abundant frequent in samples rare 84 3.3.6. A d u l t p a r a s i t o i d f o o d p l a n t s P o l l e n or honeydew a r e e s s e n t i a l f o r a d u l t p a r a s i t o i d n u t r i t i o n (Hagen and H a l e 1974) and r e p r o d u c t i o n (DeBach 1974). DeBach (1974) i n d i c a t e d t h a t e x p e r i m e n t s of R u s s i a n s c i e n t i s t s have shown the need f o r supplementary food f o r a d u l t p a r a s i t o i d s . Hagen and Hale (1974) suggested t h a t the a p p l i c a t i o n of s u p p l e m e n t a r y food where non-prey f o o d i s l a c k i n g can be used t o a t t r a c t or r e t a i n n a t u r a l enemies. DeBach (1974) r e p o r t e d t h a t i n o r d e r t o p r o v i d e f o o d f o r a d u l t p a r a s i t e s of the c o d l i n g moth, L a s p e y r e s i a  p omonella (L.) (LEPIDOPTERA: O l e t h r e u t i d a e ) , n e c t a r -p r o d u c i n g p l a n t s s h o u l d be p l a n t e d i n o r c h a r d s . I t was o b s e r v e d t h a t one of the s i t e s ( s i t e 418) had p a r t i c u l a r l y h i g h numbers and a wide v a r i e t y of p a r a s i t o i d s . T h i s s i t e s u p p o r t e d an abundance of f i r e w e e d , E p i l o b i u m  a n g u s t i f o l i u m L. ( F i g . 32) and one s p e c i e s of l u p i n , L u p i n u s sp. ( F i g . 3 3 ) . N e i t h e r of t h e s e two weed s p e c i e s grew on any of t h e o t h e r s t u d y s i t e s . I t was n o t i c e d t h a t a d u l t p a r a s i t o i d s f e d on p o l l e n of t h e s e f l o w e r i n g weeds. T h i s o b s e r v a t i o n s u g g e s t s t h a t p o l l e n p l a y s an i m p o r t a n t r o l e i n a d u l t p a r a s i t o i d n u t r i t i o n (Hagen and H a l e 1974). 85 F i g u r e 32 F i r e w e e d , E p i l o b i u m a n g u s t i f o l i u m L. i n young l o d g e p o l e p i n e s t a n d . 86 F i g u r e 33 L u p i n , L u p i n u s sp. growing i n young l o d g e p o l e p i n e s t a n d . 87 3.3.7. W e e v i l a t t a c k i n c i d e n c e and a t t a c k h i s t o r y survey A w e e v i l a t t a c k i n c i d e n c e and a t t a c k h i s t o r y s u r v e y was conducted a t E l l i s c r e e k on a 33ha l o d g e p o l e p i n e s t a n d . A t o t a l of 1088 t r e e s were sampled. R e s u l t s of the s u r v e y a r e p r e s e n t e d i n T a b l e XV. W e e v i l a t t a c k s t a r t e d i n 1983 and the number of i n f e s t e d t r e e s was 13 t i m e s h i g h e r i n 1984 than i n the p r e v i o u s y e a r . The i n f e s t a t i o n r a t e d o u b l e d i n 1985 compared t o 1984. A t t a c k i n c i d e n c e d e c l i n e d i n 1986. T h i s d e c l i n e was p r o b a b l y a r e s u l t of the u n u s u a l l y c o l d w i n t e r of 1985 ( F i g . 3 4 ) . The m i l d w i n t e r of 1986 ( F i g . 35) may have r e s u l t e d i n an i n c r e a s e i n a d u l t and l a r v a l s u r v i v a l r a t e s as t h e r e was an a l m o s t 4 - f o l d i n c r e a s e i n the number of w e e v i l l e d t r e e s i n 1987. Of the 1088 t r e e s sampled, 21 t r e e s (1.9%) were a t t a c k e d t w i c e . No t r e e was found w i t h more than two a t t a c k s . 