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Dispersal and flight behavior of Trypodendron Lineatum (Olivier) (Coleoptera: Scolytidae) as influenced… Salom, Scott Michael 1989

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DISPERSAL AND FLIGHT BEHAVIOR OF TRYPODENDRON  LINEATUM (OLIVIER) (COLEOPTERA: SCOLYTIDAE) AS INFLUENCED BY SEMIOCHEMICAL AND ENVIRONMENTAL FACTORS By SCOTT MICHAEL SALOM B.S., Iowa S t a t e U n i v e r s i t y , 1981 M.S., U n i v e r s i t y o f A r k a n s a s , 1985 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY i n THE FACULTY OF GRADUATE STUDIES (Department o f F o r e s t S c i e n c e s ) 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 t h e r e q u i r e d s t a n d a r d THE UNIVERSITY OF BRITISH COLUMBIA A p r i l 7, 1989 © S c o t t M i c h a e l Salom, 1989 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of The University of British Columbia Vancouver, Canada Date DE-6 (2/88) i i ABSTRACT L a b o r a t o r y b i o a s s a y s u s i n g a wind t u n n e l were developed t o study f l i g h t b e h a v i o r and o r i e n t a t i o n o f the s t r i p e d ambrosia b e e t l e , Trvpodendron lineatum ( O l i v i e r ) . F a c t o r s t h a t were s t u d i e d i n d e t a i l were windspeed, semiochemical c o n c e n t r a t i o n s , and semiochemical combinations. In the f i r s t o f two experiments examining the e f f e c t o f wind speed on T. li n e a t u m response t o a semiochemical-baited s u b s t r a t e , the h i g h e s t % of males (21.4) and females (25.3) caught i n a fu n n e l t r a p , o c c u r r e d a t 0.0 m/s. As windspeed was i n c r e a s e d from 0.0 t o 0.9 m/s, the per c e n t o f b e e t l e s caught decreased l i n e a r l y f o r both sexes. A second experiment showed t h a t i n the presence of wind, responding b e e t l e s o r i e n t e d a n e m o t a c t i c a l l y t o the semiochemical-baited s u b s t r a t e . With wind absent, b e e t l e s flew randomly and e r r a t i c a l l y . Upon r e a c h i n g c l o s e t o the b a i t e d s u b s t r a t e , a g r e a t e r p r o p o r t i o n o f the b e e t l e s responded t o the chemical s t i m u l i and landed on the s u b s t r a t e than when an a i r f l o w was pr e s e n t . These r e s u l t s suggest t h a t T. li n e a t u m are capable of responding under v a r i e d wind c o n d i t i o n s t y p i c a l l y p r e s e n t i n a f o r e s t , whereby they use wind t o o r i e n t t o o l f a c t o r y s t i m u l i , y e t are bes t a r r e s t e d t o the s t i m u l i under s t i l l c o n d i t i o n s . F l i g h t response of T. lineatum t o a m u l t i p l e f u n n e l t r a p b a i t e d w i t h e t h a n o l (1° a t t r a c t a n t ) and l i n e a t i n (2° a t t r a c t a n t ) a t d i f f e r e n t r e l e a s e r a t e s , i n d i c a t e d t h a t o n l y l i n e a t i n was e f f e c t i v e i n a t t r a c t i n g b e e t l e s t o t h i s k i n d o f t r a p . Maximum re s p o n s e by b o t h males and f e m a l e s o c c u r r e d a t r e l e a s e r a t e s o f l i n e a t i n between 8 and 64 ug/24 h. However, i n a more d e t a i l e d s t u d y o f T. l i n e a t u m r e s p o n s e t o e t h a n o l and l i n e a t i n u s i n g m o d i f i e d d r a i n p i p e t r a p s , s e r v i n g as a model o f a h o s t t r e e , e t h a n o l d i d p o s i t i v e l y i n f l u e n c e male f l i g h t t y p e , speed o f r e a c t i o n , and d i r e c t i o n . N e v e r t h e l e s s , l i n e a t i n was t h e most i m p o r t a n t s e m i o c h e m i c a l i n a t t r a c t i n g males t o l a n d on and e n t e r t h e t r a p s . E t h a n o l was more i m p o r t a n t f o r f e males t h a n f o r males, and when combined w i t h l i n e a t i n , p r o v i d e d t h e o p t i m a l s t i m u l i f o r a t t r a c t i n g f e m ales t o l a n d on and e n t e r t h e t r a p s . P o p u l a t i o n movement o f s p r i n g d i s p e r s i n g T. l i n e a t u m was s t u d i e d u s i n g m a r k - r e c a p t u r e t e c h n i q u e s w i t h l i n e a t i n -b a i t e d f u n n e l t r a p s . I n a f i r s t s e t o f s t u d i e s c o n d u c t e d i n an even-aged second-growth c o a s t a l f o r e s t i n B r i t i s h C o l umbia, b e e t l e r e c a p t u r e d i s t r i b u t i o n was compared w i t h w i nd d i r e c t i o n a t d i s t a n c e s between 5 and 500 m from t h e b e e t l e r e l e a s e s i t e . A t 5 and 25 m, b e e t l e r e c a p t u r e was p r e d o m i n a n t l y upwind. W i t h t r a p s p l a c e d o n l y a t 100 m from t h e r e l e a s e s i t e , b e e t l e s were r e c a p t u r e d i n a l l d i r e c t i o n s i r r e s p e c t i v e o f wind. However, w i t h t r a p s p l a c e d o n l y 500 m from t h e r e l e a s e s i t e , b e e t l e s were o n l y r e c a p t u r e d i n t h e downwind t r a p s . I n m a r k - r e c a p t u r e e x p e r i m e n t s c o n d u c t e d i n a v a l l e y , b e e t l e s r e l e a s e d from a f o r e s t m a r g i n i n f l u e n c e d by p r e v a i l i n g u p - v a l l e y winds, f l e w upwind w i t h i n t h e f o r e s t t o i v l i n e a t i n - b a i t e d f u n n e l t r a p s p l a c e d 25 m from t h e r e l e a s e s i t e . B e e t l e r e c a p t u r e i n an open s e t t i n g was h i g h e r a l o n g t h e edge o f t h e open s e t t i n g t h a n i n i t s c e n t e r , 325 m c l o s e r t o t h e r e l e a s e s i t e . B e e t l e s were r e c a p t u r e d 1 km d o w n - v a l l e y (upwind) and 1.9 km u p - v a l l e y (downwind) from t h e r e l e a s e s i t e . I n one exp e r i m e n t (two r e l e a s e s ) , 10.6 and 7.8% o f t h e marked b e e t l e s r e c a p t u r e d were c o l l e c t e d i n t r a p l i n e s > 700 m and > 1 km from t h e r e l e a s e s i t e , r e s p e c t i v e l y . I n a d d i t i o n a l m a r k - r e c a p t u r e e x p e r i m e n t s i n t h e v a l l e y , b e e t l e s were r e l e a s e d s i m u l t a n e o u s l y from a windward and l e e w a r d s i d e o f a f o r e s t margin i n t h e v a l l e y t h r o u g h two e x p e r i m e n t s o f f o u r r e l e a s e s each. W i t h l o n g d i s t a n c e f l i g h t emphasized and no s e m i o c h e m i c a l - b a i t e d t r a p s p l a c e d w i t h i n 200 m o f e i t h e r r e l e a s e s i t e , p o p u l a t i o n movement was p r e d o m i n a n t l y downwind. B e e t l e s a l s o f l e w a c r o s s t h e v a l l e y t o t r a p s on t h e o p p o s i t e f a c i n g s l o p e a t a f a i r l y h i g h f r e q u e n c y (38% o f t h e r e c a p t u r e d b e e t l e s ) , d u r i n g t h e f i r s t e x p e r i m e n t . B e e t l e s were r e c a p t u r e d a t a much h i g h e r f r e q u e n c y i n t r a p s p l a c e d w i t h i n a f o r e s t as compared t o t h o s e i n an open s e t t i n g . T h i s was l i k e l y a r e s u l t o f t h e c a l m e r wind c o n d i t i o n s under t h e f o r e s t canopy, f a c i l i t a t i n g b e t t e r f l y i n g c o n d i t i o n s and re s p o n s e t o o l f a c t o r y s t i m u l i f o r t h e b e e t l e s . The i m p l i c a t i o n s t h e s e f i n d i n g s have on t h e g e n e r a l knowledge o f s c o l y t i d b e e t l e d i s p e r s a l and o r i e n t a t i o n t o o l f a c t o r y s t i m u l i a r e d i s c u s s e d . New c o n s i d e r a t i o n s t o w a r d V improving p e s t management s t r a t e g i e s f o r T. l i n e a t u m as a r e s u l t of these s e t s of s t u d i e s are presented. y i TABLE OF CONTENTS Page ABSTRACT i i TABLE OF CONTENTS v i LIST OF TABLES i x LIST OF FIGURES x i i ACKNOWLE DGEMENTS xv i i i CHAPTER 1: OVERVIEW 1 1.1 I n t r o d u c t i o n 1 1.2 O b j e c t i v e s 7 1.3 Format 10 CHAPTER 2: THE EFFECT OF WIND SPEED ON TRYPODENDRON  LINEATUM FLIGHT BEHAVIOR IN A WIND TUNNEL 12 2.1 I n t r o d u c t i o n 12 2.2 M a t e r i a l s and Methods 14 2.3 R e s u l t s 21 2.4 D i s c u s s i o n 32 CHAPTER 3: RESPONSE OF TRYPODENDRON LINEATUM TO VARIED RELEASE RATES OF ETHANOL AND LINEATIN IN A WIND TUNNEL 39 3.1 I n t r o d u c t i o n 39 3.2 M a t e r i a l s and Methods 41 3.3 R e s u l t s 44 3.4 D i s c u s s i o n 46 CHAPTER 4: WIND TUNNEL EVALUATION OF FLIGHT AND LANDING BEHAVIOR OF TRYPODENDRON  LINEATUM (OLIVIER) IN RESPONSE TO A HOST ATTRACTANT AND AGGREGATION PHEROMONE 49 4.1 I n t r o d u c t i o n 49 v i i 4.2 M a t e r i a l s and Methods 51 4.3 R e s u l t s 56 4.4 D i s c u s s i o n 69 CHAPTER 5: INFLUENCE OF WIND ON THE SPRING FLIGHT OF TRYPODENDRON LINEATUM IN A SECOND-GROWTH CONIFEROUS FOREST 74 5.1 I n t r o d u c t i o n 74 5.2 M a t e r i a l s and Methods 75 5.3 R e s u l t s and D i s c u s s i o n 82 CHAPTER 6: DISPERSAL OF TRYPODENDRON LINEATUM (OLIVIER) WITHIN A VALLEY SETTING 96 6.1 I n t r o d u c t i o n 96 6.2 M a t e r i a l s and Methods 97 6.3 R e s u l t s and D i s c u s s i o n 106 CHAPTER 7: POPULATION MOVEMENT PATTERNS OF TRYPODENDRON LINEATUM AS INFLUENCED BY WIND AND VEGETATION WITHIN A VALLEY 12 9 7.1 I n t r o d u c t i o n 129 7.2 M a t e r i a l s and Methods 13 0 7.3 R e s u l t s and D i s c u s s i o n 139 CHAPTER 8: CONCLUSIONS 161 REFERENCES 170 APPENDIX 1: RESPONSE OF TRYPODENDRON LINEATUM (OLIVIER) TO DIFFERENT SEMIOCHEMICAL-BAITED SUBSTRATES IN A WIND TUNNEL 181 APPENDIX 2: RESPONSE OF FLOWN AND UNFLOWN T. LINEATUM TO SEMIOCHEMICAL-BAITED FUNNEL TRAPS IN A WIND TUNNEL 184 A. 2.1 I n t r o d u c t i o n 184 A.2.2 M a t e r i a l s and Methods 185 A.2.3 R e s u l t s and D i s c u s s i o n 189 APPENDIX 3: INFLUENCE OF DUSTING T. LINEATUM WITH v i i i FLUORESCENT POWDER ON THEIR RESPONSE TO SEMIOCHEMICALS IN A WIND TUNNEL 193 ix LIST OF TABLES Ta b l e 1. T a b l e 2 Page Frequency of Trypodendron l i n e a t u m f l i g h t i n a wind t u n n e l a t v a r i e d wind speeds i n the presence of an u n b a i t e d and semiochemical-baited s i m u l a t e d l o g S t a t i s t i c a l comparison of T. l i n e a t u m responses t o f i r s t and second year c o n t r o l treatments i n a wind t u n n e l . The numbers of the b e e t l e s r e l e a s e d f o r each sex/ treatment = 124 24 57 T a b l e 3. F l i g h t frequency of T. l i n e a t u m t o d i f f e r e n t semiochemical b a i t s i n a wind t u n n e l . Number of b e e t l e s r e l e a s e d f o r each sex i n each b a i t treatment = 124. 58 T a b l e 4. T a b l e 5, T a b l e 6, T a b l e 7, T a b l e 8. The capture of T. lineatum i n semiochemical-baited d r a i n p i p e t r a p s i n a wind t u n n e l (N = 124 r e l e a s e d / t r e a t m e n t / sex) 68 Numbers of marked T. lineatum r e l e a s e d and r e c a p t u r e d i n the f l i g h t d i r e c t i o n study on the U n i v e r s i t y Endowment Lands, Vancouver, B.C., 1986 84 R e l a t i o n s h i p of f l i g h t d i r e c t i o n of T. lineatum t o the wind d i r e c t i o n , u s i n g a one-sample t e s t f o r the mean d i r e c t i o n a l f l i g h t angle on the U n i v e r s i t y Endowment Lands, Vancouver, B.C., 1986 Percent of T. lineatum caught a t v a r i e d d i s t a n c e s i n the mark-recapture study, experiments 1 and 2, U n i v e r s i t y Endowment Lands, Vancouver, B.C., 1986 Comparison of wind d i r e c t i o n and speed between f o r e s t e d and open s i t e s w i t h i n the Cedar Creek V a l l e y ( u p - v a l l e y azimuth of 48 t o 68°) i n the Coquitlam Lake Watershed B.C., 1987 87 90 108 Table 9. T a b l e 10. F l i g h t and r e c a p t u r e success of T. lineatum i n the mark-recapture study a t the Coquitlam Lake Watershed, B.C., 1987 (Day 1 and Day 2 data were p o o l e d ) . The number of marked T. lineatum r e c a p t u r e d w i t h r e s p e c t t o time i n a 110 X Table 11, T a b l e 12 dense o l d growth f o r e s t i n the Coquitlam Lake Watershed, B.C., 1987 Mean per c e n t of T. lineatum r e c a p t u r e d a t v a r i e d d i s t a n c e s i n the Cedar Creek V a l l e y , Coquitlam Lake Watershed, B.C., 1987 , Comparison of T. lineatum f l i g h t and wind d i r e c t i o n i n an o l d growth f o r e s t i n the Coquitlam Lake Watershed, B.C., 1987 I l l 113 114 T a b l e 13 Table 14, T a b l e 15, T a b l e 16, T a b l e 17 Table 18, T a b l e 19 Weather measurements made between 113 0 and 1700 h PDT, from the Cedar Creek V a l l e y i n the Coquitlam Lake Watershed i n B r i t i s h Columbia d u r i n g 1988 141 F l i g h t and r e c a p t u r e success of Trypodendron lineatum i n the mark-r e c a p t u r e study conducted a t the Coquitlam Lake Watershed i n B r i t i s h Columbia d u r i n g 1988 Comparison between up- and down-valley movement of a d u l t Trypodendron l i n e a t u m d u r i n g the mark-recapture study conducted i n the Cedar Creek V a l l e y w i t h i n the Coquitlam Lake Watershed i n B r i t i s h Columbia d u r i n g 1988 Movement of T. lineatum through a f o r e s t , f l y i n g u p - v a l l e y ( p o p u l a t i o n A) and down-v a l l e y ( p o p u l a t i o n B), d u r i n g f o u r separate r e l e a s e s i n experiment 2, w i t h i n the Coquitlam Lake Watershed i n B r i t i s h Columbia d u r i n g 1988 Comparison between an open and f o r e s t e d r o u t e f o r l o n g d i s t a n c e f l y i n g Trypodendron lineatum, d u r i n g 4 r e l e a s e s i n experiment 2, w i t h i n the Coquitlam Lake Watershed i n B r i t i s h Columbia d u r i n g 1988 , Mean r e c a p t u r e percentage of T. l i n e a t u m t o d i f f e r e n t s u b s t r a t e s b a i t e d w i t h semiochemicals i n a wind t u n n e l . S i x r e p l i c a t e s of 50 b e e t l e s , f o r a t o t a l o f 3 00 b e e t l e s were r e l e a s e d f o r each treatment by sex category Response of flown and unflown T. l i n e a t u m t o semiochemical-baited f u n n e l t r a p s i n a 148 150 154 156 183 x i wind t u n n e l 190 T a b l e 20. The e f f e c t o f m a r k i n g t r e a t m e n t s on T. l i n e a t u m r e s p o n s e t o a s e m i o c h e m i c a l -b a i t e d f u n n e l t r a p i n a wind t u n n e l 195 x i i LIST OF FIGURES Page F i g . 1 Body o r i e n t a t i o n d i r e c t i o n s of T. lineatum scored p r i o r t o and d u r i n g f l i g h t t a k e - o f f , as viewed from above the r e l e a s e t r a y ( t o p ) ; and f l i g h t t a k e - o f f d i r e c t i o n s of the same b e e t l e s , as viewed from the s i d e (bottom). A l l b e e t l e s t a k i n g o f f t h a t d i d not f i t i n t o c a t e g o r i e s A t o D were p l a c e d i n the n o - d i r e c t i o n category. ...2 0 F i g . 2 The i n f l u e n c e of wind speed on response of male (top) and female (bottom) T. lineatum t o a semiochemical-baited f u n n e l t r a p i n a wind t u n n e l . Numbers next t o boxes r e p r e s e n t the number of data p o i n t s . White and b l a c k boxes r e p r e s e n t the % response f o r the b a i t e d and u n b a i t e d ( c o n t r o l ) treatments, r e s p e c t i v e l y . M i s s i n g data accounts f o r some wind speeds having l e s s than 9 o b s e r v a t i o n s 2 2 F i g . 3 Body o r i e n t a t i o n of T. lineatum, p r i o r t o and d u r i n g f l i g h t t a k e - o f f , a t d i f f e r e n t wind speeds, i n the presence of an u n b a i t e d and semiochemical-baited s i m u l a t e d l o g . Male and female data were pooled. Bars w i t h d i f f e r e n t l e t t e r s f o r each response by b a i t treatment c a t e g o r y are s i g n i f i c a n t l y d i f f e r e n t (X ; p < 0.05) f o r i n d i v i d u a l comparisons 25 F i g . 4 F l i g h t t a k e - o f f d i r e c t i o n of T. lineatum. a t d i f f e r e n t wind speeds, i n the presence of an u n b a i t e d and semiochemical-b a i t e d s i mulated l o g . Male and female data were pooled. Bars w i t h d i f f e r e n t l e t t e r s f o r each response by b a i t treatment category are s i g n i f i c a n t l y d i f f e r e n t (X 2; p < 0.05) f o r i n d i v i d u a l comparisons 2 6 F i g . 5 O v e r a l l f l i g h t d i r e c t i o n e x h i b i t e d by T. lineatum, a t d i f f e r e n t wind speeds, i n the presence of an u n b a i t e d and semiochemical-baited s i m u l a t e d l o g . Male and female data were pooled. Bars w i t h d i f f e r e n t l e t t e r s f o r each response by b a i t treatment category are s i g n i f i c a n t l y d i f f e r e n t (X 2; p < 0.05) f o r i n d i v i d u a l comparisons 28 x i i i F i g . 6 Type of f l i g h t e x h i b i t e d by T. lineatum, a t d i f f e r e n t wind speeds, i n the presence of an unb a i t e d and semiochemical-baited s i m u l a t e d l o g . Bars w i t h d i f f e r e n t l e t t e r s f o r each response by b a i t treatment by sex categor y are s i g n i f i c a n t l y d i f f e r e n t (X 2; p < 0.05) f o r i n d i v i d u a l comparisons 2 9 F i g . 7 P o s i t i v e l ock-on response e x h i b i t e d by T. lineatum. a t d i f f e r e n t wind speeds, i n the presence o f an u n b a i t e d and semiochemical-baited s i m u l a t e d l o g . Male and female data were pooled. Bars w i t h d i f f e r e n t l e t t e r s f o r each response by b a i t treatment category are s i g n i f i c a n t l y d i f f e r e n t (X 2; p < 0.05) f o r i n d i v i d u a l comparisons 31 F i g . 8 Mean p e r c e n t (+) S.E. of T. lin e a t u m caught i n t r a p s i n response t o combinations o f v a r i e d r e l e a s e r a t e s o f l i n e a t i n and et h a n o l i n a windtunnel: A) Males and B) Females. L i n e a t i n r e l e a s e r a t e columns, pooled a c r o s s e t h a n o l treatments, with the d i f f e r e n t l e t t e r s a re s i g n i f i c a n t l y d i f f e r e n t (SNK; p < 0.05) 45 F i g . 9 A) M o d i f i e d Norwegian d r a i n p i p e t r a p s used as a b a i t i n g s u b s t r a t e f o r T. lineatum, w i t h a t y p i c a l smoke plume p a t t e r n produced a t 0.15 m/s. B) Close-up of t r a p cover and l u r e h o l d e r , showing an upper and lower l i n e a t i n b a i t , and a medial e t h a n o l b a i t 53 F i g . 10 Mean time i n t e r v a l + S.E. ( i n d i c a t e d by a v e r t i c a l l i n e running through the top of each bar) between r e l e a s e and f l i g h t t a k e - o f f of T. lineatum, i n the presence of semiochemical b a i t s i n d r a i n p i p e t r a p s , w i t h i n a wind t u n n e l 60 F i g . 11 F l i g h t t a k e - o f f d i r e c t i o n of T. lineatum, i n the presence of semiochemical b a i t s i n d r a i n p i p e t r a p s , w i t h i n a wind t u n n e l . Male and female data were pooled. S t a r s on top of unb a i t e d (1988) bars denotes a s i g n i f i c a n t d i f f e r e n c e (X 2; p < 0.05) with the unb a i t e d (1987) treatment. Bars f o r each d i r e c t i o n , w i t h d i f f e r e n t l e t t e r s , i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e s i n response (X ; p < 0.05), \ x i v f o r i n d i v i d u a l c omparisons. The numbers o f o b s e r v a t i o n s f o r each t r e a t m e n t were: u n b a i t e d (1988) - 137; u n b a i t e d (1987) -138; e t h a n o l - 146; l i n e a t i n - 150; and l i n e a t i n and e t h a n o l - 152 61 F i g . 12 O b s e r v a t i o n s o f f l i g h t t y p e e x h i b i t e d by T. l i n e a t u m . i n t h e p r e s e n c e o f s e m i o c h e m i c a l s i n d r a i n p i p e t r a p s , w i t h i n a wind t u n n e l . Male and female d a t a were p o o l e d . P r o p o r t i o n s a r e based on t h e t o t a l number o f o b s e r v a t i o n s , where each ' i n d i v i d u a l c o u l d be s c o r e d f o r more t h a n one t y p e o f f l i g h t . B a r s f o r each t y p e o f f l i g h t , w i t h d i f f e r e n t l e t t e r s , i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e s i n i n r e s p o n s e ( X 2 ; p < 0.05), f o r i n d i v i d u a l c omparisons. The numbers o f o b s e r v a t i o n s f o r each t r e a t m e n t were: u n b a i t e d - 140; e t h a n o l - 152; l i n e a t i n -158; and l i n e a t i n and e t h a n o l - 156 63 F i g . 