88 T a b l e XV. R e s u l t s of the w e e v i l a t t a c k i n c i d e n c e and a t t a c k h i s t o r y s u r v e y conducted on a 33ha, 1 4 - y e a r - o l d spaced l o d g e p o l e p i n e s t a n d a t E l l i s Creek i n J u l y , 1 9 8 7 ( n = l 0 8 8 ) . Tree c o n d i t i o n Number of P e r c e n t a g e of t r e e s t o t a l H e a l t h y (never a t t a c k e d ) 842 78.9 A t t a c k e d i n 1983 2 0.2 A t t a c k e d i n 1984 26 2.4 A t t a c k e d i n 1985 53 4.9 A t t a c k e d i n 1986 36 3.3 A t t a c k e d i n 1987 129 1 1.9 T o t a l 1088 100.0 F i g u r e 34: 1985 and 30-year monthly minimum and maximum average t e m p e r a t u r e s a t the P e n t i c t o n a i r p o r t ( C o u r t e s y of Environment Canada). CO 1 1 1 1 1 Oct Nov D«c Jan Feb Mar Apr months 1986 average maximum + 30-year average maximum 1986 average minimum A 30-year average minimum F i g u r e 1986 and 30-year monthly minimum and maximum average t e m p e r a t u r e s a t the P e n t i c t o n a i r p o r t ( C o u r t e s y of Environment Canada 91 3.4 Management i m p l i c a t i o n s Emergence p a t t e r n s of the w e e v i l s showed t h a t t i m i n g i s i m p o r t a n t i n p l a n n i n g l e a d e r c l i p p i n g p r o j e c t s . E a r l y emergence of M. g e n t i l i s s u g g e s t s t h a t l e a d e r c l i p p i n g s h o u l d be c a r r i e d out by e a r l y s p r i n g . O b s e r v a t i o n s suggest t h a t the b e s t time f o r l e a d e r c l i p p i n g i s F e b r u a r y . Most of the a t t a c k e d l e a d e r s have changed c o l o r by t h i s time and r e c o g n i t i o n of i n f e s t e d t e r m i n a l s i s r e l a t i v e l y easy. E x p e r i e n c e has a l s o shown t h a t the t o p of the snow i s h a r d i n F e b r u a r y which f a c i l i t a t e s easy a c c e s s t o s t a n d s . In a d d i t i o n , the h e i g h t of the snow h e l p s a person r e a c h t h e t e r m i n a l s and i n many c a s e s bending the t r e e (which might l e a d t o breakage) can be a v o i d e d . As mentioned e a r l i e r , a wide v a r i e t y of p a r a s i t o i d s have been o b s e r v e d on f l o w e r s of weeds. F l o w e r i n g weeds i n l o d g e p o l e p i n e s t a n d s c o u l d p r o v i d e n e c t a r or p o l l e n t o p a r a s i t o i d s e n h a n c i n g n a t u r a l p a r a s i t o i d p o p u l a t i o n s a l r e a d y p r e s e n t i n the f i e l d . C u l t i v a t i n g f i r e w e e d , E. a n g u s t i f o l i u m , i s i m p r a c t i c a l because t h i s weed i s a s e r i o u s c o m p e t i t o r f o r m o i s t u r e and space, which c o u l d s u p p r e s s s m a l l e r t r e e s . However, c u l t i v a t i n g l u p i n , L u p i n u s sp., would be b e n e f i c i a l not o n l y f o r the p a r a s i t o i d s but a l s o f o r the t r e e s as l u p i n f i x e s n i t r o g e n enhancing the n i t r o g e n c o n t e n t of the s o i l . 92 3.5. Other s t r a t e g i e s f o r w e e v i l c o n t r o l Hulme e t a l . (1987) s u g g e s t e d new methods f o r the c o n t r o l of the S i t k a s p r u c e w e e v i l , P i s s o d e s s t r o b i ( P e c k ) . They suggested p u t t i n g i n f e s t e d S i t k a s p r u c e l e a d e r s i n t o c o n t a i n e r s w i t h a f i n e s c r e e n on the t o p w i t h openings 1.7x1.7mm wide. Emerging p a r a s i t o i d s and p r e d a t o r s a r e a b l e t o c r a w l through the s c r e e n but P. s t r o b i i s p r e v e n t e d from e s c a p i n g . A l t h o u g h t h i s method i s s i m p l e and i n e x p e n s i v e , i t i s not as a p p l i c a b l e f o r the c o n t r o l of t e r m i n a l w e e v i l s i n l o d g e p o l e p i n e s t a n d s because of the v a r i e t y of w e e v i l s p e c i e s and t h e i r p a r a s i t o i d s . The t h r e e w e e v i l s p e c i e s and the p a r a s i t o i d s v a r y i n s i z e w hich makes t h i s method i n a p p l i c a b l e . Hulme e t §_1. (1986) u t i l i z e d the r e l a t i v e l a c k of c o l d -h a r d i n e s s of P. s t r o b i t o k i l l t he w e e v i l and m a i n t a i n p a r a s i t o i d p o p u l a t i o n s . When i n f e s t e d t e r m i n a l s were c o o l e d t o -26°C, w e e v i l s of a l l d e v e l o p m e n t a l s t a g e s were k i l l e d damaging the p a r a s i t o i d s or p r e d a t o r s . T h i s method may be a p p l i c a b l e f o r the c o n t r o l of t e r m i n a l w e e v i l s on. l o d g e p o l e p i n e . However, the c o l d - h a r d i n e s s of P. t e r m i n a l i s , M. g e n t i l i s and C y l i n d r o c o p t u r u s sp. have y e t t o be det e r m i n e d . F u r t h e r e x p e r i m e n t s s h o u l d be c a r r i e d out t o add r e s s t h i s q u e s t i o n . 93 4. GENERAL DISCUSSION AND CONCLUSIONS The major f i n d i n g s of the p r e s e n t study can be summarized as f o l l o w s : a. A t e r m i n a l w e e v i l complex c o n s i s t i n g of t h r e e w e e v i l s p e c i e s , P i s s o d e s t e r m i n a l i s Hopping, M a q d a l i s g e n t i l i s LeConte and C y l i n d r o c o p t u r u s sp., i s p r e s e n t on l o d g e p o l e p i n e i n B r i t i s h Columbia. b. B e s i d e k i l l i n g e l o n g a t i n g t e r m i n a l s M. g e n t i l i s and C y l i n d r o c o p t u r u s sp. a d u l t s a l s o f e e d on n e e d l e s . c. B i o l o g y of P. t e r m i n a l i s d i f f e r s s l i g h t l y from t h a t of the two o t h e r w e e v i l s p e c i e s . T h i r d i n s t a r P. t e r m i n a l i s l a r v a e m i g r a t e i n t o the p i t h , whereas M. g e n t i l i s and C y l i n d r o c o p t u r u s sp. l a r v a e f e e d and pupate between the bark and the wood. d. M. g e n t i l i s emerges i n mid-May and i s f o l l o w e d by P. t e r m i n a l i s and C y l i n d r o c o p t u r u s sp. The l a t t e r two w e e v i l s emerge from e a r l y June t h r o u g h m i d - J u l y . e. The t e r m i n a l w e e v i l s have a p a r a s i t o i d complex i n B r i t i s h C o l u m b i a . f . The most w i d e l y d i s t r i b u t e d p a r a s i t o i d i s R h o p a l i c h u s  p u l c h r i p e n n i s Cwfd. Two s p e c i e s of Eurytoma c o l l e c t i v e l y rank second i n abundance. g. Emergence of t h e p a r a s i t o i d s c l o s e l y f o l l o w s t h a t of the w e e v i l s . 