13 O v e r a l l f l i g h t d i r e c t i o n e x h i b i t e d by T. l i n e a t u m . i n t h e p r e s e n c e o f s e m i o c h e m i c a l b a i t s i n d r a i n p i p e t r a p s , w i t h i n a wind t u n n e l . Male and female d a t a were p o o l e d . B a r s f o r each d i r e c t i o n , w i t h d i f f e r e n t l e t t e r s , i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e s i n r e s p o n s e (X ; p < 0.05), f o r i n d i v i d u a l c o mparisons. The numbers o f o b s e r v a t i o n s f o r each t r e a t m e n t were : u n b a i t e d - 141; e t h a n o l - 158; l i n e a t i n - 166; and l i n e a t i n and e t h a n o l - 166 62 F i g . 14 P o s i t i v e l o c k - o n r e s p o n s e , e x h i b i t e d as l a n d i n g o r n o n - l a n d i n g b e h a v i o r , by T. l i n e a t u m i n t h e p r e s e n c e o f s e m i o c h e m i c a l b a i t s i n d r a i n p i p e t r a p s , w i t h i n a wind t u n n e l . B a r s f o r each b e h a v i o r and sex, w i t h d i f f e r e n t l e t t e r s , i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e s i n r e s p o n s e (X ; p < 0.05), f o r i n d i v i d u a l c o m p a r i s o n s . The numbers o f males o b s e r v e d f o r each t r e a t m e n t were: u n b a i t e d (U) - 67; e t h a n o l (E) - 79; l i n e a t i n (L) - 83; and l i n e a t i n and e t h a n o l (L+E) - 79. The numbers o f females o b s e r v e d were; U - 80; E - 85; L - 90; and L+E - 89 65 F i g . 15 Mean t i m e i n t e r v a l + S.E. ( i n d i c a t e d by v e r t i c a l l i n e s r u n n i n g t h r o u g h t h e t o p o f each bar) between f l i g h t t a k e - o f f and t r a p l a n d i n g o f T. l i n e a t u m f i n t h e X V presence of semiochemical b a i t s i n d r a i n p i p e t r a p s , w i t h i n a wind t u n n e l . Means were compared on combined data f o r the sexes, where d i f f e r e n t l e t t e r s on top of the bars i n d i c a t e s i g n i f i c a n t d i f f e r e n c e s between semiochemical b a i t s (SNK; p < 0.05) 67 F i g . 16 Trap placement i n mark-recapture s t u d i e s of T. lineatum on the U n i v e r s i t y Endowment lands i n Vancouver, B r i t i s h Columbia. In the f i r s t experiment, 4, 8, and 16 l i n e a t i n - b a i t e d f u n n e l t r a p s were p l a c e d a t d i s t a n c e s of 5, 25, and 100 m, r e s p e c t i v e l y , from the r e l e a s e p o i n t . Experiment 2 u t i l i z e d the 100 m t r a p l i n e o n l y . In experiment 3, t r a p s were s e t up 500 m from the r e l e a s e p o i n t o n l y . The f i r s t r e p l i c a t i o n had 4 t r a p s p l a c e d i n each c a r d i n a l d i r e c t i o n , and the second and t h i r d r e p l i c a t i o n s has 2 t r a p s a t 45° i n t e r v a l s as shown here 77 F i g . 17 D i r e c t i o n a l data based on frequency d i s t r i b u t i o n s of T. lineatum f l i g h t and wind o b s e r v a t i o n s f o r experiment 1 i n the U n i v e r s i t y Endowment Lands study 85 F i g . 18 D i r e c t i o n a l data based on frequency d i s t r i b u t i o n s of T. lineatum f l i g h t and wind o b s e r v a t i o n s f o r experiments 2 and 3 i n the U n i v e r s i t y Endowments Land study. Dashed l i n e i n c i r c l e s r e p r e s e n t s mean d i r e c t i o n . Numbers i n c i r c l e s are the r - v a l u e s . Those marked wi t h a s t a r are s i g n i f i c a n t (p < 0.05). The d o t t e d l i n e s and a s s o c i a t e d v a l u e s o u t s i d e the c i r c l e s are the s c a l e s f o r the frequency bars 91 F i g . 19 A p e r s p e c t i v e view of Cedar Creek V a l l e y , Coquitlam Lake Watershed, B r i t i s h Columbia. F i v e mark-recapture experiments f o r s t u d y i n g T. lineatum d i s p e r s a l were c a r r i e d out i n t h i s v a l l e y from June-August, 1987. The u p - v a l l e y azimuth ranges from 48 t o 68°. The road moving from the southwest t o the n o r t h e a s t covers a d i s t a n c e of 4 km i n t h i s view. Slope peaks reach 1200 m i n e l e v a t i o n , w h i l e creek e l e v a t i o n averages 550 m 98 F i g . 20 D i r e c t i o n a l data based on frequency d i s t r i b u t i o n s of T. lineatum f l i g h t and xv i wind o b s e r v a t i o n s f o r experiments 1 - 3, i n the Cedar Creek V a l l e y , Coquitlam Lake Watershed, B.C., 1987. L i n e i n c i r c l e s r e p r e s e n t mean d i r e c t i o n . Numbers i n c i r c l e s r e p r e s e n t s the r - v a l u e s . Those marked w i t h a s t a r are s i g n i f i c a n t (p < 0.05). The number and a s s o c i a t e d l i n e o u t s i d e the c i r c l e s are the s c a l e s f o r the frequency bars 115 F i g . 21 Recapture of marked T. lineatum i n pheromone t r a p s , f o r experiment 3, i n the Cedar Creek V a l l e y , Coquitlam Lake Watershed, B.C., 1987. Traps were p l a c e d c i r c u l a r l y around the r e l e a s e p o i n t and i n t r a p l i n e s 50, 375, and 700 m e a s t and 1000 m west of i t . These catches are based on 4 r e l e a s e s where a t o t a l of 11 196 b e e t l e s flew. A) The t o t a l number of b e e t l e s r e c a p t u r e d on the day of r e l e a s e ; B) The number of b e e t l e s r e c a p t u r e d d u r i n g the next c o l l e c t i o n date (2 - 10 days l a t e r ) 120 F i g . 22 Recapture of marked T. lineatum i n pheromone t r a p s f o r experiment 4 i n the Cedar Creek V a l l e y , Coquitlam Lake Watershed, B.C., 1987. T r a p l i n e s were s e t up 50, 700, 1100, 1900 m e a s t and 1000 m west of the r e l e a s e p o i n t . A d d i t i o n a l l y , l i n e E-F was p l a c e d on the o t h e r s i d e of the v a l l e y , ca. 1300 m e a s t of the r e l e a s e p o i n t . These catches are based on 2 r e l e a s e s where a t o t a l of 6 698 b e e t l e s flew. A) The number of b e e t l e s r e c a p t u r e d on the day of r e l e a s e ; B) The number of b e e t l e s r e c a p t u r e d d u r i n g the next c o l l e c t i o n date ( 3 - 5 days l a t e r ) 12 3 F i g . 23 Contour map of the Cedar Creek V a l l e y i n the Coquitlam Lake Watershed, w i t h the designs of the f i r s t (A) and second (B) mark-recapture experiments, c a r r i e d out i n 1988. Dotted polygons r e p r e s e n t r e g e n e r a t i o n openings surrounded by old-growth f o r e s t . The u p - v a l l e y azimuth = 48 - 68° 131 F i g . 24 Wind d i r e c t i o n and T. lineatum f l i g h t r e c a p t u r e f o r experiment 1, r e l e a s e s 1 (A) and 2 (B), r e p r e s e n t e d by c i r c u l a r and xy a x i s histograms, r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n along the l e n g t h x v i i of the v a l l e y . The x - a x i s r e p r e s e n t s the v a l l e y azimuth. L i n e s i n the c i r c l e s a re the mean d i r e c t i o n f o r the f i r s t 2 h (d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g the r - v a l u e f o r the v e c t o r 143 F i g . 25 Wind d i r e c t i o n and T. lineatum f l i g h t r e c a p t u r e f o r experiment 1, r e l e a s e s 3 (A) and 4 (B), re p r e s e n t e d by c i r c u l a r and xy a x i s histograms, r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n along the l e n g t h of the v a l l e y . The x - a x i s r e p r e s e n t s the v a l l e y azimuth. L i n e s i n the c i r c l e s a re the mean d i r e c t i o n f o r the f i r s t 2 h (d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g the r - v a l u e f o r the v e c t o r 144 F i g . 2 6 Wind d i r e c t i o n and T. lineatum f l i g h t r e c a p t u r e f o r experiment 2, r e l e a s e s 1 (A) and 2 (B), re p r e s e n t e d by c i r c u l a r and xy a x i s histograms, r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n a l o n g the l e n g t h of the v a l l e y . The x - a x i s r e p r e s e n t s the v a l l e y azimuth. L i n e s i n the c i r c l e s are the mean d i r e c t i o n f o r the f i r s t '2 h (d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g the r - v a l u e f o r the v e c t o r 14 5 F i g . 27 Wind d i r e c t i o n and T. lineatum f l i g h t r e c a p t u r e f o r experiment 2, r e l e a s e s 3 (A) and 4 (B), rep r e s e n t e d by c i r c u l a r and xy a x i s histograms, r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n along the l e n g t h of the v a l l e y . The x - a x i s r e p r e s e n t s the v a l l e y azimuth. L i n e s i n the c i r c l e s are the mean d i r e c t i o n f o r the f i r s t 2 h (d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g the r - v a l u e f o r the v e c t o r 146 ACKNOWLEDGEMENTS xv i i i The e d u c a t i o n and e x p e r i e n c e s I have r e c e i v e d d u r i n g my t e n u r e a t t h e U n i v e r s i t y o f B r i t i s h Columbia have been e x t r a o r d i n a r y . Many i n d i v i d u a l s and o r g a n i z a t i o n s have p r o v i d e d i n v a l u a b l e a s s i s t a n c e i n h e l p i n g me r e a l i z e t h e g o a l o f c o m p l e t i n g t h i s d i s s e r t a t i o n , and I would l i k e t o thank them a l l . Dr. J . McLean, as my t e a c h e r , r e s e a r c h s u p e r v i s o r , and f r i e n d has been a c o n s t a n t s o u r c e o f i n s p i r a t i o n d u r i n g t h e p a s t f o u r y e a r s w i t h h i s p a t i e n c e and o v e r t w i l l i n g n e s s t o h e l p and a d v i s e me i n a m u l t i t u d e o f s i t u a t i o n s . D rs. S. L i n d g r e n , L. S a f r a n y i k , B. van d e r Kamp, and D. Lavender have p r o v i d e d exemplary s e r v i c e as members o f my g r a d u a t e committee. My a p p r e c i a t i o n i s extended t o D r s . M. F u r n i s s , H. MacCarthy, J . Sweeney, R. B r i d g e s , T. Shore, L. P h e l a n , and J . Borden f o r r e v i e w i n g i n d i v i d u a l c h a p t e r s . N. Ryant, J . G l a u b i t z , D. C h a l l e n g e r , L. F r i s k i e , and E. Kovacs p r o v i d e d i n v a l u a b l e t e c h n i c a l a s s i s t a n c e i n c a r r y i n g o u t t h e s t u d i e s . Drs. A. Kozak, P. M a r s h a l l , and M. Stephens, Mr. G. T h e r i e n , and F. Ho c o n t r i b u t e d s i g n i f i c a n t l y t o t h e s t a t i s t i c a l a n a l y s i s o f much o f t h e s e d a t a . My t h a n k s a r e extended t o t h e G r e a t e r Vancouver R e g i o n a l D i s t r i c t , and G. J o y c e i n p a r t i c u l a r , f o r a l l o w i n g me t o c a r r y o u t t h e f i e l d s t u d i e s d u r i n g each o f t h r e e summers on l a n d t h e y a d m i n i s t e r . M e t e o r o l o g i c a l d a t a d u r i n g t h e summer o f 1986 xix were p r o v i d e d t o me by A. Neighbour o f t h e U.B.C. C l i m a t o l o g i c a l S t a t i o n . The M i n i s t r y o f Environment l e n t me a S t e v e n s o n s c r e e n and Dr. D. G o l d i n g and K. Rae l e n t me t h e i r CR 21 d a t a l o g g i n g equipment. Dr. J . W o r r a l l g r a c i o u s l y a l l o w e d me t o use much o f h i s workspace and w a l k -i n c o o l e r i n t h e basement o f Header House. Dr. P. Murtha and R. W i a r t p r o v i d e d t e c h n i c a l a s s i s t a n c e and a c c e s s t o t h e T e r r a s o f t program used t o produce t h e v a l l e y p e r s p e c t i v e ( F i g . 1 9 ) . N. K r a j c i , L. F r i s k i e , and M. Salom c o n t r i b u t e d t h e i r d r a f t i n g e x p e r t i s e t o s e v e r a l f i g u r e s . I am g r a t e f u l t o t h e U n i v e r s i t y o f B r i t i s h C o lumbia, t h e I.W. K i l l a m F o u n d a t i o n , and t h e N a t u r a l S c i e n c e s and E n g i n e e r i n g R e s e a r c h C o u n c i l f o r p r o v i d i n g me w i t h f i n a n c i a l s u p p o r t t h r o u g h o u t t h e c o u r s e o f t h e s e s t u d i e s , and Dr. J . W i l s o n f o r h e l p i n g t o f a c i l i t a t e t h e f u n d i n g p r o c e s s . My s i n c e r e s t t h a n k s go t o my f a m i l y who have had t r u s t i n my a b i l i t i e s and have encouraged me t o c o n t i n u e my e d u c a t i o n . Most o f a l l my w i f e Mar has ne v e r l e t me c o n s i d e r a n y t h i n g o t h e r t h a n t h e c o u r s e t h a t I have chosen f o r m y s e l f . W i t h o u t h e r l o v e and p a t i e n c e t h i s p r o j e c t would have been much more d i f f i c u l t t o complete. 1 CHAPTER 1 : OVERVIEW 1 . 1 I n t r o d u c t i o n The s t r i p e d ambrosia b e e t l e , Trypodendron l i n e a t u m ( O l i v i e r ) ( C o l e o p t e r a : S c o l y t i d a e ) , i s the most numerous (S. Wood 1982) and most damaging of the ambrosia b e e t l e s p e c i e s i n B r i t i s h Columbia (Prebble and Graham 1957). These dark brown i n s e c t s , 2.7 - 3.5 mm long and 1.7 mm wide, bore i n t o the sapwood of l o g s , stumps, and windthrown t r e e s o f almost a l l s p e c i e s o f c o n i f e r s (S. Wood 1982). Tree genera e s p e c i a l l y s u s c e p t i b l e t o a t t a c k i n c l u d e Pseudotsuqa, Tsuqa, P i c e a , A b i e s , and Thuja (Prebble and Graham 1957). Trees are g e n e r a l l y s u s c e p t i b l e t o a t t a c k f o r a p e r i o d o f 3 - 12 months f o l l o w i n g t r e e m o r t a l i t y (Dyer and Chapman 1965; Mathers 1935; Prebble and Graham 1957). C h a r a c t e r i s t i c o f the ambrosia b e e t l e group,, a d u l t s of t h i s s p e c i e s i n t r o d u c e a s y m b i o t i c f u n g i i n t o the g a l l e r y , M o n i l i a f e r r u g i n e a Mathiesen-Kaarik, upon which immature and a d u l t b e e t l e s feed ( F i s h e r e t a l . 1953; Francke-Grossman 1963; Funk 1965). Degrade o f sawtimber, from the p i n h o l e s caused by g a l l e r y p r o d u c t i o n and the b l a c k s t a i n from the fungus, alone account f o r more than $63 m i l l i o n (Canadian) y e a r l y l o s s t o the B r i t i s h Columbia f o r e s t i n d u s t r y (McLean 1985). Other c o s t s i n c l u d e l o s s o f export o p p o r t u n i t i e s f o r h i g h q u a l i t y l o g s ; repackaging bundles o f l o g s t o remove a t t a c k e d m a t e r i a l ; m o d i f i c a t i o n o f h a r v e s t i n g p r a c t i c e s , i n v e n t o r y 2 management, and y a r d i n g p r o c e d u r e s ; and a p p l i c a t i o n o f p e s t management s t r a t e g i e s (Borden and McLean 1981). These economic r e a l i t i e s have s t i m u l a t e d a g r e a t d e a l o f r e s e a r c h on t h e l i f e h i s t o r y , b i o l o g y , and c h e m i c a l e c o l o g y o f T. l i n e a t u m (Borden 1988a; N i j h o l t 1979), as w e l l as t h e development o f p e s t management s t r a t e g i e s f o r use i n a t t e m p t i n g t o m i n i m i z e b e e t l e a t t a c k (Borden 1988b; L i n d g r e n e t a l . 1983; Shore and McLean 1985). M a s s - t r a p p i n g b e e t l e and removal o f nearby o v e r w i n t e r i n g s i t e s ( i e . f o r e s t margins) a t l o g s o r t s and booming grounds ( l o g g r a d i n g and s t o r a g e a r e a s ) , i s an i m p o r t a n t c o n t r i b u t i o n t o w a r d r e d u c i n g b e e t l e p o p u l a t i o n s . Y e t i t has n o t and w i l l n o t s o l v e t h e p r o b lem u n l e s s i n v e n t o r y management, wh i c h i n c l u d e s removal o f l o g i n v e n t o r y from t h e f o r e s t as soon as p o s s i b l e , i s p r a c t i c e d . The e v i d e n c e j u s t i f y i n g t h i s recommendation has become a p p a r e n t i n r e c e n t y e a r s . F o r example, Gray and Borden (1985) showed t h a t o f a l l t h e l o g s a t t a c k e d a t a l o g s o r t on n o r t h e r n Vancouver I s l a n d , 45% o f t h e l o g s had come i n t o t h e s o r t a l r e a d y a t t a c k e d by a mbrosia b e e t l e s . I n a s u r v e y o f t h e l o g i n v e n t o r y o f one woodlands d i v i s i o n on Vancouver I s l a n d , 126 000 m 3 o f i n v e n t o r y was l e f t on t h e ground d u r i n g t h e s p r i n g . About 80% o f t h i s m a t e r i a l was a t t a c k e d by a mbrosia b e e t l e s b e f o r e b e i n g t a k e n from t h e f i e l d , r e s u l t i n g i n an e s t i m a t e d l o s s t o t h e f o r e s t company o f o v e r $500 000 (Daust 1985). I n a s t u d y on n o r t h e a s t Vancouver I s l a n d , s e m i o c h e m i c a l - b a i t e d f u n n e l t r a p s ( L i n d g r e n 1983) p l a c e d i n a commercial o l d - g r o w t h c o n i f e r o u s 3 f o r e s t more than 1 .5 km from a r e c e n t l o g g i n g s e t t i n g , c a ptured over 22 000 T. lineatum i n 12 f u n n e l t r a p s d u r i n g the summer (McLean and Salom u n p u b l i s h e d ) . T h i s emphasizes how p r e v a l e n t the b e e t l e s are i n these f o r e s t s and t h e i r i n n a t e a b i l i t y t o f i n d a t t r a c t i v e host m a t e r i a l . D e s p i t e i n c r e a s i n g awareness by p r i v a t e companies of the m e r i t s o f moving t h e i r l o g i n v e n t o r y more q u i c k l y , t h e r e appear t o be some unavoidable f a c t o r s t h a t c o n s t r a i n i n v e n t o r y removal b e f o r e the s p r i n g b e e t l e f l i g h t . These i n c l u d e : the d i f f i c u l t y i n removing r i g h t - o f - w a y l o g s u n t i l roads are completed and s e t t l e d ; l o a d i n g equipment a v a i l a b i l i t y ; manpower; need t o m a i n t a i n a s p r i n g i n v e n t o r y so t h a t workers i n v o l v e d i n l o a d i n g and t r a n s p o r t o p e r a t i o n s can remain employed; m i l l c a p a c i t y ; m a r k e t a b i l i t y o f l o g s p e c i e s ; and weather, such as e a r l y s n o w f a l l o r l a t e snowmelt. I t i s c l e a r t h a t o t h e r woods management o p t i o n s must be co n s i d e r e d . Borden (1988a) recommended t h a t c u t b l o c k s be e s t a b l i s h e d f a r apart from each ot h e r i n s u c c e s s i v e y e a r s , making i t more d i f f i c u l t f o r b e e t l e s t o d i s p e r s e and f i n d s u s c e p t i b l e h o s t s . T h i s c o u l d c e r t a i n l y be c o n s i d e r e d by h a r v e s t p l a n n e r s and would be even more p r a c t i c a l as we s t a r t h a r v e s t i n g second-growth f o r e s t s where road networks w i l l a l r e a d y be e s t a b l i s h e d . However, t h i s approach w i l l be d i f f i c u l t t o f o l l o w without knowing the dynamics of T. lineatum p o p u l a t i o n movement. Previo u s s t u d i e s o f T. line a t u m b i o l o g y and p h y s i o l o g y have p r o v i d e d some 4 understanding of t h e i r f l i g h t p a t t e r n s and c a p a b i l i t i e s d u r i n g the s p r i n g (Bennett and Borden 1971; Chapman 1962; Chapman and Kinghorn 1958; Dyer 1961; Graham 1959; Rudinsky and Daterman 19 64) and l a t e summer-early f a l l d i s p e r s a l p e r i o d s (Dyer and Kinghorn 1961; Kinghorn and Chapman 1959). A s y n o p s i s of t h i s r e s e a r c h i s presented below. In the s p r i n g , when temperatures r i s e above 15°C and the snow on the ground has melted, T. lineatum w i l l b e g i n t o emerge and f l y (Chapman and Kinghorn 1958; Rudinsky and Daterman 1964) from t h e i r o v e r w i n t e r i n g s i t e s i n the f o r e s t d u f f and bark of stumps and t r e e s (Dyer and Kinghorn 1961; Kinghorn and Chapman 1959), i n search of host m a t e r i a l . Mass f l i g h t s , c h a r a c t e r i s t i c o f t h i s s p e c i e s , occur d u r i n g d a y l i g h t hours when temperatures reach near 20°C or above, w i t h optimal f l i g h t temperatures r a n g i n g from 19 - 2 6°C (Rudinsky and Daterman 1964). F l i g h t a c t i v i t y i s h i g h e s t under calm wind c o n d i t i o n s , and i s reduced markedly a t wind speeds of 5 km/h (Chapman 1962; Rudinsky and Daterman 1964), w i t h e s s e n t i a l l y no f l i g h t observed above 6.4 km/h (Chapman 1962). Chapman (1962) a l s o observed t h a t b e e t l e s f l y 3.0 -4.5m above the ground w i t h i n the f o r e s t , u s u a l l y above the u n d e r s t o r y brush (Shore and McLean 1984), dropping t o 1.8 -2.4 m i n exposed areas, and o r i e n t i n g upwind toward nearby a t t r a c t i v e h ost m a t e r i a l (Chapman 1962). With r e s p e c t t o f l i g h t d i s t a n c e c a p a b i l i t i e s , one b r i e f r e f e r e n c e by Dyer (1961) i n d i c a t e d t h a t i n a mark-recapture study, one T. lineatum b e e t l e was r e c a p t u r e d 4 km away from the r e l e a s e 5 p o i n t , a year f o l l o w i n g the r e l e a s e . F u r n i s s and F u r n i s s (1972) c o l l e c t e d specimens of T. lineatum on s n o w f i e l d s , 1200 m above t i m b e r l i n e . The s p r i n g d i s p e r s a l p e r i o d i s dominated by two types of b e h a v i o r . I n i t i a l l y , b e e t l e s tend t o be p h o t o p o s i t i v e and w i l l f l y toward the sun (Graham 1959). The l e n g t h o f time t h i s b e h a v i o r i s e x h i b i t e d v a r i e s g r e a t l y , a v e r a g i n g 30 min b e f o r e the b e e t l e s become a r r e s t e d t o host s t i m u l i (Bennett and Borden 1971). Females are the p i o n e e r b e e t l e s and a re a t t r a c t e d t o s u s c e p t i b l e l o g s by ho s t v o l a t i l e s , of which et h a n o l has been i d e n t i f i e d as the most important (Moeck 1970, 1971). A f t e r females t u n n e l i n t o the host, they w i l l r e l e a s e a ve r y potent a g g r e g a t i o n pheromone, a t r i - c y c l i c k e t a l g i v e n the name l i n e a t i n ( 3 , 3 , 7 - t r i m e t h y l -2 , 9 - d i o x a t r i c y c l o [3.3.1.0 4' 7] nonane) t o which both sexes respond (Borden e t a l . 