94 5. RECOMMENDATIONS The f o l l o w i n g recommendations a r e made based on t h e r e s u l t s of the s t u d y : a. E a r l y emergence of M. g e n t i l i s s u g g e s t s t h a t l e a d e r c l i p p i n g p r o j e c t s s h o u l d be c a r r i e d out b e f o r e the s p r i n g . The bes t time f o r t h i s i s F e b r u a r y when a t t a c k e d l e a d e r s have a l r e a d y changed c o l o r and dead t e r m i n a l s a r e e a s i l y r e c o g n i z e d . b. R e s e a r c h s h o u l d be c a r r i e d out on the taxonomy of the w e e v i l s t o ensure s p e c i e s i d e n t i f i c a t i o n when l a r v a e a l o n e a re a v a i l a b l e , and t o ensure c o r r e c t a s s o c i a t i o n between p a r a s i t o i d s and h o s t w e e v i l spec i e s . c. L u p i n , L u p i n u s sp., might be amenable t o c u l t i v a t i o n near or a t young l o d g e p o l e p i n e s t a n d s t o enhance p a r a s i t o i d s u r v i v a l by p r o v i d i n g a f e e d i n g s i t e f o r a d u l t s . d. F u r t h e r r e s e a r c h i s needed t o demonstrate how the p a r a s i t o i d complex from c l i p p e d l e a d e r s c o u l d be p r e s e r v e d and r e t u r n e d t o t h e f o r e s t . 95 BIBLIOGRAPHY A l e x a n d e r , R.R. 1960. T h i n n i n g l o d g e p o l e p i n e i n the C e n t r a l Rocky Mountains. J . F o r . 58:99-104 A l l e n , J.C. 1976. A m o d i f i e d s i n e wave method f o r c a l c u l a t i n g degree days. E n v i r o n . Entomol. 5:388-396 Amman, G.D. and L. S a f r a n y i k . 1984. I n s e c t s of l o d g e p o l e p i n e : impact and c o n t r o l . IN: Baumgartner, D.M. R.G. K r e b i l l , J.T. A r n o t t and G.F. Weetman ( e d s . ) : Lodgepole p i n e - the s p e c i e s and i t s management. Symposium P r o c e e d i n g s . Wash. S t a t e U n i v . pp.107-124 Anderson, W.H. 1941. The l a r v a and pupa of C. f u r n i s s i Buchanan ( C o l e o p t e r a : C u r c u l i o n i d a e ) . P r o c . E n t . Soc. Wash. 43:152-155 A r n o l d , C.Y. 1959. The d e t e r m i n a t i o n and s i g n i f i c a n c e of the base temperature i n a l i n e a r heat u n i t system. J . Am. Soc. H o r t i c . S c i . 74:430-435 B e l l a , I.E. 1985a. P e s t damage i n c i d e n c e i n n a t u r a l and t h i n n e d l o d g e p o l e p i n e i n A l b e r t a . F o r . Chron. 62:233-238 B e l l a , I.E. 1985b. Western g a l l r u s t and i n s e c t l e a d e r damage t o t r e e s i z e i n young l o d g e p o l e p i n e i n A l b e r t a . Can. J . F o r . Res. 15:1008-1010 Brown, J.K. 1973. F i r e c y c l e s and community dynamics i n l o d g e p o l e p i n e f o r e s t s . IN: Baumgartner, D.M. (ed.) Management of l o d g e p o l e p i n e ecosystems. Baumgartner, D.M. (Ed.) pp.429-456 Buchanan, L.L. 1940. Three new s p e c i e s of the l o n q u l u s group of C y l i n d r o c o p t u r u s ( C o l e o p t e r a : C u r c u l i o n i d a e ) . P r o c . E ntomol. Soc. Wash. 42:177-181 Bukzeeva, O.N. 1965. B i o l o g y of M a q d a l i s f r o n t a l i s G y l l . Z o o l o g i c h e s k i Zhournal 46:55-59 ( i n R u s s i a n ) . DeBach, P. 1964. B i o l o g i c a l c o n t r o l of i n s e c t p e s t s and weeds. Chapman and H a l l L t d . , London, 844pp. DeBach, P. 1974. B i o l o g i c a l c o n t r o l by n a t u r a l enemies. 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C a r o l i n . 1977. Western f o r e s t i n s e c t s . USDA F o r e s t S e r v i c e M i s c . P u b l . No.1339 654pp G r a n t , J . 1958. O b s e r v a t i o n s on a p i n e shoot moth, Eucosma  sonomana. J . Entomol. Soc. B.C. 55:26-27 Gre a t h e a d , D.J. 1986. P a r a s i t o i d s i n c l a s s i c a l b i o l o g i c a l c o n t r o l . IN: Waage, J . and D. Great h e a d (eds.) I n s e c t p a r a s i t o i d s . 13th Symposium of the R o y a l E n t o m o l o g i c a l S o c i e t y of London, Academic P r e s s , New York pp.289-318 Hagen, K.S. and R. H a l e . 1974. I n c r e a s i n g n a t u r a l enemies t h r o u g h use of supplementary f e e d i n g and n o n - t a r g e t p r a y . IN: M a x w e l l , F.G. and F.A. H a r r i s ( e d s . ) : P r o c e e d i n g s of the Summer I n s t i t u t e of B i o l o g i c a l C o n t r o l of P l a n t I n s e c t s and D i s e a s e s . J a c k s o n , pp.170-181 97 H i g l e y , L.G., L.P. Pedigo and K.R. O s t l i e . 1986. DEGDAY: A program f o r c a l c u l a t i n g degree-days, and assumptions be h i n d the degree-day approach. E n v i r o n . Entomol. 15:999-1016 Ho p k i n s , A.D. 1911. C o n t r i b u t i o n toward a monograph of the bark w e e v i l s of the genus P i s s o d e s . T e c h n i c a l S e r i e s , No.20, P a r t I , Bur. of Entomology Hopping, A. 1920. A new s p e c i e s of the genus P i s s o d e s ( C o l e o p t e r a ) . Can. E n t . 52:132-134 Hulme, M.A., A.F. Dawson and J.W.E. H a r r i s . 1986. E x p l o i t i n g c o l d - h a r d i n e s s t o s e p a r a t e P i ssodes s t r o b i (Peck) ( C o l e o p t e r a : C u r c u l i o n i d a e ) from a s s o c i a t e d i n s e c t s i n l e a d e r s of P i c e a s i t c h e n s i s (Bong.) C a r r . Can. E n t . 118:1115-1122 Hulme, M.A., J.W.E. H a r r i s and A.F. Dawson. 1987. E x p l o i t i n g a d u l t g i r t h t o s e p a r a t e P i ssodes s t r o b i (Peck) (COLEOPTERA: C u r c u l i o n i d a e ) from a s s o c i a t e d i n s e c t s i n l e a d e r s of P i c e a s i t c h e n s i s (Bong.) C a r r . Can. E n t . 119:751-753 J o h n s t o n e , W.D. 1985. T h i n n i n g l o d g e p o l e p i n e . IN: Lodgepole p i n e - the s p e c i e s and i t s management. Baumgartner D.M., R.G. K r e b i l l , J.T. A r n o t t and G.F. Weetman ( e d s . ) ; Symposium P r o c e e d i n g s . Wash. S t a t e U n i v e r s i t y , pp.251-262 Kennedy, R.W. 1985. Lodgepole p i n e as a commercial r e s o u r c e i n Canada. IN: Baumgartner D.M., R.G. K r e b i l l , J.T. A r n o t t and G.F. Weetman ( e d s . ) : Lodgepole p i n e - the s p e c i e s and i t s management. Symposium P r o c e e d i n g s . Wash. S t a t e U n i v e r s i t y , pp.21-23 Keen, F.P. 1928. I n s e c t enemies of C a l i f o r n i a p i n e s and t h e i r c o n t r o l . C a l i f . . D e p t . Nat. Res., F o r . D i v . , B u l l e t i n No.7. 113pp. Keen, F.P. 1952. I n s e c t enemies of western f o r e s t s . USDA M i s c . P u b l . 