1979; MacConnell e t a l . 1977). Graham (1959) hypothesized t h a t the i n i t i a l response t o l i g h t by T. lineatum served as a mechanism t o f o r c e a t l e a s t some b e e t l e s i n a p o p u l a t i o n t o d i s p e r s e i n se a r c h o f new hos t s , u s u a l l y s c a r c e i n nature. T h i s i s now c o n s i d e r e d a f a i r l y common beha v i o r f o r many i n s e c t s t h a t d i s p e r s e o r migrate d u r i n g t h e i r l i f e c y c l e (Kennedy 1975). T h i s b e h a v i o r may be r e l e a s e d a t a f a t content t h r e s h o l d below which p h o t o p o s i t i v e dominance i s l o s t and arrestment t o host s t i m u l i o c c u r s , as A t k i n s (1966a, 1969) and Bennett and Borden (1971) proposed f o r the D o u g l a s - f i r b e e t l e , Dendroctonus ponderosae Hopkins. 6 From e a r l y J u l y t o l a t e October, brood a d u l t s and some pare n t s w i l l emerge from t h e i r b r e e d i n g areas and f l y t o o v e r w i n t e r i n g s i t e s (Kinghorn and Chapman 1959). Dyer and Kinghorn (1961) found t h a t b e e t l e s are p h o t o p o s i t i v e when emerging from the wood m a t e r i a l and w i l l f l y ca. 15 m above the ground b e f o r e heading toward the n e a r e s t f o r e s t . They a l s o determined t h a t t h i s p h o t o p o s i t i v e dominance was l o s t as they e n t e r the shade of the f o r e s t . I n t e r e s t i n g l y , and t o the dismay of p e s t managers, f a l l d i s p e r s i n g brood a d u l t s are p r o b a b l y not r e s p o n s i v e t o semiochemicals, as they are not i n c l i n e d t o mate a t t h i s time ( F o c k l e r and Borden 1972). Yet c o n c l u s i v e t e s t s are needed t o v e r i f y t h i s . With r e s p e c t t o f l i g h t c a p a c i t y , f a l l d i s p e r s e r s were found t o f l y f o r much s h o r t e r p e r i o d s than s p r i n g d i s p e r s e r s (Chapman 1956). S i n c e s p r i n g f l y i n g b e e t l e s appear t o be the more important of the two d i s p e r s a l stages (Chapman 1956), and because b e e t l e s from t h i s stage would be e a s i e r t o study, due t o t h e i r p o s i t i v e response t o semiochemicals, I w i l l h e r e a f t e r , c o n c e n t r a t e almost e x c l u s i v e l y on t h i s p e r i o d i n the l i f e c y c l e of T. lineatum. At the p r e s e n t time, b a s i c f l i g h t c h a r a c t e r i s t i c s of T. lineatum. w i t h r e s p e c t t o s e a s o n a l and d i u r n a l p e r i o d s , has been w e l l e s t a b l i s h e d i n both North America (Chapman and Kinghorn 1958; Rudinsky and Daterman 1964) and Europe ( A n n i l a e t a l . 1972), w i t h l i t t l e d i f f e r e n c e o c c u r r i n g between b e e t l e s from both r e g i o n s . However, l i t t l e i s known about the dynamics of p o p u l a t i o n 7 movement. S p e c i f i c a l l y , data are needed c o n c e r n i n g f l i g h t d i s t a n c e and d i r e c t i o n , and d i s p e r s a l r a t e , as they r e l a t e t o wind p a t t e r n s , topography, v e g e t a t i v e cover, and ho s t l o c a t i o n s . A d d i t i o n a l l y , b e e t l e response and f l i g h t o r i e n t a t i o n t o wind and d i f f e r e n t semiochemical components have y e t t o be adequately examined (Borden 1977). The i n f l u e n c e o f wind, as i t i n t e r a c t s w i t h the environment, i s c l e a r l y one of the most important f a c t o r s i n f l u e n c i n g i n s e c t d i s p e r s a l (Johnson 1969; Pedgley 1982). The c o l l e c t i o n of data concerning p o p u l a t i o n movement and f l i g h t o r i e n t a t i o n t o hos t s , can l e a d t o s i g n i f i c a n t p e s t management recommendations f o r h a r v e s t e r s and woodlands managers concerned w i t h ambrosia b e e t l e damage. Cut b l o c k s can be d i s t r i b u t e d s p a t i a l l y t o minimize r i s k o f b e e t l e a t t a c k . A l s o p o p u l a t i o n s u p p r e s s i o n s t r a t e g i e s w i t h semiochemicals, s i m i l a r t o those used a t the l o g s o r t s and booming grounds, may be a p p l i e d i n the f o r e s t s e t t i n g s . As d i s c u s s e d e a r l i e r , the focus o f t h i s problem needs t o be extended from the l o g s o r t and y a r d i n g o p e r a t i o n s i n t o the h a r v e s t areas. 1.2 O b j e c t i v e s F l i g h t o r i e n t a t i o n and response of T. lineatum, as w e l l as o t h e r s c o l y t i d s , has not been adequately i n v e s t i g a t e d i n d e t a i l e d b e h a v i o r a l s t u d i e s (Borden 1977). Lack of such s t u d i e s o f t e n make i n t e r p r e t a t i o n o f b e e t l e t r a p p i n g f i e l d experiments more d i f f i c u l t . C o n t r o l l e d b e h a v i o r s t u d i e s 8 concerning f l i g h t are u s u a l l y c a r r i e d out i n wind t u n n e l s (Baker and L i n n J r . 1984). T h i s b i o a s s a y has been i n h e r e n t l y d i f f i c u l t t o use f o r the study of s c o l y t i d b e e t l e f l i g h t b e h a v i o r ( S c h l y t e r e t a l . 1987a), w i t h o n l y one s u c c e s s f u l b e h a v i o r study p u b l i s h e d (Choudhury and Kennedy 1980). T h e r e f o r e , my f i r s t o b j e c t i v e was t o determine the optimal wind t u n n e l c o n d i t i o n s needed f o r s t u d y i n g T. l i n e a t u m f l i g h t b e havior i n the presence of semiochemicals. P r e l i m i n a r y o b s e r v a t i o n s i n d i c a t e d the importance of wind speed, and s i n c e t h i s f a c t o r had y e t t o be examined, f o r t h i s or any o t h e r s c o l y t i d b e e t l e (Borden 1977), i t was the s u b j e c t of the experiments presented i n Chapter 2 of t h i s r e p o r t . Another c r i t i c a l component f o r d e t e r m i n i n g o p t i m a l wind t u n n e l c o n d i t i o n s i n c l u d e the d e t e r m i n a t i o n of o p t i m a l r e l e a s e r a t e s of o l f a c t o r y s t i m u l i (Baker and L i n n J r . 1984; S c h l y t e r e t a l . 1987b). T h i s l e d t o an experiment, pre s e n t e d i n Chapter 3, t h a t examined the number o f T. l i n e a t u m captured i n f u n n e l t r a p s b a i t e d w i t h v a r i e d r e l e a s e r a t e s of e t h a n o l and l i n e a t i n , alone and i n combination. Current p e s t management procedures f o r T. l i n e a t u m r e l y h e a v i l y on semiochemical-based mass t r a p p i n g programs (Borden 1988b). The use of primary and secondary a t t r a c t a n t s i n t r a p p i n g systems has come under debate i n North America (Borden e t a l . 1982; Salom and McLean 1988; Shore and McLean 1983), which was s t i m u l a t e d by s t u d i e s i n Europe, showing p o s i t i v e response of T. lineatum t o h o s t 9 a t t r a c t a n t s (Bauer and V i t e 1975) t h a t a c t s y n e r g i s t i c a l l y w i t h the a g g r e g a t i o n pheromone l i n e a t i n (Paiva and K i e s e l 1985; V i t e and Bakke 1979). Thus, i t was f e l t t h a t a d e t a i l e d i n v e s t i g a t i o n i n t o the b e h a v i o r a l responses of T. l i neatum from North America t o e t h a n o l and l i n e a t i n , alone and t o g e t h e r , were needed t o a i d i n i n t e r p r e t i n g p r e v i o u s s t u d i e s , as w e l l as s e t a c l e a r e r path f o r f u r t h e r improvements i n t r a p p i n g technology. T h i s i n v e s t i g a t i o n i s p r e s e n t e d i n Chapter 4. Our l a c k of knowledge concerning p o p u l a t i o n movement of T. l ineatum prompted me t o c a r r y out a s e r i e s of f i e l d experiments of i n c r e a s i n g complexity, i n an e f f o r t t o c h a r a c t e r i z e b e e t l e f l i g h t p a t t e r n s i n the s p r i n g w i t h r e s p e c t .to wind, topography, l a n d cover, and l o c a t i o n of o l f a c t o r y s t i m u l i . Such f i e l d experiments g e n e r a l l y employ some form of mark-recapture techniques, which can i n c l u d e marking i n s e c t s w i t h f l u o r e s c e n t powders and p a i n t s , l a b e l s , f e e d i n g w i t h dyes, genotype d i f f e r e n t i a t i o n , and r a d i o a c t i v e i s o t o p e s (Southwood 1978). The most common form of marking technique used f o r bark and wood b o r i n g b e e t l e s has been the use of f l u o r e s c e n t powders (Helland e t a l . 1984; L i n t o n e t a l . 1987; Schmitz 1979; Shore and McLean 1988). T h i s technique seemed s u i t a b l e f o r use i n these f i e l d s t u d i e s . The f i r s t s e t of f i e l d experiments was designed w i t h the f o l l o w i n g o b j e c t i v e s : 1) develop competence i n u s i n g mark-recapture as a t o o l f o r s t u d y i n g T. l i n e a t u m movement; and 2) c h a r a c t e r i z e s h o r t (5 - 25 m), i n t e r m e d i a t e (100 m), 10 and long d i s t a n c e (500 m) b e e t l e f l i g h t i n a second-growth c o n i f e r o u s f o r e s t . T h i s work i s presented i n Chapter 5. In a second s e t of f i e l d experiments, pr e s e n t e d i n Chapter 6, the study s i t e was moved t o a v a l l e y s e t t i n g , more t y p i c a l of commercial f o r e s t s i t u a t i o n s a l o n g the c o a s t of B r i t i s h Columbia. Some of the same s h o r t d i s t a n c e f l i g h t p a t t e r n s were examined again, however, t h i s time w i t h i n an old-growth c o n i f e r o u s f o r e s t . A d d i t i o n a l l y , emphasis was p l a c e d on T. lineatum long d i s t a n c e f l i g h t c a p a b i l i t i e s and r a t e of d i s p e r s a l . The l a s t two f i e l d experiments, presented i n Chapter 7, were c a r r i e d out i n the same v a l l e y as were the p r e v i o u s s e t of experiments. They were designed t o c h a r a c t e r i z e l o n g d i s t a n c e T. lineatum movement, without the presence of semiochemicals near the r e l e a s e s i t e s , and w i t h equal emphasis p l a c e d on up- and down-valley b e e t l e movement. Trap c a t c h comparisons were made between f o r e s t and open s e t t i n g s , and use of roads as flyways by b e e t l e s was i n v e s t i g a t e d . Equal numbers of b e e t l e s were r e l e a s e d s i m u l t a n e o u s l y from two separate r e l e a s e p o i n t s , a l l o w i n g f o r a b e t t e r r e p r e s e n t a t i o n of b e e t l e d i s p e r s a l throughout the v a l l e y (Banks e t a l . 1988). 1.3 Format Chapters 2 - 7 have been w r i t t e n up as s e p a r a t e papers i n p u b l i c a t i o n format. They should be a b l e t o stand alone 11 except a l l r e f e r e n c e s are l i s t e d a f t e r Chapter 8. Chapters 2 - 4 are w r i t t e n f o r j o u r n a l s t h a t separate the r e s u l t s and d i s c u s s i o n s e c t i o n s . In c o n t r a s t , c h a p t e r s 5 - 7 are w r i t t e n f o r j o u r n a l s t h a t combine these two s e c t i o n s . A l l ot h e r a s p e c t s of the format between cha p t e r s w i l l be c o n s i s t e n t throughout the d i s s e r t a t i o n . While I have attempted t o exclude redundant p r e s e n t a t i o n s and d i s c u s s i o n s between the ch a p t e r s , i t was i m p o s s i b l e t o remove a l l of i t completely. H o p e f u l l y t h i s w i l l not d e t r a c t too much from the r e a d i n g . 12 CHAPTER 2: THE EFFECT OF WIND SPEED ON TRYPODENDRON LINEATUM FLIGHT BEHAVIOR IN A WIND TUNNEL 2.1 I n t r o d u c t i o n O r i e n t a t i o n behavior of f l y i n g i n s e c t s i n response t o semiochemicals has been a s u b j e c t of r e c e n t i n v e s t i g a t i o n s and reviews ( B e l l and Carde 1984; Borden 1977; Farkas and Shorey 1974; Kennedy 1978; Payne e t a l . 1986; Shorey 1973). O l f a c t o r y o r i e n t a t i o n has been shown t o v a r y w i t h r e s p e c t t o chemical g r a d i e n t s , wind, l i g h t , as w e l l as p h y s i o l o g i c a l requirements and c a p a b i l i t i e s of the i n s e c t . Upwind f l i g h t i s g e n e r a l l y accepted as the behavior of most i n s e c t s i n response t o the d e t e c t i o n of a t t r a c t i v e m a t e r i a l (Baker and L i n n J r . 1984). S e v e r a l s p e c i e s of s c o l y t i d b e e t l e s have demonstrated upwind f l i g h t t o a t t r a c t i v e m a t e r i a l i n the f i e l d (Botterweg 1982; Byers 1988; Chapman 1962; Gara 1963; McMullen and A t k i n s 1962;). Under more c o n t r o l l e d c o n d i t i o n s , wind t u n n e l s have become a v a l u a b l e t o o l f o r d e c i p h e r i n g the complex f l i g h t p a t t e r n s of i n s e c t s (mostly Lepidoptera) (Baker and L i n n J r . 1984). They al l o w the i n v e s t i g a t o r t o t e s t f o r i n d i v i d u a l e f f e c t s under a r e l a t i v e l y r i g i d experimental d e s i g n and procedure not p o s s i b l e i n the f i e l d . While upwind o r i e n t a t i o n t o semiochemicals has been demonstrated f o r many s p e c i e s of L e p i d o p t e r a ( P r e i s s and Kramer 1986), i t has been demonstrated f o r o n l y two s p e c i e s of s c o l y t i d s (Choudhury and Kennedy 1980; Lindgren and McLean, unp u b l i s h e d d a t a ) , 13 one a n o b i i d s p e c i e s ( B i r c h and White, 1988), and one c l e r i d s p e c i e s ( M i z e l l e t a l . 1984). The s t r i p e d ambrosia b e e t l e , Trypodendron li n e a t u m ( O l i v i e r ) , uses semiochemicals t o f i n d s u i t a b l e h o s t s f o r c o l o n i z a t i o n , mating, and r e p r o d u c t i o n . During s p r i n g f l i g h t d i s p e r s a l , T. lineatum must d e t e c t and respond t o s u i t a b l e host m a t e r i a l , mainly c o n i f e r o u s l o g s t h a t have been h a r v e s t e d f o r a t l e a s t 3 months (Mathers 1935). Temperature, time of day, and wind d i r e c t i o n are known t o a f f e c t g e n e r a l f l i g h t a c t i v i t y , but l i t t l e i s known about c l o s e - r a n g e f l i g h t b e h a v i o r t o host m a t e r i a l , w i t h r e s p e c t t o v a r i e d environmental c o n d i t i o n s f o r t h i s i n s e c t as w e l l as f o r e s t C o l e o p t e r a i n g e n e r a l (Borden e t a l . 1986; Carde 1984). Two wind t u n n e l s t u d i e s presented here examine the e f f e c t wind speed has on clo s e - r a n g e response of T. lineatum t o a t t r a c t i v e s u b s t r a t e s . The f i r s t study was designed t o determine the wind speed a t which the h i g h e s t p r o p o r t i o n of b e e t l e s r e l e a s e d would respond p o s i t i v e l y t o a semiochemical-baited f u n n e l t r a p . I t was h y p o t h e s i z e d t h a t the h i g h e s t p r o p o r t i o n of r e l e a s e d b e e t l e s would be r e c a p t u r e d i n the b a i t e d t r a p s p l a c e d upwind, a t the lower end o f the wind speed range (0.0 t o 0.9 m/s) t e s t e d . A second study was designed t o see i f the mechanisms by which T. lineatum o r i e n t e d t o a semiochemical-baited s u b s t r a t e d i f f e r e d throughout the same range of wind speeds t h a t were t e s t e d i n study 1. One hyp o t h e s i s t e s t e d was t h a t c h e m i c a l l y - i n d u c e d anemotaxis would occur when wind i s p r e s e n t and a second h y p o t h e s i s was t h a t a d i f f e r e n t o r i e n t a t i o n mechanism f o r a t t r a c t i o n i s used by the i n s e c t when wind i s absent. 2 . 2 M a t e r i a l s and Methods Both s t u d i e s were conducted i n a wind t u n n e l ( A n g e r i l l i and McLean 1984) 3.6m long and 1.2 m h i g h and wide. Cool white f l u o r e s c e n t l i g h t s (660 W) s e t 1.5 m above the wind t u n n e l s i m u l a t e d d i u r n a l c o n d i t i o n s and f a c i l i t a t e d d i r e c t o b s e r v a t i o n s of f l y i n g b e e t l e s . A sheet o f c e l l u l o s e a c e t a t e was p l a c e d over the top of the t u n n e l t o d i f f u s e the incoming l i g h t and reduce g l a r e . I n c i d e n t i l l u m i n a t i o n r e a d i n g s from w i t h i n the t u n n e l were measured wi t h a Sekonic® model L-398 exposure meter. V e r t i c a l l i g h t measured 0.8 m from the c e i l i n g i n the exhaust, middle, and wind entrance s e c t i o n s of the t u n n e l were 112, 125, and 140 l x , r e s p e c t i v e l y . Thus, a s l i g h t l i g h t g r a d i e n t from the c e i l i n g was pr e s e n t . B e e t l e s were c o l l e c t e d d a i l y d u r i n g May and June of 1986 and 1988, i n l i n e a t i n - b a i t e d , m u l t i p l e - f u n n e l t r a p s , near a l a r g e l o g boom storage area a t the mouth of the North arm o f the F r a s e r R i v e r , B r i t i s h Columbia. Captured b e e t l e s were s t o r e d a t 4°C, under a 14:10 h (L:D) pho t o p e r i o d , i n 1 L p l a s t i c j a r s c o n t a i n i n g s l i g h t l y moistened absorbent c l o t h towels. The c o n t a i n e r s were checked twice weekly t o monitor 15 moisture l e v e l s . B e e t l e s s u r v i v e d f o r more than 100 days under these c o n d i t i o n s . S i n c e the b e e t l e s were captured i n pheromone t r a p s , they were judged t o have had s u f f i c i e n t f l i g h t e x e r c i s e (Graham, 1959; Bennett & Borden, 1971) t o ensure subsequent response t o semiochemicals. Study 1 A m u l t i p l e - f u n n e l t r a p , p l a c e d i n the c e n t e r of the upwind s e c t i o n of the t u n n e l , ca. 0.6 m from the screen, was b a i t e d w i t h two l i n e a t i n Biolures® (Consep Membranes, Bend, Oregon) i n the upper and lower p o r t i o n of the t r a p , and a d i s p e n s e r c o n t a i n i n g 95% ethanol i n the middle. Release r a t e s , determined at 2 0°C i n the l a b o r a t o r y were 100 - 12 0 ug/24 h f o r each l i n e a t i n d i s p e n s e r and 75 mg/24 h f o r the e t h a n o l d i s p e n s e r (B.S. Lindgren, p e r s o n a l c o mmunication)^ The l u r e s and r e l e a s e r a t e s were i d e n t i c a l t o those used i n mass-trapping programs of T. lineatum throughout southwestern B.C. A r e l e a s e p l a t f o r m c o n s i s t i n g of a c i r c u l a r , f l a t g l a s s p l a t e 13 cm i n diam, was suspended h o r i z o n t a l l y 35 cm above the t u n n e l f l o o r and 1 m from the downwind scr e e n . P r e l i m i n a r y t r i a l s suggested t h a t optimal response by T. l i n e a t u m o c c u r r e d when the r e l e a s e p l a t f o r m was p l a c e d a t t h i s h e i g h t . I t was h i g h enough f o r the b e e t l e s t o be c o n t i n u o u s l y exposed t o the semiochemical plume p r i o r t o t a k e - o f f , y e t c l o s e enough t o the s t r i p e d f l o o r t o use l l . Research D i r e c t o r , Phero Tech. Inc., 1140 C l a r k Dr., Vancouver, B.C. V5L 3K3. 