273 280pp. L i n d g r e n , B.S. 1980. P e s t s of l o d g e p o l e p i n e , P i n u s c o n t o r t a , w i t h p a r t i c u l a r r e f e r e n c e t o p o t e n t i a l impact i n Sweden. F o r e s t Entomology R e p o r t s No.3, Swedish U n i v . of A g r i c . S c i . - , U p p s a l a , 125pp. Maher, T.C. 1982. The b i o l o g y and impact of the l o d g e p o l e t e r m i n a l w e e v i l i n the C a r i b o o F o r e s t R e g i o n . M.Sc. T h e s i s , U n i v . of B.C. Vancouver, 63pp. Manna, G.K. and S.G. Smith. 1958 Chromosomal polymorphism and i n t e r r e l a t i o n s h i p s among bark w e e v i l s of the genus P i s s o d e s German. The N u c l e u s . 2:179-208 98 M a r t y , R.J. 1959. P r e d i c t i n g w e e v i l - c a u s e d volume l o s s i n w h i t e p i n e . F o r . S c i . 5:269-274 McDougal, F.W. 1973. The importance of l o d g e p o l e p i n e i n Canada, pp.10-26. IN: Baumgartner, D.M. (ed.) Management of l o d g e p o l e p i n e ecosystems. Wash. S t a t e U n i v e r s i t y , pp.10-26 M i n i s t r y of F o r e s t s . 1981-1986. Annual R e p o r t . P r o v i n c e of B.C. M i n i s t r y of F o r e s t s and Lands. 1987. P e s t Management P r o g r e s s . 6(2):25 Murdoch, W.W., J . Chesson and P.L. Chesson. 1985. B i o l o g i c a l c o n t r o l i n t h e o r y and p r a c t i c e . Amer. Nat. 125:344-366 Plumb, G.H. 1950. The a d u l t f e e d i n g h a b i t of some c o n i f e r -i n f e s t i n g w e e v i l s . Can. E n t . 82:53-57 P r i c e , D.S. 1980. The major e n t o m o l o g i c a l p e s t s of young l o d g e p o l e p i n e i n the C a r i b o o . B.S.F. T h e s i s , F a c u l t y of F o r e s t r y , U n i v . of B r i t i s h C o l u m b i a , 74pp. S a f r a n y i k , L., D.M. Shrimpton and H.S. Whitney. 1974. Management of l o d g e p o l e p i n e t o reduce l o s s e s from The mountain p i n e b e e t l e . F o r . Techn. Report 1, Pac. F o r . C e n t r e 24pp. Salman, K.A. 1935. The e f f e c t s of a t t a c k by P i s s o d e s t e r m i n a l i s Hopping on l o d g e p o l e p i n e i n C a l i f o r n i a . J . Econ. Entomol. 28:496-497 S a t t e r l a n d , D.R. 1973. C l i m a t i c f a c t o r s and l o d g e p o l e p i n e . IN: Baumgartner, D.M. ( e d . ) : Management of l o d g e p o l e p i n e ecosystems. Wash. S t a t e U n i v e r s i t y , pp.297-309 Schmidt, W.C. and R.R. A l e x a n d e r . 1985. S t r a t e g i e s f o r managing l o d g e p o l e p i n e . IN: Baumgartner, D.M., R.G. K r e b i l l , J.T. A r n o t t and G.F. Weetman (eds.) Lodgepole p i n e - the s p e c i e s and i t s management. Symposium P r o c e e d i n g s . Wash. S t a t e U n i v e r s i t y , pp.201-210 Smith, S.G. and Y. T a k e n o u c h i . 1969. Chromosomal polymorphism i n P i ssodes w e e v i l s : F u r t h e r on i n c o m p a t i b i l i t y i n P. t e r m i n a l i s . Can. J . of G e n e t i c s and C y t o l . 11:761-782 S t a r k , R.W. and D.L. Wood. 1964. The b i o l o g y of P i s s o d e s  t e r m i n a l i s Hopping ( C o l e o p t e r a : C u r c u l i o n i d a e ! i n C a l i f o r n i a . Can. E n t . 