16 optomotor cues i f needed. The g l a s s p l a t e was covered w i t h a white i n d u s t r i a l paper towel t o enhance t r a c t i o n f o r the i n s e c t s . A 16 cm 2 x 0.7 cm deep p l a s t i c weighing boat, covered by the same m a t e r i a l , was taped t o the upper s i d e of the g l a s s p l a t e . T h i s allowed the b e e t l e s two p e r c h i n g o p p o r t u n i t i e s ; the f i r s t around the edge of the weighing boat, and i f they f e l l o f f t h a t , a second o p p o r t u n i t y around the edge of the g l a s s p l a t e . B e e t l e s were sexed and g i v e n a walking t e s t i n the morning p r i o r t o f l i g h t i n the wind t u n n e l . They were allowed t o a c c l i m a t e t o the warmer room temperatures (22 -25°C) f o r 10 min b e f o r e r e l e a s e , t o ensure maximal response of the b e e t l e s t o the t r a p . A n i n e - r e p l i c a t e , randomized complete b l o c k d e s i g n was used, i n which the treatments were wind speeds of 0.0, 0.15, 0.30, 0.45, 0.60, 0.75, and 0.90 m/s (0 - 3.2 km/h). For each treatment, males and females were t e s t e d s e p a r a t e l y . Groups of 50 males and 30 - 50 females were p l a c e d on the r e l e a s e t r a y . A l l treatments w i t h i n one b l o c k ( r e p l i c a t i o n ) were run f o r each sex on the same day, and the experiment was run on 9 days over a 21 day p e r i o d . Each r e p l i c a t i o n f o r a g i v e n treatment l a s t e d f o r 20 min. B e e t l e s were never used more than once. An a d d i t i o n a l t h r e e - r e p l i c a t e , c o n t r o l experiment assessed male and female catches a t 0.0 m/s i n u n b a i t e d t r a p s . P r i o r t o these r e p l i c a t e s , the f u n n e l t r a p , t u n n e l , 17 and r e l e a s e t r a y were cleaned w i t h a 0.5% sodium h y p o c h l o r i t e s o l u t i o n . With treatment v a l u e s s e t a t equal i n t e r v a l s throughout the wind speed range t e s t e d , the data were analyzed u s i n g o r t h o g o n a l polynomials, f o l l o w e d by r e g r e s s i o n a n a l y s i s . For males, the v a r i a n c e s d e v i a t e d s i g n i f i c a n t l y from homogeneity, u s i n g Box's t e s t (Fox and G u i r e 1976); thus a weighted r e g r e s s i o n a n a l y s i s was conducted, w i t h the weights e q u a l i n g the r e c i p r o c a l s of the e r r o r v a r i a n c e s (SAS 1985). Male and female r e g r e s s i o n equations were then compared u s i n g an F - t e s t of c o i n c i d e n t a l r e g r e s s i o n s (Zar 1984). Study 2 A p r e l i m i n a r y study was conducted t o determine the number of b e e t l e s l a n d i n g i n or on v a r i o u s semiochemical-b a i t e d s u b s t r a t e s . These i n c l u d e d a 28 cm diam x 71 cm long l o g o f western hemlock, Tsuga h e t e r o p h y l l a (Raf.) Sarg., o r i e n t e d e i t h e r v e r t i c a l l y , or h o r i z o n t a l l y a c r o s s the width of the t u n n e l ; a simulated l o g (SL) of the same dimensions as the r e a l l o g , made of cardboard, c o t t o n f i b e r t a g paper, and c o n s t r u c t i o n paper, o r i e n t e d i n the same two ways as the r e a l l o g ; and a f u n n e l t r a p . The b e e t l e s showed no p r e f e r e n c e f o r any s u b s t r a t e or o r i e n t a t i o n (Appendix 1). T h e r e f o r e a h o r i z o n t a l SL was used t o assess the e f f e c t of wind speed on f l i g h t behavior, because i t allowed f o r c o n t r o l of r e l e a s e d v o l a t i l e s , not p o s s i b l e w i t h a r e a l l o g , and assumed the p o s i t i o n of most l o g s i n the f i e l d . The SL 18 was p l a c e d on two concre t e b l o c k s w i t h h o l e s t h a t reduced the d i s r u p t i o n of a i r flow. Two l i n e a t i n l u r e s and an et h a n o l d i s p e n s e r , i d e n t i c a l t o those used i n study 1, were hooked t o the SL along the upwind s i d e , p r o v i d i n g a l a r g e , r e l a t i v e l y even downwind plume, as v e r i f i e d by r e l e a s i n g T i C l 4 smoke a t the d i f f e r e n t wind speeds. The r e l e a s e t r a y and i t s placement i n the t u n n e l were the same as i n study 1. To study T. lineatum f l i g h t behavior, the o v e r a l l experiment was d i v i d e d up i n t o two p a r t s . P a r t 1 examined b e e t l e responses t o a SL b a i t e d w i t h l i n e a t i n and e t h a n o l , w h i l e p a r t 2 examined b e e t l e responses t o an u n b a i t e d SL, s e r v i n g as a c o n t r o l . Each p a r t examined b e e t l e responses a t wind speeds of 0.0, 0.3, 0.6, 0.9 m/s. P a r t s 1 and 2 were conducted i n 1986 and 1988, r e s p e c t i v e l y . The t u n n e l and the r e l e a s e t r a y were c a r e f u l l y c l e a n e d w i t h a 0.5% sodium h y p o c h l o r i t e s o l u t i o n p r i o r t o running p a r t 2. The 0.0 m/s treatment i n p a r t 1 was d i v i d e d i n t o two treatments, one i n an unaerated t u n n e l and the ot h e r i n the same t u n n e l a e r a t e d f o r 10 min. T h i s was c a r r i e d out t o determine i f contamination o c c u r r e d i n the unaerated t u n n e l . No d i f f e r e n c e s were observed between the two treatments, and as a r e s u l t , the data were pooled, and are pre s e n t e d as one treatment h e r e a f t e r . A l l ot h e r procedures used i n conducting both p a r t s of the experiment were the same. During the experiment, two b e e t l e s of the same sex were r e l e a s e d per r e p l i c a t e and observed f o r up t o 8 min. F i f t y r e p l i c a t e s were run f o r each l e v e l o f wind speed and b a i t 19 t e s t e d . Observations i n c l u d e d occurrence o f f l i g h t , body o r i e n t a t i o n p r i o r t o and d u r i n g t a k e - o f f ( F i g . 1), t a k e - o f f d i r e c t i o n ( F i g . 1), o v e r a l l f l i g h t d i r e c t i o n , type o f f l i g h t , and loc k - o n behavior. O v e r a l l f l i g h t d i r e c t i o n was scored as uptunnel, downtunnel, or n o n - d i r e c t i o n a l , depending upon the predominant d i r e c t i o n flown by the i n s e c t throughout the o b s e r v a t i o n p e r i o d . I f b e e t l e s d i d not e x h i b i t predominant up- or downtunnel f l i g h t , they were c o n s i d e r e d n o n - d i r e c t i o n a l f l i e r s . Type of f l i g h t was scored as: e r r a t i c - quick, sharp changes i n movement, o f t e n o c c u r r i n g when b e e t l e s bounce o f f the t u n n e l w a l l s ; steady -a s e t t l e d , s t a b l e movement t y p i c a l o f f l i g h t i n the f i e l d ( p e r s o n a l o b s e r v a t i o n ) ; and d i r e c t - a f a s t , s t r a i g h t - l i n e f l i g h t t o some endpoint. Lock-on be h a v i o r can be d e s c r i b e d by steady f l i g h t i n an upwind d i r e c t i o n , o r uptunnel toward the l o g i n the absence of an a i r f l o w , w i t h the i n s e c t a p p a r e n t l y o r i e n t i n g s p e c i f i c a l l y t o the SL, e i t h e r by c a s t i n g up and downtunnel, h o v e r i n g around the SL, or l a n d i n g on i t . A l l o b s e r v a t i o n s were recorded on a c a s s e t t e tape and l a t e r t r a n s c r i b e d on t o data sheets. The data were summarized as f r e q u e n c i e s and analyzed s e p a r a t e l y f o r each b e h a v i o r a l category, as m u l t i d i m e n s i o n a l contingency t a b l e s u s i n g the CATMOD procedure i n SAS (1985). T h i s procedure uses a l i n e a r models approach t o the a n a l y s i s of c a t e g o r i c a l data, a p p l y i n g g e n e r a l w e i g h t e d - l e a s t squares r e g r e s s i o n techniques t o estimates o f a p p r o p r i a t e ( l i n e a r and/or l o g l i n e a r ) f u n c t i o n s of the c e l l p r o p o r t i o n s i n ORIENTATION PRIOR T O T A K E - O F F W I N D D I R E C T I O N FLIGHT T A K E - O F F DIRECTION NO DIRECTION F i g . 1. Body o r i e n t a t i o n d i r e c t i o n s o f T . l i n e a t u m s c o r e d p r i o r t o and d u r i n g f l i g h t t a k e - o f f , as viewed from above t h e r e l e a s e t r a y ( t o p ) ; and f l i g h t t a k e - o f f d i r e c t i o n s o f t h e same b e e t l e s , as viewed from the s i d e (bottom). A l l b e e t l e s t a k i n g o f f t h a t d i d not f i t i n c a t e g o r i e s A t o D were p l a c e d i n t h e n o - d i r e c t i o n c a t e g o r y . 21 complex c a t e g o r i c a l data l a y o u t s (Kleinbaum & Kupper, 1978). T h i s approach allowed me t o t e s t f o r f i r s t and second-order i n t e r a c t i o n s , main e f f e c t s , and c o n t r a s t between l e v e l s of main e f f e c t s , f o r a s p e c i f i c response, u s i n g the c h i - s q u a r e (X 2) s t a t i s t i c . 2.3 R e s u l t s Study 1 The r e l a t i o n s h i p between the numbers o f T. l i n e a t u m caught a t d i f f e r e n t wind speeds was s i m i l a r f o r both sexes; the h i g h e s t numbers of b e e t l e s were caught a t 0.0 m/s. The p r o p o r t i o n of males and females caught were 21.4 and 25.3%, r e s p e c t i v e l y ( F i g . 2). The p e r c e n t response decreased l i n e a r l y as wind speed i n c r e a s e d . However, much of the v a r i a t i o n i n these data i s not e x p l a i n e d by the r e g r e s s i o n e q u a t ions, w i t h r 2 v a l u e s of 0.28 f o r both sexes. The h i g h v a r i a t i o n i n these r e s u l t s may be due i n p a r t t o the v a r i e d p h y s i o l o g i c a l s t a t e s (e.g. v a r y i n g energy r e s e r v e s ) of the b e e t l e s , r e s u l t i n g i n a wide range of b e h a v i o r a l responses, or the c o n d i t i o n s i n the wind t u n n e l were not o p t i m a l f o r T. lineatum response, or both. The s l o p e s and i n t e r c e p t s of the r e g r e s s i o n equations were not s i g n i f i c a n t l y d i f f e r e n t ( F120,115 = °- 9 8 6'* P > 0.05) from each other, i n d i c a t i n g no d i f f e r e n c e i n response between sexes. B e e t l e responses t o the u n b a i t e d c o n t r o l treatment a t 0.0 m/s averaged o n l y 3.3% f o r males and 2.0% f o r females ( F i g . 2, b l a c k boxes), 22 .50 MALES 40 Y = 21.4 - 17.2X r 2 = 0.2807 P < 0.0001 30 5! O < U W 20 HI 10 -'t2 tu CQ Q W to I o 60 F EMALES 50 -ii-2 40 Y = 253 - 19.7X r 2 = 0.2753 P < 0.0001 WIND SPEED (M/S) F i g . 2 The i n f l u e n c e o f wind speed on response o f male ( t o p ) and female ( b o t t o m ) T. l i n e a t u m t o a s e m i o c h e m i c a l - b a i t e d funne l t r a p i n a wind t u n n e l . Numbers next t o boxes r e p r e s e n t the number o f d a t a p o i n t s . White and b lack boxes rep resen t the % response f o r the b a i t e d and u n b a i t e d ( c o n t r o l ) t r e a t m e n t s , r e s p e c t i v e l y . M i s s i n g d a t a accounts f o r some wind speeds hav ing less than 9 o b s e r v a t i o n s . 23 i n d i c a t i n g l a c k o f c o n t a m i n a t i o n i n t h e t u n n e l and l i t t l e v i s u a l a t t r a c t i o n t o t h e t r a p under t h e s e c o n d i t i o n s . Study 2 The number o f b e e t l e s t a k i n g f l i g h t was n o t i n f l u e n c e d by w i n d speed ( T a b l e 1 ) . Frequency o f f l i g h t was a n a l y z e d by sex and b a i t s e p a r a t e l y because o f a s i g n i f i c a n t i n t e r a c t i o n o c c u r r i n g between t h e s e f a c t o r s ( X ^ = 17.1; p < 0.05), w i t h males r e s p o n d i n g a t l o w e r f r e q u e n c i e s t h a n f e m a l e s t o a b a i t e d SL. Body o r i e n t a t i o n p r i o r t o and d u r i n g f l i g h t t a k e - o f f d i d n o t d i f f e r between males and f e m a l e s , t h e r e f o r e t h e d a t a f o r t h e sexes were p o o l e d . U p t u n n e l body o r i e n t a t i o n i n c r e a s e d s i g n i f i c a n t l y i n t h e p r e s e n c e o f an a i r f l o w f o r b o t h b a i t e d and u n b a i t e d t r e a t m e n t s ( F i g . 3 ) . The p r e s e n c e o f s e m i o c h e m i c a l s d i d n o t r e s u l t i n i n c r e a s e d u p t u n n e l o r i e n t a t i o n , b u t i t d i d r e s u l t i n some i n c r e a s e i n t h e o c c u r r e n c e o f n o n - d i r e c t i o n a l o r i e n t a t i o n , s u g g e s t i n g more a c t i v e movement on t h e r e l e a s e t r a y . T a k e - o f f d i r e c t i o n o f T. l i n e a t u m d i d n o t d i f f e r between males and f e m a l e s , and t h e r e f o r e t h e d a t a f o r t h e sexes were p o o l e d . A t 0.0 m/s, t a k e - o f f d i r e c t i o n was b a s i c a l l y u p t u n n e l / v e r t i c a l , c h a n g i n g t o a d o w n t u n n e l / v e r t i c a l d i r e c t i o n a t 0.3 - 0.6 m/s, and t h e n dominated by downtunnel t a k e - o f f a t 0.9 m/s i n b o t h b a i t e d and u n b a i t e d t r e a t m e n t s ( F i g . 4 ) . A h i g h e r p e r c e n t a g e o f T. l i n e a t u m appear t o t a k e - o f f u p t u n n e l i n t h e p r e s e n c e o f T a b l e l . Frequency o f Trypodendron l i n e a t u m f l i g h t i n a wind t u n n e l a t v a r i e d wind speeds i n t h e p r e s e n c e o f an u n b a i t e d and s e m i o c h e m i c a l -b a i t e d s i m u l a t e d l o g . 1 , 2 Wind speed % F l y i n g (m/s) M a l e s Females U n b a i t e d 0.0 61 67 0.3 70 74 0.6 77 68 0.9 70 80 B a i t e d 0.0 56 72 0.3 44 74 0.6 47 76 0.9 45 67 100 b e e t l e s r e l e a s e d f o r e v e r y l e v e l o f w i n d speed and b a i t t r e a t m e n t (50 f o r each s e x ) . No s i g n i f i c a n t d i f f e r e n c e s i n number o f b e e t l e s f l y i n g , where each sex by b a i t t r e a t m e n t were a n a l y z e d s e p a r a t e l y ( X 2 ; P > 0.05). a a B z w O w to Ed Z o b a a _a 80 -, 60 40 20 o TOTAL NUMBER OF BEETLES OBSERVED WINDSPEED (M/S) UNBAITED BAITED 0.0 111 112 0.3 125 113 0.6 135 108 0.9 137 100 b b a a b b b be b ab ab a a A J , — i — i — i — i — 0.0 0.6 0.3 0.9 — i 1 1 1 — 0.0 0.6 03 0.9 — i 1 1 1 — 0.0 0.6 03 0.9 — i 1 1 r 0.0 0.6 BAITED 0.3 0.9 UPTUNNEL CROSSTUNNEL DOWNTUNNEL NO DIRECTION BODY ORIENTATION PRIOR TO TAKE - OFF UNBAITED WIND SPEED (M/S) F i g . 3. Body o r i e n t a t i o n o f T. l i n e a t u m , p r i o r t o and d u r i n g f l i g h t t a k e - o f f , a t d i f f e r e n t w i n d speeds, i n t h e presence o f an u n b a i t e d and s e m i o c h e m i c a l -b a i t e d s i m u l a t e d l o g . Male and female d a t a were p o o l e d . Bars w i t h d i f f e r e n t l e t t e r s f o r each r e s p o n s e by b a i t t r e a t m e n t c a t e g o r y a r e s i g n i f i c a n t l y d i f f e r e n t ( X 2 ; p < 0.05) f o r i n d i v i d u a l comparisons. B z w C W as (A w z o TOTAL NUMBER OF BEETLES OBSERVED WINDSPEED (M/S) UNBAITED BATTED 0.0 118 116 0.3 135 114 0.6 135 118 0.9 139 106 BAITED ~i 1 1 r 0.0 0.6 0.3 0.9 UPTUNNEL ~l 1 1 r 0.0 0.6 0.3 0.9 UPTUNNEL/ VERTICAL i r 0.0 0.6 03 0.9 DOWNTUNNEL/ VERTICAL UNBAITED ~i r r 0.0 0.6 WIND SPEED 0.3 0.9 ( M / S ) DOWNTUNNEL FLIGHT TAKE - OFF DIRECTION F i g . 4 . F l i g h t t a k e - o f f d i r e c t i o n o f T. l i n e a t u m , a t d i f f e r e n t wind speeds, i n t h e p r e s e n c e o f an u n b a i t e d and s e m i o c h e m i c a l - b a i t e d s i m u l a t e d l o g . Male and female d a t a were p o o l e d . Bars w i t h d i f f e r e n t l e t t e r s f o r each response by b a i t t r e a t m e n t c a t e g o r y a r e s i g n i f i c a n t l y d i f f e r e n t ( X 2 ; p < 0.05) f o r i n d i v i d u a l c o m p a r i s o n s . N3 27 semiochemicals than without them, but t h i s c o u l d not be t e s t e d s t a t i s t i c a l l y because of the occurrence of a s i g n i f i c a n t wind speed by b a i t i n t e r a c t i o n (X g = 23.5; p < 0.05). O v e r a l l f l i g h t d i r e c t i o n changed from uptunnel and non-d i r e c t i o n a l t o downtunnel upon exposure t o wind f o r both the b a i t e d and u n b a i t e d treatments ( F i g . 5). Increases i n wind speed above 0.3 m/s d i d not cause any f u r t h e r change i n f l i g h t d i r e c t i o n f o r b a i t e d treatments, but d i d cause an i n c r e a s e i n downwind f l i g h t f o r u n b a i t e d treatments. A h i g h e r frequency of b e e t l e s e x h i b i t e d uptunnel f l i g h t i n the presence of semiochemicals, w i t h s t a t i s t i c a l d i f f e r e n c e s o c c u r r i n g a t windspeeds of 0.3 ( X 2 2 = 7.1; p < 0.05), 0.6 ( X 2 2 = 10.9; p < 0.05), and 0.9 m/s ( X 2 2 = 8.8; p < 0.05). F l i g h t type data show s i g n i f i c a n t speed by b a i t ( X 2 6 = 20.3; p < 0.05) and speed by sex ( X 2 6 = 16.4; p < 0.05) i n t e r a c t i o n s . The frequency of e r r a t i c f l i g h t d i s p l a y e d by female b e e t l e s i n response t o both b a i t e d and u n b a i t e d SI/s. decreased s i g n i f i c a n t l y as wind speed i n c r e a s e d from 0.6 t o 0.9 m/s ( F i g . 6). The frequency of e r r a t i c f l i g h t o f male b e e t l e s v a r i e d s i g n i f i c a n t l y w i t h no apparent p a t t e r n t o changing wind speed i n response t o the b a i t e d SL, but was unchanged i n response t o the u n b a i t e d treatment ( F i g . 6). At a wind speed of 0.0 m/s, steady f l i g h t o f both sexes was more fre q u e n t i n response t o the b a i t e d treatment than t o the u n b a i t e d treatment, however t h i s d i f f e r e n c e between b a i t e d and u n b a i t e d treatments d e c l i n e d a t h i g h e r wind WINDSPEED (M/S) TOTAL NUMBER OF BEETLES OBSERVED UNBAITED BATTED 0.0 0.3 0.6 0.9 117 145 144 148 128 118 123 112 100 50 b_b_ A b b O S> C=J — I 1 1 r — 0.0 03 0.6 0.9 UPTUNNEL 0 b ab, bj — i 1 1 1 — 0.0 0.3 0.6 0.9 DOWNTUNNEL • B B / BAITED b b b • S B f UNBAITED — i 1 1 r 0.0 0.3 0.6 0.9 WINDSPEED (M/S) NO DIRECTION OVERALL FLIGHT DIRECTION O v e r a l l f l i g h t d i r e c t i o n e x h i b i t e d by T. l i n e a t u m , a t d i f f e r e n t wind speeds, i n t h e p r e s e n c e o f an u n b a i t e d and s e m i o c h e m i c a l - b a i t e d s i m u l a t e d l o g . Male and female d a t a were p o o l e d . Bars w i t h d i f f e r e n t l e t t e r s f o r each r e s p o n s e by b a i t t r e a t m e n t c a t e g o r y a r e s i g n i f i c a n t l y d i f f e r e n t (X ; p < 0.05) f o r i n d i v i d u a l comparisons. TOTAL NUMBER OF BEETLES OBSERVED MALES WTNDSPEED (M/S) UNBAITED BATTED FEMALES UNBAITED BAITED 0.0 0.3 0.6 0.9 61 64 75 65 56 40 44 41 66 72 64 74 69 74 73 67 a a i 1 1 r 0.0 03 0.6 0.9 STEADY ~i r 0.0 03 0.6 0.9 ERRATIC BAITED FEMALES BAITED MALES UNBAITED FEMALES UNBAITED MALES WIND SPEED (M/S) DIRECT TYPE OF FLIGHT Fig. 6. Type of f l i g h t exhibited by T. lineatum. at different wind speeds, in the presence of an unbaited and semiochemical-baited M simulated log. Bars with different l e t t e r s for each response by bait treatment by sex category are s i g n i f i c a n t l y different (X ; p < 0.05) for individual comparisons. 