96:1208-1218 99 S t e v e n s , R.E. 1973. A s s o c i a t i o n of P i t y o p h t h o r u s opimus Blackman w i t h P i ssodes t e r m i n a l i s i n C o l o r a d o l o d g e p o l e p i n e ( C o l e o p t e r a : S c o l y t i d a e and C u r c u l i o n i d a e ) . C o l e o p t . B u l l . 27:141-142 S t e v e n s , R.E. and J.A.E. Knopf. 1074. Lodgepole t e r m i n a l w e e v i l i n i n t e r i o r l o d g e p o l e p i n e f o r e s t s . E n v i r o n . Entomol. 3:998-1002 St e v e n s o n , R.E. and J . J . P e t t y . 1968. Lodgepole t e r m i n a l w e e v i l ( P i s s o d e s t e r m i n a l i s Hopping) i n the A l b e r t a / Northwest T e r r i t o r i e s R e g i o n . Can. Dept. of F o r . B i - m o n t h l y Res. Notes 24:6 T a c k l e , D. 1959. S i l v i c s of l o d g e p o l e p i n e . USDA F o r e s t S e r v i c e , I n t e r m o u n t a i n F o r e s t and Range E x p e r i m e n t a l S t a t i o n M i s c . P u b l . No.19 24pp. Weetman, G.F., R.C. Yang and I.E. B e l l a . 1985. N u t r i t i o n and f e r t i l i z a t i o n of l o d g e p o l e p i n e . IN: Baumgartner, D.M., R.G. K r e b i l l , J.T. A r n o t t and G.F. Weetman ( e d s . ) : Lodgepole p i n e - the s p e c i e s and i t s management. Symposium P r o c e e d i n g s . Wash. S t a t e U n i v e r s i t y , pp.225-232 Wheeler, N.C. and W.B. C r i t c h f i e l d . 1985. The d i s t r i b u t i o n and b o t a n i c a l c h a r a c t e r i s t i c s of l o d g e p o l e p i n e . IN: Baumgartner, D.M., R.G. K r e b i l l , J.T. A r n o t t and G.F. Weetman ( e d s . ) : Lodgepole p i n e - the s p e c i e s and i t s management. Symposium P r o c e e d i n g s . Wash. S t a t e U n i v e r s i t y , pp.1-13 Wood, G.S., G.A. V a n S i c k l e , L. Humble. 1987. F o r e s t I n s e c t and D i s e a s e C o n d i t i o n s , B r i t i s h Columbia and Yukon 1987 Can. F o r . S e r v i c e , Pac. F o r . C e n t r e BC-X-1987 32pp. Woodhead, P.V. 1934. T h i n n i n g l o d g e p o l e p i n e s t a n d s i n the C e n t r a l Rocky Mountain R e g i o n . J . F o r . 32:594-597 APPENDIX I Form used f o r r e c o r d i n g i n s e c t l i f e s t a g e s found d u r i n g d i s s e c t i o n M i-> (71 a II II =1 II $ <a fD Cu (0 ro c 3 < 1—» n> rr i-« h-1 (TO • fD o. Oi 01 u II u •o tu 01 M 1 <: 01 < U) — fD H-rr O o u o. II •a c CU •a ffl f» Cu "3 _o Leader _ Length i p i c a l diam. b a s a l dlam. pi 3 03 Cu c c 31 ro K ro rs < 01 <: 01 ro <d 01 1 - 01 en ro 2 m e r g l o l e s o o H n r*i 55 m > T0T 102 APPENDIX I I Form used f o r r e c o r d i n g emerged specimens from r e a r i n g boxes 103 (NJ IT) (Nl <T tNJ CNI CNI (Nl 1— 3 i — ! (Nl C -o (NJ 3 CM i—! 3 00 p_ t H 3 r-i — I C -3 i — l i - 3 i H 3 ro i — l C-. 3 CNI 1—! 3 I—l i — l C -3 c 1—1 3-3 ON 3 cc C -3 C~ 3 C -3 NT CM 3 ro 3 : N I En1 - i CM 3 No. «-i CNI ro -a uo vD r~ ao o> O T—I i - l i-H (N| i — I ro i — i <T r - l uo i-H vC •H oo i-H ON I-H O CNI i-H CNI C N I CNI ro ( N J -3 C N uo CNI v O C N r> C N 00 C N I ON C N I O CO I — c-C N C I C I C I APPENDIX I I I P r o j e c t e d f i e l d emergence of w e e v i l s c a l c u l a t e d w i t h t h r e e methods of degree-day c a l c u l a t i o n f o r d i f f e r e n t d e v e l o p m e n t a l minimum t e m p e r a t u r e s , (TMIN) when the maximum d e v e l o p m e n t a l t e m p e r a t u r e , (TMAX) was unknown and s e t a t 30°C 105 Projected f i e l d emergence based on Penticton Airport data Start of lab T M I N 2 Accumulated 1 3 emergence heat units TMAX unknown Rectang. Triang. Sine Rectang. Triang. Sine Magda1i s gent i1 i s May 12 4°C 7G May 13 May 15 May 14 May 13 May 15 May 14 May 12 5°C 72 May 13 May 16 May 15 May 13 May 16 May 16 May 12 G°C €8 May 13 May 18 May 16 May 13 May 18 May 16 May 12 7°C 64 May 13 May 20 May 18 May 14 May 20 May 18 May 12 8°C 60 May 14 May 2 1 May 20 May 14 May 22 May- 20 P1ssodes term i na1 is June 3 4°C 532 June 19 June 2 1 June 20 June 19 June 19 June 19 June 3 5°C 504 June 20 June 20 June 20 June 2 1 June 21 June 2 1 June 3 6°C 476 June 21 June 22 June 22 June 2 1 June 22 June 22 June 3 7°C 448 June 22 June 23 June 23 June 22 June 23 June 23 June 3 8°C 420 June 23 June 25 June 24 June 23 June 26 June 25 CylIndrocopturus sp June 9 4°C 627 June 26 June 27 June 26 June 26 June 27 June 27 June 9 5°C 594 June 26 June 28 June 27 June 27 June 28 June 28 June 9 €°C 561 June 27 June 29 June 28 June 28 June 29 June 29 June 9 7°C 528 June 27 June 30 June 29 June 28 June 30 June 30 June 9 B°C 495 June 30 July 1 July 1 June 29 July 2 Ju 1 y 1 Specimens set up for rearing on May 7, 1987. TMIN - minimum developmental temperature TMAX - maximum developmental temperature APPENDIX IV Weekly measurements of 25 e l o n g a t i n g l o d g e p o l e p i n e l e a d e r s 107 Date May 14 1 May 24 Jun 4 Jun 14 Jun 24 Jul 4 Jul 14 Jul 24 Aug 3 Aug 13 No. Length of leader 1 8 .2 15 . 7 23 .0 36 .0 38 .6 46 .4 49 .4 49, .5 49 .8 50. .4 2 6 . a 16 . 2 21 .4 33 .2 38 .O 47 .2 50 .2 51 , 9 53 . 1 53 , . e 3 7 . 1 1 1 .5 15 .9 27 . 1 31 .5 37 .2 40. ,6 42 .3 44 .0 44 , 9 4 6 . 3 22 .8 29 .6 42 . 1 47 .9 55 .6 56, .5 57 .8 58 .6 58, .8 5 7 .6 9 .9 14 . 3 29 .9 37 .4 44 .7 47 .6 50 .6 54 .3 54 , .8 6 18 .9 25 .3 41 .4 47 .3 56 .8 59, .0 59 .5 60 .0 60 .8 7 13 .0 18 .6 32 .6 37 .2 45 .5 49 .7 52 .3 53 .5 53 .5 8 16 .8 24 .6 39 .6 41 .5 45 .9 47 .3 47 .5 47 .5 47 .8 9 17 .9 24 .9 39 .2 42 .2 49 .6 50 .8 51 . 1 52 .3 52 .4 10 8 .4 11 .2 20 .7 24 .4 28 .6 29 .6 29 .6 29 .6 29 .6 11 14 . 4 20 .3 31 .4 34 .5 39 .2 41 .9 42 .5 42 .5 42 .8 12 13 .5 17 .7 25 .2 28 .5 28 .7 28 .7 28 .7 29 .6 29 .6 13 13 . 1 19 .2 33 . 1 37 .0 44 .4 46 .6 46 .8 49 .5 49 .5 14 20 .3 26 .9 38 .5 43 .6 54 .2 58 .2 59 .5 60 .0 60 .8 15 18 .3 24 .2 36 .3 4 1 .0 45 .6 46 .7 46 .8 4S .3 48 .5 1G 19 . 7 26 .5 35 .9 38 .9 48 .3 51 .0 51 .6 52 .8 53 .4 17 21 .0 26 .5 40 .0 45 .8 53 .5 56 .9 57 . 3 56 . 4 59 . 4 18 17 .9 24 . 1 37 .7 42 .6 52 .0 55 .2 56 .4 58 .0 58 .0 19 14 .0 17 .7 29 .8 35 .0 41 . 1 41 .5 41 .6 42 .8 43 .6 20 1 1 .2 14 .9 25 .4 28 .7 34 .9 37 .8 39 .8 4 1 .4 4 1 .8 21 18 .5 26 .4 39 .5 44 .6 53 .8 58 . 1 61 .2 63 . 1 64 .0 22 15 . 1 22 .0 37 .0 42 .9 51 .3 53 .0 56 .0 58 .8 59 .5 23 16 .8 21 .2 33 .5 36 .8 39 .2 39 .5 39 .7 39 .7 39 .7 24 20 .7 25 .9 37 .9 4 1 .2 48 .8 51 .4 53 .7 53 .7 54 . 1 25 20 .7 27 .6 43 .5 48 . 1 59 . 1 62 . 1 65 .8 68 .8 70 .2 1 leaders were c o l l e c t e d on s i t e 120 near Kelowna 

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