30 speeds. F o r b a i t e d males, i t was an a b r u p t change, whereas f o r f e m a l e s , i t was g r a d u a l , w i t h s i m i l a r f r e q u e n c i e s o f s t e a d y f l i g h t n o t o c c u r r i n g u n t i l w i nd speeds r e a c h e d 0.6 m/s. These d i f f e r e n c e s c o u l d n o t be t e s t e d s t a t i s t i c a l l y because o f t h e speed by b a i t i n t e r a c t i o n , b u t t h e y were a p p a r e n t when p l o t t e d two d i m e n s i o n a l l y on a r e c t a n g u l a r c o o r d i n a t e system.• Observed d e c r e a s e s i n e r r a t i c and s t e a d y f l i g h t f o r b o t h males and f e m a l e s , as wind speed i n c r e a s e d , r e s u l t e d from marked i n c r e a s e s i n d i r e c t f l i g h t , e s p e c i a l l y n o t i c e a b l e from 0.6 t o 0.9 m/s. No d i f f e r e n c e s i n p o s i t i v e l o c k - o n r e s p o n s e s were o b s e r v e d between sexes w i t h r e s p e c t t o wind speed and b a i t ; t h e r e f o r e male and female d a t a were p o o l e d . ( F i g . 7 ) . The f r e q u e n c y o f T. l i n e a t u m e x h i b i t i n g l o c k - o n r e s p o n s e t o a b a i t e d and u n b a i t e d SL i n t e r a c t e d s i g n i f i c a n t l y w i t h wind speed ( X 2 6 = 32.9; p < 0.05), p r e v e n t i n g s t a t i s t i c a l c o m parisons between t h e two. However, l o c k - o n r e s p o n s e was g e n e r a l l y h i g h e r i n t h e p r e s e n c e o f s e m i o c h e m i c a l s a t a l l wind speeds t h a n when t h e s e m i o c h e m i c a l s were a b s e n t . The d i f f e r e n c e s i n re s p o n s e between t h e s e t r e a t m e n t s were e s p e c i a l l y n o t i c e a b l e i n t h e absence o f wind, where 54% o f t h e f l y i n g b e e t l e s l o c k e d on t o t h e b a i t e d SL, w i t h 29% l a n d i n g on i t , compared t o o n l y 8% l o c k i n g on t o t h e c o n t r o l SL, o f w h i c h l e s s t h a n 5% l a n d e d . P o s i t i v e r e s p o n s e d e c r e a s e d i n t h e p r e s e n c e o f an a i r f l o w , y e t i n c r e a s i n g w ind speed from 0.3 t o 0.9 m/s d i d n o t r e s u l t i n any f u r t h e r d e c r e a s e s . TOTAL NUMBER OF BEETLES OBSERVED W1NDSPEED (M/S) UNBAITED BAITED i i 1 1 1 1 1 1 1 1 1 1 — 0.0 03 0.6 0.9 0.0 0.3 0.6 0.9 0.0 03 0.6 0.9 WIND SPEED (M/S) LOCK-ON WITH LOCK-ON WITHOUT N O RESPONSE LANDING LANDING POSITIVE LOCK - ON RESPONSE F i g . 7. P o s i t i v e l o c k - o n r e s p o n s e e x h i b i t e d by T. l i n e a t u m , a t d i f f e r e n t wind speeds, i n t h e p r e s e n c e o f an u n b a i t e d and s e m i o c h e m i c a l - b a i t e d s i m u l a t e d l o g . Male and female d a t a were p o o l e d . Bars w i t h d i f f e r e n t l e t t e r s f o r each r e s p o n s e by b a i t t r e a t m e n t c a t e g o r y a r e s i g n i f i c a n t l y d i f f e r e n t (X p < 0.05) f o r i n d i v i d u a l comparisons. 32 2.4 D i s c u s s i o n I n s t u d y 1, t h e d e c r e a s e i n t r a p c a t c h w i t h i n c r e a s e d w i nd speed was e x p e c t e d s i n c e T. l i n e a t u m p r e f e r t o f l y under low wind speed c o n d i t i o n s (Rudinsky and Daterman 19 6 4 ) . However, t h e o c c u r r e n c e o f h i g h e s t b e e t l e c a t c h f r e q u e n c y a t 0.0 m/s was u n a n t i c i p a t e d . T h i s s u g g e s t s t h a t t h e b e e t l e s c a p t u r e d w i t h i n an a i r f l o w o r i e n t e d upwind t o t h e t r a p , y e t b e e t l e s c a p t u r e d w i t h o u t an a i r f l o w must have responded by u s i n g a d i f f e r e n t o r i e n t a t i o n mechanism o r f l i g h t p a t t e r n . I n t h e absence o f wind and s e m i o c h e m i c a l s , t h e 3:1 r a t i o o f u p t u n n e l v s . downtunnel t a k e - o f f d i r e c t i o n ( F i g . 4) may be a r e s u l t i n p a r t from t h e l i g h t g r a d i e n t coming t h r o u g h t h e c e i l i n g o f t h e t u n n e l . F r a n c i a and Graham (1967) found t h a t T. l i n e a t u m were p h o t o p o s i t i v e b e f o r e and a f t e r f l i g h t , a l t h o u g h v a r i a t i o n a s s o c i a t e d w i t h t h i s b e h a v i o r was h i g h . I n a d d i t i o n , b e e t l e s may have v i s u a l l y d e t e c t e d t h e a v a i l a b i l i t y o f more f l y i n g space u p t u n n e l t h a n downtunnel from t h e r e l e a s e t r a y . These e x p l a n a t i o n s may a l s o a c c o u n t f o r t h e h i g h e r p r o p o r t i o n o f u p t u n n e l v s . downtunnel f l i e r s i n t h e absence o f wind and s e m i o c h e m i c a l s ( F i g . 5 ) . I n t h i s case however, i n c r e a s e d space u p t u n n e l from t h e r e l e a s e t r a y would not be j u s t p e r c e i v e d t h r o u g h v i s i o n , b u t would be based on t h e b e e t l e ' s a c t u a l f l i g h t a c t i v i t i e s . 33 Both wind and semiochemicals s i g n i f i c a n t l y i n f l u e n c e d b e e t l e b e h a v i o r i n most of the response c a t e g o r i e s s t u d i e d . The h i g h e r frequency of p o s i t i v e response t o a semiochemical-baited SL i n the absence of wind may have r e s u l t e d from i n i t i a l p h o t o p o s i t i v e f l i g h t or random f l i g h t b r i n g i n g the b e e t l e s c l o s e t o the SL by chance, f o l l o w e d by chemo-orientation near the SL. The i n c r e a s e i n steady f l i g h t , l o c k - o n response, and l a n d i n g i n the presence of the semiochemical-baited SL i n d i c a t e s some form of chemo-o r i e n t a t i o n . Wind appears t o o v e r r i d e any p h o t o p o s i t i v e or random f l i g h t b e h a v i o r the b e e t l e s might e x h i b i t , as t a k e -o f f d i r e c t i o n ( F i g . 4) and o v e r a l l f l i g h t d i r e c t i o n ( F i g . 5) s h i f t e d s u b s t a n t i a l l y from uptunnel t o downtunnel upon t h e i r exposure t o a wind speed as low as 0.3 m/s. While upwind chemo-anemotaxis (a d i r e c t e d response t o a chemical mediated by wind) appears t o be the mechanism by which T. lineatum f i n d and o r i e n t t o a pheromone source (Chapman, 1962; Chapters 5 and 6), c l o s e - r a n g e ( i . e . < 1 m) o r i e n t a t i o n and l a n d i n g on the host s u b s t r a t e does not appear t o r e q u i r e wind. The chemo-anemotactic responses, r e p o r t e d i n wind t u n n e l s t u d i e s by Choudhury and Kennedy (1980) f o r S c o l y t u s m u l t i s t r i a t u s (Marsh.), B i r c h and White (1988) f o r Anobium punctatum (Deg.), and M i z e l l e t a l . (1984) f o r Thanasimus dubius (F.) d i d not i n c o r p o r a t e a pheromone source w i t h i n a no-wind environment. With L e p i d o p t e r a , Farkas and Shorey (1972) demonstrated t h a t the p i n k bollworm, Pectinophora g o s s y p i e l l a (Saund.), d i d not 34 r e q u i r e moving a i r t o f l y toward a pheromone s o u r c e , and s u g g e s t e d t h a t t h e moths o r i e n t e d t o t h e c h e m i c a l s t i m u l i by s e n s i n g d i f f e r e n c e s i n c o n c e n t r a t i o n . Baker and Kuenen (1982) had s i m i l a r r e s u l t s w i t h t h e o r i e n t a l f r u i t moth, G r a p h o l i t h a m o l e s t a ( B u s k . ) , y e t t h e y showed t h a t l o n g i t u d i n a l k l i n o t a x i s was t h e mechanism used by t h e s e moths t o o r i e n t t o t h e odor s o u r c e . I n b o t h s t u d i e s , f l i g h t was i n i t i a t e d w i t h i n an a i r f l o w . I n t h e s t u d y r e p o r t e d i n t h i s c h a p t e r , f l i g h t was i n i t i a t e d w i t h o u t an a i r f l o w . The mechanism o f o r i e n t a t i o n used by T. l i n e a t u m p o s s i b l y r e s u l t e d from a c o m b i n a t i o n o f f i r s t p h o t o p o s i t i v e o r random f l i g h t , and t h e n c h e m i c a l l y s t i m u l a t e d o r i e n t a t i o n . I t i s p o s s i b l e t h a t b e e t l e s combined v i s u a l and o l f a c t o r y s t i m u l i t o l a n d on t h e SL (Borden e t a l . 1986; Kennedy 1986). Combined use o f v i s u a l and o l f a c t o r y cues i n S c o l y t i d a e have been r e p o r t e d by Pitman and V i t e (1969) f o r t h e mountain p i n e b e e t l e , Dendroctonus ponderosae Hopk., and by T i l d e n e t a l . (1983) f o r t h e w e s t e r n p i n e b e e t l e , Dendroctonus  b r e v i c o m i s LeConte. F u r t h e r e x p e r i m e n t s a r e needed t o s e p a r a t e o u t t h e e f f e c t s o f each o f t h e s e v a r i a b l e s , b e f o r e a p r e c i s e d e s c r i p t i o n o f T. l i n e a t u m o r i e n t a t i o n t o a h o s t i n t h e absence o f wind can be made. W i t h wind p r e s e n t , b e e t l e s t e n d t o t a k e - o f f and f l y downwind, away from t h e s o u r c e o f s e m i o c h e m i c a l s . D e s p i t e t h e c o n s t a n t exposure o f t h e b e e t l e s t o t h e s e m i o c h e m i c a l plume, o n l y 2 0% o f t h e b e e t l e s appeared t o r e s p o n d chemo-a n e m o t a c t i c a l l y . The h i g h f r e q u e n c y o f non-responders may be a r e s u l t o f t h e a r t i f i c i a l c o n d i t i o n s imposed on t h e b e e t l e s by t h e wind t u n n e l , i n w h i c h t h e s m a l l d i m e n s i o n s ( r e l a t i v e t o a f i e l d s e t t i n g ) may have i n h i b i t e d t h e b e e t l e s from c a r r y i n g o u t t h e f u l l complement o f t h e i r f l i g h t b e h a v i o r , such as f l i g h t e x e r c i s e , b e f o r e r e s p o n d i n g p o s i t i v e l y t o s e m i o c h e m i c a l s . However, f u r t h e r work has shown t h a t a d d i t i o n a l f l i g h t e x e r c i s e p r i o r t o t e s t i n g o f t h e a l r e a d y f l o w n b e e t l e s i n t h e wind t u n n e l d i d n o t enhance t h e f r e q u e n c y o f p o s i t i v e r e s p o n s e t o s e m i o c h e m i c a l s (Appendix 2 ) . A n o t h e r e x p l a n a t i o n f o r low res p o n s e i s t h e v a r i a t i o n i n p h y s i o l o g i c a l s t a t e i n i n d i v i d u a l b e e t l e s , c o n s i d e r e d an i m p o r t a n t f a c t o r i n i n f l u e n c i n g o r i e n t a t i o n o f b a r k b e e t l e s t o o l f a c t o r y s t i m u l i (Borden e t a l . 1986; Wood 1982). These a u t h o r s h y p o t h e s i z e d t h a t some b e e t l e s r e spond i m m e d i a t e l y t o s e m i o c h e m i c a l s and c o l o n i z e s u i t a b l e h o s t t r e e s nearby, w h i l e o t h e r s f l y f o r a p e r i o d o f t i m e b e f o r e r e s p o n d i n g , and t h u s s t a r t new c e n t e r s o f a t t r a c t i o n a t g r e a t e r d i s t a n c e s . A t k i n s (1966a, 1969) l i n k e d t h i s d i s p e r s a l p a t t e r n t o f a t c o n t e n t i n Dendroctonus pseudotsuqae. P h y s i c a l s t a t e o f t h e b e e t l e s such as nematode o r m i c r o o r g a n i s m i n f e c t i o n may a l s o p l a y a r o l e i n t h i s . A l t h o u g h I am n o t s u r e o f t h e mechanism i n T. l i n e a t u m , I b e l i e v e t h e v a r i a b i l i t y i n re s p o n s e b e h a v i o r w i t h i n a p o p u l a t i o n may ac c o u n t f o r t h e h i g h v a r i a t i o n i n response f o r t h i s s t u d y . A p p a r e n t l y , low f r e q u e n c y o f ambrosia b e e t l e r e s p o n s e t o s e m i o c h e m i c a l s i s q u i t e common. I n b i o a s s a y s o f w a l k i n g i n s e c t s w i t h i n o l f a c t o m e t e r s , 40 t o 50% response i s t h e b e s t t h a t has been o b s e r v e d (Borden e t a l . 1977; Borden e t a l . 1980; M a c C o n n e l l e t a l . 1977). The d e c r e a s e s i n T. l i n e a t u m r e s p o n s e t o s e m i o c h e m i c a l s a t i n c r e a s i n g wind speeds c o r r e s p o n d w e l l w i t h o b s e r v a t i o n s i n f i e l d s i t u a t i o n s , where t w i c e as many marked T. l i n e a t u m were r e c a p t u r e d i n a f o r e s t when wind speed was h a l f t h e normal average o f 0.4 m/s (Chapter 6 ) . Other s c o l y t i d b e e t l e s have a l s o been c a p t u r e d i n h i g h e r numbers d u r i n g p e r i o d s o f calm, o f t e n a s s o c i a t e d w i t h oncoming t h u n d e r s t o r m s ( V i t e e t a l . 1964; Edson 1978). I t may be argued t h a t by i n c r e a s i n g t h e w i n d speed i n t h e t u n n e l , a d e c r e a s e i n t h e s i z e o f t h e s e m i o c h e m i c a l plume would r e s u l t , r e d u c i n g t h e chance t h a t b e e t l e s w i l l r e s p o n d w h i l e i n f l i g h t . Sanders (1985) showed t h a t a plume o f smoke r e l e a s e d from a p o i n t s o u r c e d e c r e a s e d i n s i z e w i t h an i n c r e a s e i n wind speed. However, i n our s t u d i e s , t h e plumes were r e l e a s e d from a much l a r g e r a r e a t h a n a p o i n t s o u r c e , and a t e s t w i t h T i C l 4 smoke showed t h a t t h e plume s i z e s d i d n o t d e c r e a s e s i g n i f i c a n t l y between 0.25 and 0.75 m/s. A n o t h e r c o n s i d e r a t i o n i s t h a t a t l o w e r wind speeds t h e s e m i o c h e m i c a l c o n c e n t r a t i o n p e r u n i t volume o f a i r i n t h e t u n n e l would be i n c r e a s e d , p o s s i b l y a c c o u n t i n g f o r t h e i n c r e a s e d r e s p o n s e by t h e b e e t l e s . Y e t r e l e a s e r a t e s f o r b o t h s t u d i e s were 10 t i m e s h i g h e r t h a n t h o s e w h i c h e l i c i t o p t i m a l r e s p o n s e s i n t h e same t u n n e l ( C hapter 3 ) . Thus, a t l o w e r wind speeds when s e m i o c h e m i c a l s a r e above o p t i m a l r e l e a s e r a t e s , p o s i t i v e r e s ponse would most l i k e l y d e c r e a s e r a t h e r t h a n i n c r e a s e . I n summary, t h e p r e s e n c e o f wind as w e l l as i t s speed has a s i g n i f i c a n t i n f l u e n c e on T. l i n e a t u m r e s p o n s e t o a s e m i o c h e m i c a l - b a i t e d s u b s t r a t e . I n t h e s e e x p e r i m e n t s I was a b l e t o s t u d y t h e i n f l u e n c e o f t h e s e f a c t o r s a t a l o n g i t u d i n a l range o f c a . 3 m. D i f f e r e n t b e h a v i o r p a t t e r n s by T. l i n e a t u m were o b s e r v e d i n t h e p r e s e n c e and absence o f wind. When a i r f l o w was a b s e n t , f l i g h t t e n d e d t o be random, a l t h o u g h more b e e t l e s ended u p t u n n e l n e a r t h e SL p o s s i b l y due t o t h e l i g h t g r a d i e n t i n t h e t u n n e l , g i v i n g them an o p p o r t u n i t y t o be a r r e s t e d by s e m i o c h e m i c a l s d i f f u s i n g from t h e s o u r c e . W i t h wind p r e s e n t , t h e m a j o r i t y o f b e e t l e s t o o k o f f and f l e w downwind. B e e t l e s t h a t d i d r e s pond t o t h e s e m i o c h e m i c a l s f l e w upwind and o r i e n t e d t o t h e s o u r c e i n s t e a d o f b e i n g a r r e s t e d by i t . I n d e s c r i b i n g T. l i n e a t u m d i s p e r s a l and subsequent a t t r a c t i o n t o h o s t m a t e r i a l , i t i s i m p o r t a n t t o keep t h e e n v i r o n m e n t a l c o n d i t i o n s as w e l l as t h e p h y s i o l o g i c a l s t a t u s o f t h e b e e t l e s i n c o n t e x t . I n a f i e l d s e t t i n g where wind speeds were l i g h t (< 0.5 m/s), T. l i n e a t u m g e n e r a l l y f l e w i n a l l d i r e c t i o n s (Chapter 5 ) , y e t a t l o n g d i s t a n c e s , such as 500 m, b e e t l e s were caught downwind, whereas a t s h o r t d i s t a n c e s (< 25 m) r e s p o n d i n g b e e t l e s f l e w upwind t o w a r d s e m i o c h e m i c a l - b a i t e d t r a p s . F i e l d d a t a , r e l a t i n g h i g h e s t b e e t l e c a t c h e s t o s t i l l w ind c o n d i t i o n s , as w e l l as t h i s 38 wind t u n n e l s t u d y , suggest t h a t once T. l i n e a t u m a r r i v e w i t h i n 1 o r 2 m o f t h e h o s t , a r r e s t m e n t , o r i e n t a t i o n , and subsequent l a n d i n g on t h e h o s t i s b e s t f a c i l i t a t e d d u r i n g p e r i o d s o f l i g h t o r n i l wind c o n d i t i o n s . I n l e s s o p t i m a l p e r i o d s , when t h e wind speed i n c r e a s e s , T. l i n e a t u m can o r i e n t t o and l a n d on a h o s t u s i n g chemo-anemotaxis, b u t w i l l do so a t l o w e r f r e q u e n c i e s . The d i f f e r e n t mechanisms used by s p r i n g d i s p e r s i n g T. l i n e a t u m as t h e y s e a r c h f o r a t t r a c t i v e h o s t m a t e r i a l a l l o w them t o o p t i m i z e t h e i r s e a r c h by a d a p t i n g t o t h e v a r y i n g e n v i r o n m e n t a l c o n d i t i o n s e n c o u n t e r e d . 39 CHAPTER 3: RESPONSE OP TRYPODENDRON LINEATUM TO VARIED RELEASE RATES OF ETHANOL AND LINEATIN IN A WIND TUNNEL. 3.1 I n t r o d u c t i o n The a m b r o s i a b e e t l e , Trypodendron l i n e a t u m ( O l i v i e r ) , l i k e most s p e c i e s o f t h e f a m i l y S c o l y t i d a e , r e l i e s on i t s o l f a c t o r y p e r c e p t i o n o f c h e m i c a l s f o r h o s t a t t r a c t i o n and ma t i n g (Borden 1985). Because t h i s i n s e c t i s a major p e s t o f l o g g e d t i m b e r on t h e P a c i f i c c o a s t o f B r i t i s h C o lumbia, a s u b s t a n t i a l e f f o r t has been p u t i n t o i d e n t i f y i n g and q u a n t i f y i n g t h e c h e m i c a l s t o wh i c h t h e b e e t l e r esponds ( L i n d g r e n e t a l . 1983). T h i s has r e s u l t e d i n . i m p r o v e d methodology i n s u r v e y i n g and managing t h e p e s t (Borden and McLean 1981; L i n d g r e n and Borden 1983). Three c h e m i c a l s have been i d e n t i f i e d as a t t r a c t a n t s f o r T. l i n e a t u m i n N o r t h A m e r i c a ; e t h a n o l (Moeck 1970, 1971) and c<.-pinene ( N i j h o l t and Schonherr 1976) as h o s t a t t r a c t a n t s , and l i n e a t i n , a t r i - c y c l i c k e t a l (MacConnell e t a l . 1977; Borden e t a l . 1979) as t h e a g g r e g a t i o n pheromone. C o m b i n a t i o n s o f t h e s e c h e m i c a l s , f o r use i n t r a p p i n g programs, have been i n v e s t i g a t e d i n Europe and w e s t e r n N o r t h A m e r i c a where T. l i n e a t u m o c c u r s . Borden e t a l . (1982) c o n f i r m e d r e p o r t s from Europe ( P a i v a and K i e s e l 1985; V i t e and Bakke 1979) t h a t e t h a n o l , oC-pinene, and l i n e a t i n a c t e d s y n e r g i s t i c a l l y i n a t t r a c t i n g T. l i n e a t u m . However, Borden e t a l . (1982) found t h a t e t h a n o l and o<-pinene d i d n o t 40 enhance t h e a t t r a c t i o n o f T. l i n e a t u m t o s t i c k y w i r e mesh o r d r a i n p i p e t r a p s i n B r i t i s h Columbia. I n c o n t r a s t , Shore and McLean (1983) found t h a t e t h a n o l and £K.-pinene t o g e t h e r d i d a c t s y n e r g i s t i c a l l y w i t h l i n e a t i n i n a t t r a c t i n g N o r t h A merican T. l i n e a t u m a d u l t s t o d r a i n p i p e t r a p s . The d i f f e r e n t r e s u l t s may be a t t r i b u t e d t o s e v e r a l f a c t o r s . F i r s t l y , Borden e t a l . (1982) used r e l e a s e r a t e s o f e t h a n o l and oC-pinene 3 - 6 and 2 - 3 t i m e s g r e a t e r , r e s p e c t i v e l y , t h a n d i d Shore and McLean (1983). S e c o n d l y , Shore and McLean (1983) a c t u a l l y t e s t e d f o r i n t e r a c t i o n s between t h e s e m i o c h e m i c a l s o f G n a t h o t r i c h u s s u l c a t u s (LeConte) and T. l i n e a t u m . These i n t e r a c t i o n s may have i n f l u e n c e d b e e t l e r e s p o n s e t o t r a p s b a i t e d o n l y w i t h T. l i n e a t u m s e m i o c h e m i c a l s . I n a d d i t i o n , t h e e x p e r i m e n t a l d e s i g n used by Shore and McLean (1983) was n o t p l a n n e d t o d i f f e r e n t i a t e t h e e f f e c t s o f e t h a n o l and oC-pinene on b e e t l e r e s p o n s e . The c o n f l i c t i n g r e s u l t s o f t h e s e s t u d i e s , a l o n g w i t h t h e development and w i d e s p r e a d use o f t h e m u l t i p l e - f u n n e l t r a p ( L i n d g r e n 1983), prompted t h e i n v e s t i g a t i o n o f t h e imp o r t a n c e o f t h e s e s e m i o c h e m i c a l s i n a t t r a c t i n g T. l i n e a t u m t o t h e e f f i c i e n t new t r a p s . A f i e l d s t u d y was f i r s t c o n d u c t e d i n B r i t i s h Columbia by Salom and McLean (1988) , where i t was shown t h a t l i n e a t i n was t h e o n l y e f f e c t i v e c h e m i c a l i n a t t r a c t i n g T. l i n e a t u m t o f u n n e l t r a p s . No s y n e r g i s t i c e f f e c t s r e s u l t e d from a d d i n g e t h a n o l ( r e l e a s e r a t e = 75 mg/24 h) o r c<_-pinene ( r e l e a s e r a t e = 30 mg/24 h) , 41 a l o n e o r t o g e t h e r , t o t h e t r a p s . T h i s was f o l l o w e d w i t h t h e s t u d y r e p o r t e d h e r e w h i c h a t t e m p t e d t o d e t e r m i n e t h e o p t i m a l r e l e a s e r a t e s o f s e m i o c h e m i c a l s f o r use i n wind t u n n e l b i o a s s a y s (Baker and L i n n , J r . 1984). The s t u d y was d e s i g n e d t o compare t h e number o f T. l i n e a t u m caught i n t r a p s b a i t e d w i t h e t h a n o l and l i n e a t i n , u s i n g v a r i o u s c o m b i n a t i o n s and r e l e a s e r a t e s . 3.2 M a t e r i a l s and Methods The a t t r a c t i o n o f b e e t l e s t o f u n n e l t r a p s b a i t e d w i t h v a r i o u s r e l e a s e r a t e s o f e t h a n o l and l i n e a t i n , was s t u d i e d i n 1987 i n a wind t u n n e l d e s c r i b e d i n C h a p t e r 2. Three 8-f u n n e l t r a p s were p l a c e d i n t h e upwind s e c t i o n o f t h e t u n n e l , 0.5 m from t h e s c r e e n . The t h r e e t r a p s p r o v i d e d a l a r g e r plume o f s e m i o c h e m i c a l s w i t h i n t h e t u n n e l t h a n c o u l d be a c h i e v e d w i t h a s i n g l e t r a p . Treatments f o r t e s t i n g t h e re s p o n s e o f T. l i n e a t u m i n t h e wind t u n n e l i n c l u d e d 4 l i n e a t i n r e l e a s e r a t e s (0, 8, 64, and 512 ug/24 h ) , each t e s t e d i n a l l c o m b i n a t i o n s w i t h 3 r e l e a s e r a t e s o f 95% e t h a n o l (0, 75, and 150 mg/24 h ) . T h i s r e s u l t e d i n 12 t r e a t m e n t c o m b i n a t i o n s o f l i n e a t i n and e t h a n o l . The 0,0 r e l e a s e r a t e was t h e c o n t r o l t r e a t m e n t . S l o w - r e l e a s e l i n e a t i n l u r e s were used, i n t h e form o f Hereon c o n t r o l l e d r e l e a s e d i s p e n s e r s (Kydonieus and B e r o z a 1981), w i t h r e l e a s e r a t e s based on l u r e s i z e (7.0 ug/24 42 h/cm2)!2-. The l u r e s were aged f o r one week a t ambient t e m p e r a t u r e s t o a l l o w t h e r e l e a s e r a t e s t o s t a b i l i z e . S i x l u r e s were used i n each l i n e a t i n t r e a t m e n t . A l u r e was p l a c e d i n t h e t o p and bottom o f each t r a p . The s i z e o f each l u r e used f o r r e l e a s e r a t e s o f 8, 64, and 512 ug/24 h, were 0.19, 1.51, and 12.2 cm 2, r e s p e c t i v e l y . E t h a n o l (95%) was r e l e a s e d from a 40 mL p l a s t i c c o n t a i n e r w i t h a 1 mm-diam-aperture ( r e l e a s e r a t e = 75 mg/24 I i h) 1^. F o r a r e l e a s e r a t e o f 75 mg/24h, one l u r e was p l a c e d i n t h e c e n t r a l t r a p , and f o r t r e a t m e n t s o f 150 mg/24 h, one l u r e was p l a c e d i n each o f t h e o u t e r t r a p s . D i f f e r e n t placement o f e t h a n o l may have i n f l u e n c e d t h e odor plume, y e t t h e t y p e o f d i s p e n s e r used l i m i t e d me t o t h i s d e s i g n . Traps used f o r t h e c o n t r o l t r e a t m e n t s were n o t t h e same as t h o s e used f o r t h e b a i t e d t r e a t m e n t s . To see i f b a i t e d t r a p s c a r r i e d r e s i d u e , a c o m p a r a t i v e t e s t was r u n t o compare b e e t l e r e s p o n s e t o t h e s e t r a p s f o l l o w i n g b a i t r e m o v a l , w i t h t h e r e s p o n s e t o t h e c o n t r o l t r a p s . The b e e t l e s used i n t h i s s t u d y had been c o l l e c t e d d u r i n g t h e s p r i n g o f 1987 w i t h l i n e a t i n - b a i t e d m u l t i p l e -f u n n e l t r a p s a t t h e same f i e l d s i t e used i n C h a p t e r 2. B e e t l e s were c o l l e c t e d d a i l y and p l a c e d i n 1 L p l a s t i c c o n t a i n e r s w i t h m o i s t e n e d c l o t h t o w e l s . The c o n t a i n e r s were s t o r e d i n a w a l k - i n c o o l e r a t 4°C under a 14:10 h (L:D) p h o t o p e r i o d . The c o n t a i n e r s were checked t w i c e w e e k l y t o |2_ Phero Tech. I n c . , 1140 C l a r k Dr., Vancouver, B.C. V5L 3K3. 43 m o n i t o r m o i s t u r e l e v e l s . These b e e t l e s s u r v i v e d f o r more t h a n 100 days. P r i o r t o t e s t i n g , t h e b e e t l e s were removed from t h e p l a s t i c c o n t a i n e r s and s e l e c t e d f o r t e s t i n g based on t h e i r h e a l t h y appearance ( i . e . p r e s e n c e o f a l l body p a r t s ) and t h e i r a b i l i t y t o walk n o r m a l l y . The s e l e c t e d b e e t l e s were t h e n p l a c e d i n p e t r i d i s h e s w i t h c l o t h t o w e l s and s t o r e d a t 4°C u n t i l needed f o r t h e t e s t , always on t h e same day. The b e e t l e s were g i v e n a warm-up p e r i o d o f 15 min a t room t e m p e r a t u r e p r i o r t o r e l e a s e . The b e e t l e s were r e l e a s e d from a h o r i z o n t a l t r a y , 3 5 cm above t h e t u n n e l f l o o r , 1.5 m downwind from t h e t r a p s . The wind speed was h e l d c o n s t a n t a t 0.15 m/s (Chapter 2 ) . A 4 X 3 f a c t o r i a l e x periment was s e t up as a randomized complete b l o c k d e s i g n w i t h days s e r v i n g as b l o c k s . M a l e s and f e m a l e s were t e s t e d s e p a r a t e l y . Because o f t h e l a r g e number o f t r e a t m e n t s , one r e p l i c a t i o n was t e s t e d p e r day f o r each o f 15 e x p e r i m e n t a l days. F o r each r e p l i c a t i o n , 25 b e e t l e s were r e l e a s e d and a l l o w e d 10 min t o r e s p o n d . The p e r c e n t a g e o f b e e t l e s caught i n t h e t r a p s was used as t h e dependent v a r i a b l e . The a r c s i n e t r a n s f o r m e d d a t a were a n a l y z e d by a n a l y s i s o f v a r i a n c e and mean s e p a r a t i o n between t r e a t m e n t s was c a r r i e d out w i t h t h e Student-Newman-Kuel's t e s t (P < 0.05) (SAS 1985). 44 3.3 R e s u l t s The number o f T. l i n e a t u m caught i n f u n n e l t r a p s i n t h e wind t u n n e l was n o t s i g n i f i c a n t l y a f f e c t e d by t h e p r e s e n c e o f e t h a n o l e i t h e r f o r males ( F 2 1 5 4 = 0.4; P > 0.05) o r fem a l e s ( F 2 ^54 = 1.4; P > 0.05). I n c o n t r a s t , a s i g n i f i c a n t i n c r e a s e i n t h e number o f b e e t l e s caught o c c u r r e d i n t h e p r e s e n c e o f l i n e a t i n a l o n e f o r b o t h males ( F i g 8A) and females ( F i g . 8B). Males responded b e s t a t t h e l o w e s t r e l e a s e r a t e s o f l i n e a t i n w i t h an average t r a p c a t c h o f 2 3.5 and 21.6% f o r t h e 8 and 64 ug/24 h r e l e a s e r a t e s , r e s p e c t i v e l y . A s i g n i f i c a n t r e d u c t i o n i n r e s p o n s e o c c u r r e d a t 512 ug/24 h w i t h a mean c a t c h o f 16.6% o f t h e b e e t l e s t e s t e d . D e s p i t e somewhat l o w e r c a t c h e s , s i m i l a r r e s p o n s e s were found f o r f e m a l e s , i n wh i c h t h e h i g h e s t c a t c h e s , 20.0 and 16.1%, were o b t a i n e d a t 8 and 64 ug/24 h, r e s p e c t i v e l y . Mean c a t c h a t 512 ug/24 h was 14.3 %, a s i g n i f i c a n t l y l o w e r r e s p o n s e t h a n f o r t h e 8 ug t r e a t m e n t . No s i g n i f i c a n t d i f f e r e n c e s i n r e s p o n s e by T. l i n e a t u m were o b s e r v e d between t h e c o n t r o l t r a p s and u n b a i t e d t r a p s t h a t had p r e v i o u s l y h e l d e t h a n o l and l i n e a t i n d u r i n g t h e e x p e r i m e n t ( F 5 7 2 = 0.9; P > 0.05). Thus, t h e r e was no e v i d e n c e f o r c o n t a m i n a t i o n o f t h e t r a p s . 45 F i g . 8. Mean p e r c e n t (±) S.E. o f T. l i n e a t u m caught i n t r a p s i n r e s p o n s e t o c o m b i n a t i o n s o f v a r i e d r e l e a s e r a t e s o f l i n e a t i n and e t h a n o l i n a w ind t u n n e l : A) Males and B) Females. L i n e a t i n r e l e a s e r a t e columns, p o o l e d a c r o s s e t h a n o l t r e a t m e n t s , w i t h t h e d i f f e r e n t l e t t e r s a r e s i g n i f i c a n t l y d i f f e r e n t (SNK; p < 0.05). 46 3.4 D i s c u s s i o n Ethanol was hypothesized t o a c t as an a r r e s t a n t / b o r i n g s t i m u l a n t f o r b e e t l e s i n c l o s e p r o x i m i t y t o a s u i t a b l e host (McLean and Borden 1977). The r e f o r e , s i n c e d r a i n p i p e t r a p s r e q u i r e t h a t b e e t l e s l a n d on the t r a p and crawl through s m a l l diam h o l e s i n order t o e n t e r the t r a p (McLean e t a l . 1987; V i t e and Bakke 1979), et h a n o l would appear t o serve an important f u n c t i o n i n c a p t u r i n g b e e t l e s . In c o n t r a s t , f u n n e l t r a p s capture b e e t l e s t h a t f l y i n t o the t r a p s , and don't r e q u i r e them t o l a n d . T h i s type of b e h a v i o r would e x p l a i n our data, which suggests t h a t t h e r e i s no g a i n i n adding e t h a n o l b a i t s t o f u n n e l t r a p s , as the l i n e a t i n alone i s s u f f i c i e n t f o r a t t r a c t i n g and c a p t u r i n g T. lineatum. I t i s p o s s i b l e t h a t ethanol and oC-pinene may be important i n a t t r a c t i n g T. lineatum t o f u n n e l t r a p s , but t h a t the d i s p e n s e r s normally used f o r r e l e a s i n g them may account f o r our r e s u l t s . T h i s has been i n v e s t i g a t e d by Cushon (unpublished datajl 3-. He has shown i n p r e l i m i n a r y s t u d i e s t h a t newly developed closed-system host l u r e s work b e t t e r than the o l d host l u r e s i n c a p t u r i n g T. 1ineatum. However, f u r t h e r t e s t i n g i s needed t o compare b e e t l e capture i n f u n n e l t r a p s b a i t e d w i t h l i n e a t i n p l u s new h o s t l u r e s t o t r a p s b a i t e d w i t h l i n e a t i n alone. The argument a g a i n s t the use of the o l d e t h a n o l l u r e ( p l a s t i c c o n t a i n e r w i t h the [3 Mr. Geoff Cushon, Research T e c h n i c i a n , Phero Tech. Inc., 114 0 C l a r k Dr., Vancouver. B.C. V5L 3K3. s m a l l h o l e i n t h e c o ver) i s t h a t r a i n w a t e r g e t s i n t o t h e c o n t a i n e r d i l u t i n g t h e e t h a n o l s o l u t i o n , r e d u c i n g t h e a t t r a c t i o n o f t h e b a i t . Yet t h i s i s n o t a p p l i c a b l e t o t h i s w i n d t u n n e l s t u d y . I n a d d i t i o n , t h e same d i s p e n s e r s were shown t o be e f f e c t i v e i n e n h ancing t h e a t t r a c t i o n o f T. l i n e a t u m i n Europe as w e l l as a n o t h e r a m b r o s i a b e e t l e s p e c i e s , G n a t h o t r i c h u s s u l c a t u s (LeConte) t o s t i c k y w i r e mesh t r a p s (Borden e t a l . 1982). Response o f T. l i n e a t u m t o v a r i o u s r e l e a s e r a t e s o f l i n e a t i n has been p r e v i o u s l y examined i n a f i e l d s e t t i n g i n B r i t i s h Columbia by L i n d g r e n e t a l . (1983). They found t h a t t h e number o f b e e t l e s caught on c y l i n d r i c a l s t i c k y w i r e mesh t r a p s b a i t e d w i t h l i n e a t i n , i n c r e a s e d as r e l e a s e r a t e s i n c r e a s e d , from 10-40 ug/24h, and remained t h e same between 40 and 800 ug/24h. I n a l a t e r e x p e r i m e n t d e s c r i b e d i n t h e same paper, r e l e a s e r a t e s o f 40 ug/24h were found t o be o p t i m a l f o r f u n n e l t r a p s , y e t a r e l e a s e r a t e o f o n l y 10 ug/24 h o f l i n e a t i n was adequate as l o n g as t h e r e m a i n i n g l i n e a t i n (30 ug/24 h) was p l a c e d w i t h i n 1.5 - 2 m o f t h e t r a p . Our r e s u l t s i n t h e wind t u n n e l w i t h low r e l e a s e r a t e s c o r r e s p o n d w e l l w i t h t h o s e from L i n d g r e n e t a l . (1983). However, a d e c r e a s e i n r e s p o n s e was o b s e r v e d i n t h e w ind t u n n e l a t t h e h i g h e r r e l e a s e r a t e s . The d i f f e r e n c e s i n s e n s i t i v i t y o f t h e b e e t l e s a t t h e h i g h e r r a t e s may have r e s u l t e d from a r t i f i c i a l f a c t o r s imposed by t h e wind t u n n e l , an e n c l o s e d environment w i t h a c o n s t a n t , u n i d i r e c t i o n a l a i r f l o w . Both f a c t o r s r e s u l t e d i n c o n t i n u o u s e x posure o f t h e 48 b e e t l e s t o t h e pheromone, whereas i n t h e f i e l d , pheromone plumes a r e o f t e n b r o k e n up by t u r b u l e n c e and w i n d s h i f t s ( F a r e s e t a l . 1980), r e s u l t i n g i n n o n - c o n t i n u o u s e x p o s u r e . F o r t h e above r e a s o n s , as w e l l as t h e p o t e n t i a l o f a t t r a c t i n g b e e t l e s from a l a r g e r a r e a , and because t h e l u r e s weaken w i t h age, i t i s p r o b a b l y b e s t f o r m a s s - t r a p p i n g i n t h e f i e l d t o keep t h e r e l e a s e r a t e s o f l i n e a t i n h i g h e r t h a n f o r s t u d y i n g b e e t l e f l i g h t b e h a v i o r i n t h e wind t u n n e l . Host v o l a t i l e s may w e l l be i m p o r t a n t i n i n i t i a l h o s t r e c o g n i t i o n by T. l i n e a t u m i n a n a t u r a l f o r e s t s i t u a t i o n . However, from t h e r e s u l t s i n Salom and McLean (1988) and t h i s c h a p t e r , l i n e a t i n b a i t s a l o n e appear t o be s u f f i c i e n t f o r r u n n i n g an e f f e c t i v e m a s s - t r a p p i n g program w i t h m u l t i p l e - f u n n e l t r a p s around l o g booms and d r y l a n d s o r t s , where h o s t v o l a t i l e s a r e l i k e l y t o be p r e s e n t anyway. I f t h e new h o s t l u r e s a r e t o be used i n t h e s e programs, f u r t h e r s t u d i e s s h o u l d be c a r r i e d o ut t o v e r i f y and q u a n t i f y t h e i r enhanced e f f e c t on b e e t l e c a p t u r e . 49 CHAPTER 4: WIND TUNNEL EVALUATION OF FLIGHT AND LANDING BEHAVIOR OF TRYPODENDRON LINEATUM (OLIVIER) IN RESPONSE TO A HOST ATTRACTANT AND AGGREGATION PHEROMONE 4.1 Introduction The e f f e c t of o l f a c t o r y s t i m u l i on f o r e s t C o l e o p t e r a i s b e s t understood f o r the bark and ambrosia b e e t l e s ( S c o l y t i d a e ) , where o r i e n t a t i o n t o host m a t e r i a l i s c o n s i d e r e d t o be a s e t of s e q u e n t i a l events (Borden e t a l . 1986). For the s t r i p e d ambrosia b e e t l e , Trypodendron  li n e a t u m ( O l i v i e r ) , i t i s h y p o t h e s i z e d t h a t the p i o n e e r females cue i n t o the host a t t r a c t a n t s (primary a t t r a c t a n t phase), and upon s u c c e s s f u l l y b o r i n g i n t o a s u i t a b l e l o g , emit a pheromone t h a t r e s u l t s i n the mass a t t a c k of the l o g s by males and females (secondary a t t r a c t i o n phase) (Borden 1985). As mentioned i n Chapter 1, the h o s t a t t r a c t a n t f o r North American T. lineatum has been i d e n t i f i e d as e t h a n o l and the a g g r e g a t i o n pheromone has been i d e n t i f i e d , and i s known t o be a t r i - c y c l i c k e t a l c a l l e d l i n e a t i n . Over the p a s t decade, development of a f a c i l e s y n t h e s i s f o r the pheromone, along w i t h the e v o l u t i o n of t r a p technology (Lindgren 1983; Lindgren e t a l . 1983), has l e d t o semiochemical-based p e s t management s t r a t e g i e s around l o g p r o c e s s i n g areas (Borden 1988b). In an e f f o r t t o i n c r e a s e e f f i c i e n c y of t h i s program, i n v e s t i g a t o r s have looked a t the i n t r a - s p e c i f i c response of T. lineatum t o v a r i o u s 50 c o m b i n a t i o n s o f h o s t a t t r a c t a n t s and i t s pheromone (Borden e t a l . 1982; Salom and McLean 1988). I n t e r - s p e c i f i c r e s p o n s e o f T. l i n e a t u m and t h e c l o s e l y r e l a t e d G n a t h o t r i c h u s spp. t o t h e i r r e s p e c t i v e pheromones and h o s t a t t r a c t a n t s has a l s o been i n v e s t i g a t e d (Borden e t a l . 1981; Shore and McLean 1983). Host a t t r a c t a n t s were c o n c l u d e d t o be n e c e s s a r y components o f t h e s e m i o c h e m i c a l bouquet f o r a t t r a c t i n g N o r t h American T. l i n e a t u m by Shore and McLean (1983) whereas t h e y were not i n two o t h e r s t u d i e s (Borden e t a l . 1982; Salom and McLean 1988). T h i s has r e s u l t e d i n some c o n f u s i o n as t o t h e a c t u a l r o l e s o f t h e s e c h e m i c a l s i n N o r t h American T. l i n e a t u m h o s t f i n d i n g , o r i e n t a t i o n , and a t t a c k . A d d i n g t o t h e c o n f u s i o n , s t u d i e s c o n d u c t e d i n Europe have c o n s i s t e n t l y shown t h a t T. l i n e a t u m r e s p o n d t o h o s t a t t r a c t a n t s a l o n e (Bauer and V i t e 1985) and re s p o n d s y n e r g i s t i c a l l y t o l i n e a t i n when h o s t a t t r a c t a n t s a r e added t o t h e bouquet (Borden e t a l . 1982; P a i v a and K i e s e l 1985; V i t e and Bakke 1985), r e g a r d l e s s o f t h e t r a p s employed f o r c a p t u r i n g b e e t l e s (Borden e t a l . 1982; McLean e t a l . 1987). I n t h i s c h a p t e r , I a d d r e s s e d t h e d i f f e r e n c e s i n r e s u l t s f o r N o r t h American T. l i n e a t u m by c o n d u c t i n g a d e t a i l e d b e h a v i o r a l s t u d y i n a wind t u n n e l t o examine b e e t l e f l i g h t and l a n d i n g r e s p o n s e t o o l f a c t o r y s t i m u l i . By c o n d u c t i n g t h e s t u d y i n t h e wind t u n n e l , I was a b l e t o c o n t r o l most f a c t o r s and make d e t a i l e d b e h a v i o r a l o b s e r v a t i o n s o f i n d i v i d u a l b e e t l e s . 51 4 . 2 Materials and Methods The s t u d y was conducted i n a wind t u n n e l d e s c r i b e d i n C h a p t e r 2, u s i n g a c o n s t a n t wind speed o f 0.15 m/s, a t a t e m p e r a t u r e i n s i d e t h e t u n n e l o f 23 + 2°C. The b e e t l e s used i n t h i s s t u d y were c o l l e c t e d d a i l y d u r i n g May and June o f 1987 and 1988, a t t h e same l o g boom s i t e d e s c r i b e d i n C h a p t e r 2. S t o r a g e p r o c e d u r e s were a l s o s i m i l a r t o t h e d e s c r i p t i o n g i v e n i n C h a p t e r 2. T r a p s Three m o d i f i e d Norwegian d r a i n p i p e t r a p s ( B o r r e g a a r d I n d . L t d . , 1701 S a r p s b o r g , Norway) were s e t out a t e q u a l s p a c i n g s a c r o s s t h e upwind s e c t o r o f t h e w ind t u n n e l , c a . 0.6 m from t h e s c r e e n . T h i s t r a p d e s i g n was chosen o v e r o t h e r d e s i g n s because i t was j u d g e d t o b e s t s i m u l a t e a l o g . E x t e r n a l r i d g e s p r o j e c t i n g 3 mm from t h e t r a p s u r f a c e , c i r c l e t h e f u l l l e n g t h o f t h e t r a p c o n c e n t r i c a l l y , a l l o w i n g b e e t l e s t o l a n d and s u b s e q u e n t l y walk on t h e t r a p , as t h e y would on a l o g . Numerous h o l e s (3 mm d i a m ) , punched between t h e r i d g e s , s i m u l a t e g a l l e r y h o l e s i n a l o g . They p r o v i d e : 1) t h e o penings needed f o r s e m i o c h e m i c a l e m i s s i o n , and 2) e n t r a n c e s i n t o t h e t r a p f o r r e s p o n d i n g b e e t l e s . As a r e s u l t , b e e t l e s t h a t l a n d on t h e s e t r a p s a r e r e q u i r e d t o go t h r o u g h a s e a r c h i n g phase t o f i n d t h e s e m i o c h e m i c a l s o u r c e , s i m i l a r t o a f i e l d s i t u a t i o n where males s e a r c h f o r mates 52 and f e m a l e s s e a r c h f o r s u i t a b l e b o r i n g s i t e s (McLean e t a l . 1987; V i t e and Bakke 1979). S e v e r a l m o d i f i c a t i o n s were made on t h e s e t r a p s from t h o s e used i n t h e f i e l d ( F i g . 9 ) . Traps were p a i n t e d a l i g h t g r e e n - g o l d c o l o r t o a l l o w f o r e a s i e r o b s e r v a t i o n o f b e e t l e movement near and on t h e t r a p . L i n d g r e n e t a l . (1983) found no d i f f e r e n c e s i n r e s p o n s e by T. l i n e a t u m t o f u n n e l t r a p s o f 5 d i f f e r e n t c o l o r s , s u g g e s t i n g t h a t p a i n t i n g our t r a p s would n o t a l t e r b e e t l e r e s p o n s e . The bottom s e c t i o n was f i t t e d w i t h a f u n n e l t h a t t i g h t l y c o n n e c t e d t h e t r a p and t h e c o l l e c t i n g j a r . The s e m i o c h e m i c a l plume e m i t t e d from t h e t r a p was s i m u l a t e d u s i n g T i C l 4 smoke. I n i t i a l t e s t s i n d i c a t e d t h a t a f u n n e l f i t t e d i n t o t h e c o l l e c t i n g j a r and t h e t r a p c o v e r l i f t e d 1 cm above t h e t o p o f t h e t r a p , a c h i e v e d e m i s s i o n o f t h e plume o v e r t h e f u l l l e n g t h o f t h e t r a p ( F i g . 9A). The s e m i o c h e m i c a l d i s p e n s e r s were p l a c e d i n t h e t r a p by a 1 cm -diam wooden dowel suspended from t h e t r a p c o v e r ( F i g . 9B). The l e n g t h o f each t r a p was 1.1 m, p r o d u c i n g a c o m p o s i t e plume from t h e t h r e e t r a p s w h i c h c o v e r e d most o f t h e volume w i t h i n t h e t u n n e l . Treatments Traps u n b a i t e d ( c o n t r o l ) , and b a i t e d w i t h e t h a n o l and l i n e a t i n , a l o n e and i n c o m b i n a t i o n , s e r v e d as t h e f o u r t r e a t m e n t s o f s t u d y . I n t r e a t m e n t s c o n t a i n i n g e t h a n o l , 95 % e t h a n o l was r e l e a s e d from two 40 ml d i s p e n s e r s ( t o t a l B F i g . 9. A) M o d i f i e d Norwegian d r a i n p i p e t r a p s used as a b a i t i n g s u b s t r a t e f o r T. l i n e a t u m . w i t h a t y p i c a l smoke plume p a t t e r n produced a t 0.15 m/s. B) C l o s e - u p o f t r a p c o v e r and l u r e h o l d e r , showing an upper and l o w e r l i n e a t i n b a i t , and a m e d i a l e t h a n o l b a i t . 54 r e l e a s e r a t e = 150 mg/24 each p l a c e d i n t h e c e n t e r o f t h e two o u t e r t r a p s . I n t r e a t m e n t s c o n t a i n i n g l i n e a t i n , t h e pheromone was d i s p e n s e d from Hereon® c o n t r o l l e d r e l e a s e d e v i c e s (Kydonieus and Bero z a 1981), where r e l e a s e r a t e s were based on l u r e s i z e (7 ug/24h/cm 2)l 2-. Two 0.76 cm 2 l i n e a t i n l u r e s were p l a c e d i n t h e t o p and bottom o f a l l t h r e e t r a p s , r e s u l t i n g i n a t o t a l r e l e a s e r a t e o f 32 ug/24 h (Chapter 3 ) . A l l l u r e s had been aged f o r one week a t ambient t e m p e r a t u r e s t o a l l o w r e l e a s e r a t e s t o s t a b i l i z e . Removal o f a l l t r a c e s o f s e m i o c h e m i c a l s from t h e wind t u n n e l as w e l l as t h e t r a p s f o r t h e c o n t r o l t r e a t m e n t s would have been v e r y d i f f i c u l t t o c a r r y o u t d u r i n g t h e r u n o f t h e ex p e r i m e n t i n 1987. T h i s l e d t o a r e p e a t o f t h e c o n t r o l t r e a t m e n t s o n l y , d u r i n g 1988, t o t e s t f o r e v i d e n c e o f c o n t a m i n a t i o n d u r i n g t h e f i r s t s e t o f u n b a i t e d c o n t r o l t r e a t m e n t s . P r i o r t o t h e second s e t o f c o n t r o l t r e a t m e n t s , t h e c h a r c o a l f i l t e r i n t h e wind t u n n e l was r e a c t i v a t e d , t h e o l d d u s t f i l t e r was r e p l a c e d w i t h a new one, t h e w i n d t u n n e l was c o m p l e t e l y c l e a n e d w i t h a 0.5% h y p o c h l o r i t e s o l u t i o n , and p r e v i o u s l y unused m o d i f i e d d r a i n p i p e t r a p s were used. A l l subsequent p r o c e d u r e s d e s c r i b e d below were t h e same f o r b o t h t h e f i r s t ( a l l t r e a t m e n t s ) and second ( c o n t r o l s o n l y ) e x p e r i m e n t a l s e t s . T e s t P r o c e d u r e B e e t l e s were sexed and g i v e n a w a l k i n g t e s t i n t h e morning p r i o r t o f l i g h t i n t h e wind t u n n e l . They were 55 a l l o w e d t o a c c l i m a t e t o t h e warmer room t e m p e r a t u r e s (22 -25°C) f o r 15 min b e f o r e r e l e a s e , a p e r i o d s u f f i c i e n t t o ens u r e maximal r e s p o n s e o f t h e b e e t l e s t o t h e t r a p s . B e e t l e s were r e l e a s e d from t h e same p l a t f o r m a t t h e same l o c a t i o n as was used i n t h e s t u d i e s r e p o r t e d i n C h a p t e r 2. Two b e e t l e s , w i t h males and females t e s t e d s e p a r a t e l y , were r e l e a s e d f o r each t r e a t m e n t / r e p l i c a t i o n . A l l t r e a t m e n t s were s e l e c t e d randomly w i t h i n each r e p l i c a t i o n . I n 1987, a range o f 1 t o 4 r e p l i c a t i o n s were conducted/day o v e r 24 e x p e r i m e n t a l days, whereas i n 1988, 8 t o 12 r e p l i c a t i o n s o f t h e second s e t o f u n b a i t e d t r e a t m e n t s were c o n d u c t e d o v e r 6 e x p e r i m e n t a l days. I n t o t a l , 124 b e e t l e s o f each sex were t e s t e d i n each t r e a t m e n t . The d e t a i l e d o b s e r v a t i o n s o f b e e t l e r e s p o n s e t o t h e d i f f e r e n t t r e a t m e n t s i n c l u d e d ; o c c u r r e n c e o f f l i g h t , t i m e t o f l i g h t t a k e - o f f , t a k e - o f f d i r e c t i o n , f l i g h t t y p e , o v e r a l l f l i g h t d i r e c t i o n , l o c k - o n b e h a v i o r , l a n d i n g on t h e t r a p s , t i m e t o l a n d i n g , and b e h a v i o r on t h e t r a p s . F l i g h t t a k e - o f f d i r e c t i o n was s c o r e d as upwind, v e r t i c a l , o r downwind. Type o f f l i g h t was s c o r e d as s t e a d y o r e r r a t i c (see C h a p t e r 2 f o r d e s c r i p t i o n s ) . O v e r a l l f l i g h t d i r e c t i o n , upwind, downwind, o r n o n - d i r e c t i o n a l , was based on t h e predominant d i r e c t i o n f l o w n by t h e i n s e c t t h r o u g h o u t t h e o b s e r v a t i o n p e r i o d . I f b e e t l e s d i d n o t e x h i b i t predominant up- o r downtunnel f l i g h t , t h e y were c o n s i d e r e d n o n - d i r e c t i o n a l f l i e r s . Lock-on b e h a v i o r was a l s o d e s c r i b e d i n Ch a p t e r 2. F o r b e e t l e s l a n d i n g on t h e t r a p s , o b s e r v a t i o n s were made t o see i f t h e y e n t e r e d t h e t r a p s , and i f t h e y d i d , how l o n g i t t o o k . A l l o b s e r v a t i o n s were r e c o r d e d on a c a s s e t t e t a p e and l a t e r t r a n s c r i b e d onto d a t a s h e e t s . Response b e h a v i o r d a t a were summarized as f r e q u e n c i e s and a n a l y z e d as d e s c r i b e d i n C h a p t e r 2. Response t i m e d a t a were a n a l y z e d w i t h a G e n e r a l L i n e a r Model (GLM) (SAS 1985), w i t h subsequent mean s e p a r a t i o n between t r e a t m e n t s c a r r i e d o u t w i t h t h e Student-Newman-Kuel's t e s t (p < 0.05). 4.3 R e s u l t s No s i g n i f i c a n t d i f f e r e n c e s were found between t h e f i r s t and second y e a r u n b a i t e d c o n t r o l t r e a t m e n t s f o r a l l b e h a v i o r s and t i m e r e s p o n s e s o b s e r v e d , e x c e p t f o r f l i g h t t a k e - o f f d i r e c t i o n ( T a ble 2 ) . The d i f f e r e n t r e s u l t s i n f l i g h t t a k e - o f f d i r e c t i o n d i d not a p p a r e n t l y a f f e c t any o t h e r s e t o f r e s u l t s . T h e r e f o r e , t h e u n b a i t e d t r e a t m e n t r e s u l t s from y e a r 1 were c o n s i d e r e d a v a l i d c o n t r o l f o r t h e b a i t e d t r e a t m e n t s f o r a l l b e h a v i o r c a t e g o r i e s s t u d i e d e x c e p t f l i g h t t a k e - o f f d i r e c t i o n . I n t h e l a t t e r c a s e , b o t h y e a r 1 and y e a r 2 u n b a i t e d t r e a t m e n t r e s u l t s a r e p r e s e n t e d . O c c u r r e n c e o f f l i g h t was found t o be i n d e p e n d e n t o f s e m i o c h e m i c a l t r e a t m e n t and sex ( T a b l e 3 ) . A l t h o u g h n o t s i g n i f i c a n t , s l i g h t l y more b e e t l e s f l e w i n t h e p r e s e n c e o f b a i t e d t r a p s v s . u n b a i t e d t r a p s . Females d i d e x h i b i t a h i g h e r tendency t o f l y t h a n males ( X ^ = 5.72; p < 0.05), a l t h o u g h t h e d i f f e r e n c e c o u l d not be d e t e c t e d a t i n d i v i d u a l 57 T a b l e 2. S t a t i s t i c a l c omparison o f Trypodendron l i n e a t u m r e s p o n s e s t o f i r s t and second y e a r c o n t r o l t r e a t m e n t s i n a wind t u n n e l . The numbers o f b e e t l e s r e l e a s e d f o r each s e x / t r e a t m e n t = 124. B e h a v i o r T e s t S t a t i s t i c 1 P r o b a b i l i t y Observed (Degrees o f Freedom) L e v e l O c c u r r e n c e o f F l i g h t x 2 ( l ) = 0.69 0.4054 Time o f F l i g h t F ( l , 2 8 3 ) = ° - 0 1 0.9094 F l i g h t T a k e - o f f x 2 ( 2 ) = 8 - 3 3 0.0155 * 2 D i r e c t i o n F l i g h t Type x 2 ( l ) = 0 , 0 0 0.9451 O v e r a l l F l i g h t x 2 ( 2 ) = 2 , 1 0 0.3495 D i r e c t i o n Lock-on Response x 2 ( 2 ) = ° - 9 8 0.6111 Time Between T a k e - o f f 3  and L a n d i n g C a p t u r e d i n T r a p s as x 2 ( l ) = 0 , 0 ° 0.9491 % o f B e e t l e s L o c k i n g - o n 1 Degrees o f freedom f o r t h e X 2 s t a t i s t i c a r e a p r o d u c t o f t r e a t m e n t ( t - 1 ) by re s p o n s e l e v e l s ( r - 1 ) . 2 S i g n i f i c a n t p r o b a b i l i t y l e v e l i s < 0.05. 3 • Not enough r e s p o n d e r s t o make t i m e co m p a r i s o n s . 58 T a b l e 3. F l i g h t f r e q u e n c y o f Trypodendron l i n e a t u m t o d i f f e r e n t s e m i o c h e m i c a l b a i t s i n a wind t u n n e l . Number o f b e e t l e s r e l e a s e d f o r each sex i n each b a i t t r e a t m e n t = 124. A t t r a c t a n t % F l y i n g M ales Females U n b a i t e d 54.0 ( 6 7 ) 2 64.5 (80) E t h a n o l 63 .7 (79) 68.5 (85) L i n e a t i n 66.9 (83) 72.6 (90) L i n e a t i n + E t h a n o l 63.7 (79) 71.8 (89) S i g n i f i c a n t d i f f e r e n c e s i n re s p o n s e were found between sex (X ; p < 0.05), b u t n o t among b a i t s ( X 2 ; p > 0.05). Number o f f l o w n b e e t l e s a r e p r e s e n t e d i n t h e p a r e n t h e s e s . 59 t r e a t m e n t l e v e l s . Response t i m e from r e l e a s e t o i n i t i a l t a k e - o f f was found t o be independent o f t r e a t m e n t and sex ( F i g . 1 0 ) . Mean t a k e - o f f t i m e f o r a l l t r e a t m e n t s averaged 80 seconds. D i f f e r e n c e s i n upwind and v e r t i c a l t a k e - o f f d i r e c t i o n s were o b s e r v e d between t h e f i r s t and second y e a r u n b a i t e d c o n t r o l s ( F i g . 1 1 ) . Upwind t a k e - o f f was h i g h e r i n y e a r 1 and v e r t i c a l t a k e - o f f was h i g h e r i n y e a r 2 t r e a t m e n t s . I n y e a r 1, no d i f f e r e n c e s i n upwind t a k e - o f f d i r e c t i o n were o b s e r v e d f o r any t r e a t m e n t s . An i n c r e a s e i n downwind t a k e -o f f was o b s e r v e d i n t h e p r e s e n c e o f l i n e a t i n and e t h a n o l t o g e t h e r . These d i f f e r e n c e s i n r e s p o n s e s between t h e 1987 and 1988 c o n t r o l t r e a t m e n t s f o r t h i s b e h a v i o r d i d n o t appear t o a f f e c t o t h e r a s p e c t s o f T. l i n e a t u m b e h a v i o r i n t h e wind t u n n e l . F r e q u e n c i e s o f s t e a d y and e r r a t i c f l i g h t were i n f l u e n c e d by t h e p r e s e n c e o f s e m i o c h e m i c a l s , w i t h no d i f f e r e n c e s o b s e r v e d between males and fe m a l e s ( F i g . 1 2 ) . E t h a n o l caused a s i g n i f i c a n t i n c r e a s e i n s t e a d y f l i g h t from 27 t o 37% and a d e c r e a s e i n e r r a t i c f l i g h t from 72 t o 63%, when compared t o t h e u n b a i t e d t r e a t m e n t . W i t h l i n e a t i n a l o n e , t h e f r e q u e n c y o f s t e a d y f l i g h t i n c r e a s e d s i g n i f i c a n t l y t o 48%. E t h a n o l combined w i t h l i n e a t i n d i d no t f u r t h e r enhance t h e o c c u r r e n c e o f s t e a d y f l i g h t . No d i f f e r e n c e s between males and females were o b s e r v e d i n o v e r a l l f l i g h t d i r e c t i o n ( F i g . 1 3 ) . Upwind f l i g h t d i r e c t i o n i n c r e a s e d s i g n i f i c a n t l y from 15 t o 31% i n t h e 120 110 100 •—s CO 90 | 80 H 70 CO ON 60 CO 50 2 40 1 30 20 10 0 rZ71 M A L E 1X5 F E M A L E UNBAITED ETHANOL LINEATIN SEMIOCHEMICAL BAITS LIN + ETH Fig. 10. Mean time i n t e r v a l + S.E. (indicated by a v e r t i c a l l i n e running through the top of each bar) between release and f l i g h t take-off of T . lineatum. i n the presence of semiochemical baits i n drainpipe traps, within a wind tunnel. o 100 (ZZ1 UNBAITED (1988) F x X l UNBAITED (1987) UPWIND VERTICAL DOWNWIND FLIGHT TAKE-OFF DIRECTION Fig. 11. Flight take-off d i r e c t i o n of T. lineatum. i n the presence of semiochemical baits i n drainpipe traps, within a wind tunnel. Male and p female data were pooled. Stars on top of unbaited (1988) bars denotes a significant difference (X ; p < 0.05) with the unbaited — 2 (1987) treatment. Bars for each d i r e c t i o n , with different l e t t e r s , indicates significant differences in response (X ; p < 0.05), for individual comparisons. The numbers of observations for each treatment were: unbaited (1988) - 137; unbaited (1987) - 138; ethanol - 146; l i n e a t i n - 150; and l i n e a t i n and ethanol - 152. CO O 0-co CQ 100 90 80 70 60 50 40 30 20 10 0 [771 UNBAITED fV\l ETHANOL W7A LINEATIN LIN + ETH STEADY ERRATIC FLIGHT TYPE Fig. 12. Observations of f l i g h t type exhibited by T. lineatum. in the presence of semiochemicals in drainpipe traps, within a wind tunnel. Male and female data were pooled. Proportions are based on the tot a l number of observations, where each individual could be scored for more then one type of f l i g h t . Bars for each type of f l i g h t , with different l e t t e r s , indicates significant differences i n in response (X ; p < 0.05), for individual comparisons. The numbers of observations for each treatment were: unbaited - 140; ethanol - 152; l i n e a t i n - 158; and l i n e a t i n and ethanol - 156. [771 UNBAITED UPWIND DOWNWIND NO DIRECTION OVERALL FLIGHT DIRECTION Overall f l i g h t d i r e c t i o n exhibited by T. lineatum, i n the presence of semiochemical baits i n drainpipe traps, within a wind tunnel. Male and female data were pooled. Bars for each direction, with different l e t t e r s , indicates significant differences in response (X ; p < 0.05), for individual comparisons. The numbers of observations for each treatment were : unbaited - 141; ethanol - 158; l i n e a t i n - 166; and l i n e a t i n and ethanol - 166. 64 p r e s e n c e o f e t h a n o l when compared t o t h e u n b a i t e d t r a p s , w h i l e a c o r r e s p o n d i n g r e d u c t i o n i n downwind f l i g h t from 67 t o 50% o c c u r r e d . F u r t h e r i n c r e a s e s i n upwind f l i g h t o c c u r r e d i n t h e p r e s e n c e o f l i n e a t i n , y e t t h e y were n o t s t a t i s t i c a l l y s i g n i f i c a n t f o r e t h a n o l a l o n e . However, non-d i r e c t i o n a l f l i g h t d e c r e a s e d s i g n i f i c a n t l y . Responses t o e t h a n o l and l i n e a t i n combined d i d not d i f f e r from r e s p o n s e s t o l i n e a t i n a l o n e . P o s i t i v e l o c k - o n r e s p o n s e p a t t e r n s d i f f e r e d between sexes when exposed t o t h e d i f f e r e n t s e m i o c h e m i c a l t r e a t m e n t s ( F i g . 1 4 ) . The l a n d i n g o f males on d r a i n p i p e t r a p s was n o t i n f l u e n c e d s i g n i f i c a n t l y by t h e p r e s e n c e o f e t h a n o l , b u t w i t h l i n e a t i n p r e s e n t , l a n d i n g f r e q u e n c y i n c r e a s e d from 6 t o 27%. The number o f b e e t l e s l a n d i n g d i d n o t i n c r e a s e s i g n i f i c a n t l y when l i n e a t i n and e t h a n o l were combined, b u t t h e f r e q u e n c y d i d r i s e t o 34%. Lock-on r e s p o n s e w i t h o u t l a n d i n g d i d n o t appear t o be a f f e c t e d by t h e p r e s e n c e o f s e m i o c h e m i c a l s . L a n d i n g by females was s i g n i f i c a n t l y g r e a t e r i n t h e p r e s e n c e o f e t h a n o l , w i t h an i n c r e a s e i n r e s p o n s e from 1 t o 12% when compared t o female r e s p o n s e t o u n b a i t e d t r a p s ( F i g . 1 4 ) . The p r e s e n c e o f l i n e a t i n a l s o s i g n i f i c a n t l y enhanced l a n d i n g , w i t h 16% o f t h e f l o w n f e males r e s p o n d i n g . The c o m b i n a t i o n o f e t h a n o l w i t h l i n e a t i n d i d n o t r e s u l t i n an i n c r e a s e d r e s p o n s e o v e r l i n e a t i n o r e t h a n o l a l o n e . Non-l a n d i n g p o s i t i v e r e s p o n s e s i n c r e a s e d s l i g h t l y from 18 t o 24% w i t h e t h a n o l p r e s e n t , and s i g n i f i c a n t l y i n c r e a s e d o v e r t h e z o a to to* (/} Z o CM 1 0 0 5 0 -0 a a - i — i — i — i — U E L L+E LOCK-ON WITH LANDING a b a b D — i — i — i — i — U E L L+E LOCK-ON WITHOUT LANDING MALE FEMALE NO RESPONSE POSITIVE L O C K - O N RESPONSE F i g . 14. P o s i t i v e l o c k - o n r e s p o n s e , e x h i b i t e d as l a n d i n g o r n o n - l a n d i n g b e h a v i o r , by T. l i n e a t u m i n t h e p r e s e n c e o f semi o c h e m i c a l b a i t s i n d r a i n p i p e t r a p s , w i t h i n a wind t u n n e l . Bars f o r each b e h a v i o r and sex, w i t h d i f f e r e n t l e t t e r s , i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e s i n response ( X 2 ; p < 0.05), f o r i n d i v i d u a l c o m p a r i s o n s . The numbers o f males observed f o r each t r e a t m e n t were: u n b a i t e d (U) - 67; e t h a n o l (E) - 79; l i n e a t i n (L) - 83; and l i n e a t i n and e t h a n o l (L+E) - 79. The numbers o f females observed were; U - 80; E - 85; L - 90; and L+E - 89. 66 u n b a i t e d c o n t r o l t o 32% when o n l y l i n e a t i n was p r e s e n t . The c o m b i n a t i o n o f e t h a n o l and l i n e a t i n d i d n o t improve t h i s r e s p o n s e . The t i m e i n t e r v a l between t a k e - o f f and l a n d i n g on t h e t r a p a l s o d i f f e r e d w i t h s e m i o c h e m i c a l t r e a t m e n t s , y e t was independent o f sex ( F i g . 15). The s m a l l number o f e t h a n o l o n l y r e s p o n d e r s would make i t d i f f i c u l t t o d e t e c t t h e p r e s e n c e o f s t a t i s t i c a l d i f f e r e n c e s between s e x e s , t h e r e f o r e male and female r e s p o n s e t i m e s a r e p r e s e n t e d as s e p a r a t e b a r s . Time r e s p o n s e t o e t h a n o l - b a i t e d t r a p s was s i g n i f i c a n t l y s h o r t e r t h a n l i n e a t i n - b a i t e d t r a p s , a v e r a g i n g 81 and 191 s, r e s p e c t i v e l y . I n t e r m e d i a t e r e s p o n s e s were o b s e r v e d when l i n e a t i n and e t h a n o l were combined, w i t h an average r e s p o n s e t i m e o f 128 s, b u t n o t s i g n i f i c a n t l y d i f f e r e n t from e t h a n o l . The p r e s e n c e o f l i n e a t i n was c r i t i c a l i n i n d u c i n g males t o e n t e r t h e t r a p s . I n terms o f t o t a l b e e t l e s r e l e a s e d , 12% were c a p t u r e d ; o f t h e b e e t l e s t h a t f l e w , 18% were c a p t u r e d ; and o f t h e b e e t l e s t h a t showed a p o s i t i v e l o c k - o n r e s p o n s e , 43% were c a p t u r e d (Table 4 ) . Responses t o e t h a n o l were n o t s i g n i f i c a n t l y h i g h e r t h a n r e s p o n s e s t o u n b a i t e d t r a p s , and r e s p o n s e s t o l i n e a t i n and e t h a n o l t o g e t h e r were o n l y s l i g h t l y h i g h e r t h a n l i n e a t i n a l o n e . The p r e s e n c e o f e t h a n o l and l i n e a t i n , however, were b o t h found t o be c r i t i c a l i n s t i m u l a t i n g f e m a l e s t o e n t e r t h e t r a p s ( T a b l e 4 ) . F o r t r a p s b a i t e d w i t h e t h a n o l o n l y , 7% o f t h e t o t a l number o f b e e t l e s r e l e a s e d were c a p t u r e d , 9% o f CO 280 260 240 220 H 200 180 H 160 140 120 100 80 60 40 20 A 0 1771 M A L E [\3 F E M A L E ETHANOL LINEATIN LIN + ETH SEMIOCHEMICAL BAITS Fig. 15. Mean time interval + S.E. (indicated by v e r t i c a l lines running through the top of each bar) between f l i g h t take-off and trap landing of T. lineatum. in the presence of semiochemical baits i n drainpipe traps, within a wind tunnel. Means were compared on combined data for the sexes, where different letters on top of the bars indicate significant differences between semiochemical baits (SNK; p < 0.05). Table 4. The capture of Trypodendron lineatum i n semiochemical-baited drainpipe traps in a wind tunnel (N = 124 released/treatment /sex). No. Beetles % of % of % of % of Beetles 1 Attractant Entering Beetles Beetles Beetles Landing On a Trap Released That Flew Locking On a trap Unbaited 1 Ethanol 3 Lineatin 15 Lin + Eth 19 Unbaited 0 Ethanol 8 Lineatin 7 Lin + Eth 13 Males 0.8 b 2 1.5 b 2.4 b 3.8 b 12.1 a 18.1 a 15.3 a 24.1 a Females 0.0 b 0.0 b 6.5 a 9.4 a 5.6 a 7.8 a 10.5 a 14.6 a 9.1 b 100.0 13.0 b 60.0 42.9 a 71.4 47.5 a 70.4 0.0 c 0.0 27.6 ab 80.0 16.3 b 50.0 37.1 a 81.3 Individual comparisons between treatments could not be carried out due to low numbers. For each column, within each sex, percentages followed by different 2 le t t e r s are s i g n i f i c a n t l y different (X ; p < 0.05), for individual comparisons. t h e number t h a t f l e w were c a p t u r e d , and 28% o f t h e number t h a t showed a p o s i t i v e l o c k - o n r e s p o n s e were c a p t u r e d . Responses t o t r a p s b a i t e d s o l e l y w i t h l i n e a t i n were v e r y s i m i l a r t o t h o s e b a i t e d w i t h e t h a n o l a l o n e . The p r o p o r t i o n o f b e e t l e s e n t e r i n g t h e t r a p s , w i t h r e s p e c t t o a l l c a t e g o r i e s i n T a b l e 4, was h i g h e r when e t h a n o l and l i n e a t i n were combined compared t o e t h a n o l and l i n e a t i n a l o n e , y e t t h i s i n c r e a s e was not s t a t i s t i c a l l y s i g n i f i c a n t . A l t h o u g h t h e t i m e i n t e r v a l between l a n d i n g on and t h e n e n t e r i n g t h e t r a p s was r e c o r d e d , t h e number o f o b s e r v a t i o n s were t o o few t o be a n a l y z e d . No p a t t e r n c o u l d be d i s c e r n e d from t h e few b e e t l e s t h a t were r e c o r d e d . 4.4 D i s c u s s i o n N e i t h e r f l i g h t f r e q u e n c y ( T a b l e 3) n o r t i m e o f f l i g h t t a k e - o f f ( F i g . 10) were a f f e c t e d by t h e d i f f e r e n t t r e a t m e n t s t e s t e d . The i n f l u e n c e o f s e m i o c h e m i c a l s on f l i g h t t a k e - o f f d i r e c t i o n c o u l d n o t be d e t e r m i n e d from our d a t a , due t o t h e v a r y i n g r e s u l t s o b t a i n e d between t h e u n b a i t e d c o n t r o l s i n y e a r 1 and y e a r 2. The d i f f e r e n c e s between t h e two c o n t r o l t r e a t m e n t s ( F i g . 11) a r e d i f f i c u l t t o e x p l a i n because o f t h e l a c k o f d i f f e r e n c e s i n response o b s e r v e d between t h e s e t r e a t m e n t s f o r a l l o t h e r b e h a v i o r c a t e g o r i e s t e s t e d . F l i g h t t y p e was r e c o r d e d t o see i f a c h a r a c t e r i s t i c b e h a v i o r c o u l d be o b s e r v e d t h a t was n e c e s s a r y f o r b e e t l e s t o res p o n d t o s e m i o c h e m i c a l s . I n c r e a s e d s t e a d y and d e c r e a s e d 70 e r r a t i c f l i g h t s were o b s e r v e d when b e e t l e s were exposed t o s e m i o c h e m i c a l s ( F i g . 1 2 ) , s u g g e s t i n g t h a t f l i g h t t y p e may have p o t e n t i a l i n d e s c r i b i n g T. l i n e a t u m r e s p o n s e i n wind t u n n e l s t u d i e s . Steady f l i g h t may i n d i c a t e t h a t b e e t l e s a r e i n a r e c e p t i v e mode t o respond t o a t t r a c t a n t s . A l t h o u g h some b e e t l e s responded p o s i t i v e l y f o l l o w i n g a p e r i o d o f e r r a t i c f l i g h t , most p o s i t i v e r e s p o n s e s came from b e e t l e s t h a t i n i t i a l l y f l e w i n a s t e a d y manner. From f i e l d o b s e r v a t i o n s , I s u s p e c t t h a t b e e t l e s e x h i b i t i n g s t e a d y f l i g h t a r e u n a f f e c t e d by t h e wind t u n n e l , and t h o s e t h a t e x h i b i t e r r a t i c f l i g h t may be r e s p o n d i n g t o t h e u n n a t u r a l c o n d i t i o n s imposed upon t h e i n s e c t s by a c o n s t r a i n e d e nvironment. A l s o t h e s e i n d i v i d u a l s might be t h e ones t h a t a r e l o s t t o n a t u r e . F l i g h t t y p e was n o t r e p o r t e d i n t h e o n l y d e t a i l e d a c c o u n t o f a s t u d y o f a s c o l y t i d b e e t l e i n a wind t u n n e l (Choudhury and Kennedy 1980). B i o a s s a y s w i t h i n w i nd t u n n e l s t h a t account f o r a v e r t i c a l w i nd f l o w ( P h e l a n and M i l l e r 1982) may be needed t o improve b e e t l e r e s p o n s e . Males and females responded s i m i l a r l y i n terms o f o v e r a l l f l i g h t d i r e c t i o n ( F i g . 1 3 ) . E t h a n o l and l i n e a t i n were b o t h i m p o r t a n t i n i n f l u e n c i n g upwind f l i g h t o f b e e t l e s t o t h e b a i t e d t r a p s , w i t h l i n e a t i n c a u s i n g a d e c r e a s e i n n o n - d i r e c t i o n a l f l i g h t . W h i l e upwind f l i g h t r e s p o n s e t o e t h a n o l was e x p e c t e d f o r f e m a l e s , as t h e y a r e t h e p i o n e e r i n g sex f o r t h i s s p e c i e s , i t was a l s o i m p o r t a n t w i t h r e s p e c t t o upwind male r e s p o n s e . Male r e s p o n s e t o e t h a n o l d i d n o t c a r r y o v e r t o any p o s i t i v e l o c k - o n b e h a v i o r , a l t h o u g h a few more males d i d l a n d e d on e t h a n o l - b a i t e d t r a p s t h a n on u n b a i t e d t r a p s ( T a b l e 4 ) . L i n e a t i n was by f a r t h e most p o w e r f u l compound t o i n d u c e males t o l a n d on t h e t r a p s ( F i g . 1 4 ) . By c o n t r a s t , f e m a l e s r e q u i r e d e t h a n o l a l m o s t as much as l i n e a t i n t o i n d u c e t r a p l a n d i n g . B e e t l e s o f b o t h sexes responded i n d e c r e a s i n g t i m e t o t r a p s b a i t e d w i t h l i n e a t i n a l o n e , e t h a n o l and l i n e a t i n t o g e t h e r , and e t h a n o l a l o n e ( F i g . 1 5 ) . E t h a n o l p l a y s a r o l e n o t o n l y i n a t t r a c t i n g f e m a l e s , b u t a l s o t o a l e s s e r e x t e n t i n a t t r a c t i n g males t o h o s t m a t e r i a l . E t h a n o l and l i n e a t i n were o f e q u a l i m p o r t a n c e i n i n d u c i n g females t o e n t e r t r a p s , w h i l e o n l y l i n e a t i n was e f f e c t i v e f o r males ( T a b l e 4 ) . L i n e a t i n was t h e c r i t i c a l compound f o r i n f l u e n c i n g males t o f l y upwind, l a n d on, and e n t e r t h e t r a p s . F o r f e m a l e s , l i n e a t i n was more i m p o r t a n t i n s t i m u l a t i n g s t e a d y upwind f l i g h t t h a n e t h a n o l , however, b o t h compounds were e q u a l l y i m p o r t a n t i n i n d u c i n g l a n d i n g on t h e t r a p . Trap c a p t u r e i n c r e a s e d s l i g h t l y , y e t n o t s i g n i f i c a n t l y , when b o t h compounds were used t o g e t h e r . These r e s u l t s s u g g e s t t h a t t h e h o s t s t i m u l u s may s e r v e p a r t i a l l y as a c l o s e - r a n g e a t t r a c t a n t and t h e pheromone may s e r v e as a l o n g - r a n g e a t t r a c t a n t . A s i m i l a r h y p o t h e s i s has been made f o r t h e h o s t a t t r a c t a n t and pheromone o f t h e s p r u c e b a r k b e e t l e , I p s  tvp o g r a p h u s L., based on e l e c t r o p h y s i o l o g i c a l s t u d i e s ( D i c k e n s 1981). I t was found t h a t I . ty p o g r a p h u s has a l o w e r t h r e s h o l d and w i d e r range o f p e r c e p t i o n o f pheromone c o n c e n t r a t i o n , compared t o t h e c o n c e n t r a t i o n o f t h e h o s t a t t r a c t a n t c < - p i n e n e , i n d i c a t i n g t h e i n s e c t ' s a b i l i t y t o p e r c e i v e t h e pheromone a t g r e a t e r d i s t a n c e s t h a n t h e h o s t v o l a t i l e s . The r e s u l t s o f male and female T. l i n e a t u m r e s p o n s e s t o s e m i o c h e m i c a l s i n t h e wind t u n n e l d i f f e r from f i e l d s t u d i e s p e r f ormed by Borden e t a l . (1982). These a u t h o r s found t h a t b o t h male and female c a p t u r e i n d r a i n p i p e t r a p s was i n h i b i t e d by t h e p r e s e n c e o f e t h a n o l . However, t h e r e l e a s e r a t e s o f e t h a n o l i n t h e i r e x p e r i m e n t was a l m o s t 7x h i g h e r t h a n t h o s e r e p o r t e d h e r e , n o t i n c l u d i n g a d d i t i o n a l e t h a n o l v o l a t i l e s e m i t t e d from nearby l o g g i n g s l a s h . I n f a c t , t h e y r e p o r t e d t h a t h i g h r e l e a s e r a t e s o f e t h a n o l , a t g/24 h, i n h i b i t e d male r e s p o n s e when compared t o l o w e r r e l e a s e r a t e s , a t 120 mg/24 h. I n c o n t r a s t , Shore and McLean (1983) r e p o r t e d i n c r e a s e s i n male and femal e c a t c h e s t o d r a i n p i p e t r a p s b a i t e d w i t h e t h a n o l , p l u s <*=-pinene ( a n o t h e r h o s t v o l a t i l e ) , and l i n e a t i n , when compared t o t r a p s b a i t e d s o l e l y w i t h l i n e a t i n . However, Shore and McLean (1983) d i d not t e s t e t h a n o l and oC-pinene s e p a r a t e l y . Salom and McLean (1988) d e t e r m i n e d i n b o t h f i e l d and l a b o r a t o r y e x p e r i m e n t s t h a t n e i t h e r e t h a n o l n o r cxCrpinene were i m p o r t a n t i n c a p t u r i n g T. l i n e a t u m i n f u n n e l t r a p s . However, b e e t l e s must e n t e r h o l e s f o r c a p t u r e i n d r a i n p i p e t r a p s , and l a c k o f t h i s b e h a v i o r r e q u i r e m e n t i n f u n n e l t r a p may a c c o u n t f o r t h e d i f f e r e n c e i n t h e r e s u l t s between t h e two s t u d i e s . T h i s s t u d y r e p o r t s t h e f i r s t d e t a i l e d o b s e r v a t i o n s o f f l i g h t b e h a v i o r f o r t h i s i n s e c t . U s i n g o p t i m a l r e l e a s e r a t e s (Chapter 3 ) , t h e r e s u l t s c o n f i r m t h e h y p o t h e s i s t h a t p r i m a r y females r e l y p a r t i a l l y on t h e h o s t a t t r a c t a n t e t h a n o l , w h i l e males, a l t h o u g h n o t i n h i b i t e d by e t h a n o l , appear t o r e l y m o s t l y on l i n e a t i n i n s e a r c h i n g f o r f e m a l e mates. E t h a n o l appears t o speed r e s p o n s e by b o t h males and females ( F i g . 1 5 ) , r e s u l t i n g i n l e s s exposure o f t h e s e b e e t l e s t o p r e d a t o r s and o t h e r m o r t a l i t y a g e n t s , d u r i n g t h e i r s p r i n g d i s p e r s a l f l i g h t p e r i o d . I f males were as a t t r a c t e d t o e t h a n o l as females a r e , t h e y would p o s s i b l y f i n d a l t e r n a t i v e h o s t m a t e r i a l i n s t e a d o f homing i n on t u n n e l l i n g f e males who a r e r e a d y t o mate and produce t h e brood needed t o p e r p e t u a t e t h e p o p u l a t i o n . W h i l e males cannot a f f o r d t o s e a r c h f o r p o t e n t i a l s u i t a b l e h o s t s , and t a k e t h e chance o f not f i n d i n g a mate, fe m a l e s can a f f o r d t o respond t o a l t e r n a t i v e b u t e q u a l l y s u i t a b l e h o s t m a t e r i a l , s i n c e t h e y produce t h e a g g r e g a t i o n pheromone. 74 CHAPTER 5: INFLUENCE OF WIND ON THE SPRING PLIGHT OF TRYPODENDRON LINEATUM IN A SECOND-GROWTH CONIFEROUS FOREST 5.1 introduction I n v e s t i g a t i o n s i n t o i n s e c t d i s p e r s a l and f l i g h t b e h a v i o r have become i m p o r t a n t i n f o r e s t p e s t management r e s e a r c h d u r i n g t h e p a s t 2 5 y e a r s , due t o t h e r e c o g n i t i o n o f d i s p e r s a l as an o b l i g a t o r y phase i n t h e l i f e c y c l e f o r many p e s t s ( W e l l i n g t o n 1983), and as an i m p o r t a n t p r o c e s s i n t h e i r p o p u l a t i o n dynamics and sp r e a d ( S t i n n e r e t a l . 1983). F o r t h e s t r i p e d a m brosia b e e t l e , Trypodendron l i n e a t u m ( O l i v i e r ) , many advances have been made i n o u r knowledge o f i t s b i o l o g y o v e r t h i s t i m e ( N i j h o l t 1979). N e v e r t h e l e s s , a r e c e n t s t u d y on t h e o c c u r r e n c e o f ambrosia b e e t l e p o p u l a t i o n s i n commercial f o r e s t s e t t i n g s has emphasized t h e need t o l e a r n more about t h e d i s p e r s a l dynamics o f T. l i n e a t u m (McLean and Salom u n p u b l i s h e d d a t a ) . Q u a n t i t a t i v e i n f o r m a t i o n i s needed c o n c e r n i n g f l i g h t d i s t a n c e and d i r e c t i o n o f T. 1 ineatura, as i t r e l a t e s t o weather p a t t e r n s , t o p ography, and o t h e r e n v i r o n m e n t a l f a c t o r s . Such i n f o r m a t i o n can p l a y an i m p o r t a n t r o l e i n d e v e l o p i n g a p e s t management program f o r T. l i n e a t u m i n t h e f o r e s t , t h r o u g h m o d i f i c a t i o n o f h a r v e s t p l a n n i n g and l o g i n v e n t o r y management, based on where b e e t l e p o p u l a t i o n s may be ex p e c t e d t o d i s p e r s e . 75 The i m p o r t a n c e o f wind i n i n f l u e n c i n g i n s e c t d i s p e r s a l has been w e l l summarized by Johnson (1969) and P e d g l e y (1982). F o r T. l i n e a t u m , upwind f l i g h t towards a t t r a c t i v e m a t e r i a l was ob s e r v e d by Chapman and K i n g h o r n (1958) and documented from t r a p c a t c h e s by Chapman (1962) i n r e l a t i o n t o an average p r e v a i l i n g wind. To improve o ur u n d e r s t a n d i n g o f t h e p a t t e r n o f T. l i n e a t u m s p r i n g d i s p e r s a l , more i n f o r m a t i o n i s needed t o r e l a t e t h e dynamics o f wind d i r e c t i o n t o t h a t o f b e e t l e f l i g h t t o ward s o u r c e s o f s e m i o c h e m i c a l a t t r a c t i o n . A m a r k - r e c a p t u r e s t u d y was u n d e r t a k e n t o e v a l u a t e t h e i n f l u e n c e o f wind on T. l i n e a t u m d i s p e r s a l t o o l f a c t o r y s t i m u l i . 5.2 M a t e r i a l s and Methods The m a r k - r e c a p t u r e s t u d y , c o n s i s t i n g o f t h r e e e x p e r i m e n t s , was conducted from June t o August, 1986, i n a dense, second-growth, c l o s e d canopy f o r e s t o f D o u g l a s - f i r , Pseudotsucra m e n z i e s i i (Mirb.) F r a n c o and w e s t e r n hemlock, Tsuga h e t e r o p h y l l a (Raf.) S a r g . , n a t u r a l l y r e g e n e r a t e d c a . 75 y e a r s ago f o l l o w i n g a c l e a r c u t and b u r n o f t h e a r e a (Thompson 1985). The e x p e r i m e n t a l s i t e was l o c a t e d i n a 140 ha s t a n d on t h e U n i v e r s i t y Endowment Lands (UEL) a t t h e U n i v e r s i t y o f B r i t i s h Columbia, Vancouver, B.C. I t c o u l d o n l y be a c c e s s e d by s m a l l w a l k i n g t r a i l s . T h i s l o c a t i o n was p r e f e r r e d o v e r an i n d u s t r i a l f o r e s t s i t e where t h e p r e s e n c e o f competing h o s t m a t e r i a l ( l o g s ) might have a c o n f o u n d i n g i n f l u e n c e on t h e r e c a p t u r e o f r e l e a s e d b e e t l e s . The f i r s t e x p e r i m e n t i n t h e s t u d y was d e s i g n e d t o examine b e e t l e d i s p e r s a l o v e r a 100 m r a d i u s (3.1 h a ) , where o l f a c t o r y s t i m u l i were p r e s e n t i n a l l d i r e c t i o n s a t t h r e e d i f f e r e n t d i s t a n c e s from t h e r e l e a s e p o i n t . T w e n t y - e i g h t m u l t i p l e - f u n n e l t r a p s , b a i t e d w i t h t h e a g g r e g a t i o n pheromone l i n e a t i n , were s e t up i n t h r e e c o n c e n t r i c r i n g s a t d i s t a n c e s o f 5, 25, and 100 m from t h e r e l e a s e p o i n t , w i t h 4, 8, and 16 t r a p s p l a c e d a t t h e t h r e e d i s t a n c e s , r e s p e c t i v e l y ( F i g . 16). T h i s e x p e r i m e n t was r e p l i c a t e d s i x t i m e s . The f i r s t h y p o t h e s i s t e s t e d was t h a t b e e t l e s f l y upwind t o o l f a c t o r y s t i m u l i . The second h y p o t h e s i s t e s t e d was t h a t most b e e t l e s would f l y upwind t o t h e c l o s e s t s o u r c e o f o l f a c t o r y s t i m u l i . Thus t h e f r e q u e n c y o f c a t c h e s was compared a t t h e t h r e e d i s t a n c e s . The second e x p e r i m e n t examined b e e t l e d i s p e r s a l a t o n l y a 100 m d i s t a n c e from t h e r e l e a s e p o i n t . T h i s f o l l o w e d t h e removal o f t h e 5 and 25 m t r a p s . T h i s e x p e r i m e n t was r e p l i c a t e d f o u r t i m e s . The hypotheses t e s t e d were t h a t b e e t l e f l i g h t would s t i l l be upwind even though c l o s e - r a n g e a t t r a c t a n t s ( i e . 5 and 25 m t r a p s ) were not p r e s e n t , and s e c o n d l y , t h a t a h i g h e r p r o p o r t i o n o f r e l e a s e d b e e t l e s would f l y t h e l o n g e r d i s t a n c e t h a n when c l o s e - r a n g e a t t r a c t a n t s were i n p l a c e , as i n experiment 1. The t h i r d e x p e r i m e n t e v a l u a t e d b e e t l e d i s p e r s a l a t a 500 m r a d i u s from t h e r e l e a s e p o i n t , e q u a l i n g 78.5 ha i n 77 F i g . 16. Trap placement i n mark-recapture studies of T. lineatum on the University Endowment lands i n Vancouver, B r i t i s h Columbia. In the f i r s t experiment, 4, 8, and 16 l i n e a t i n - b a i t e d funnel traps were placed at distances of 5, 25, and 100 m, respectively, from the release point. Experiment 2 u t i l i z e d the 100 m t r a p l i n e only. In experiment 3, traps were set up 500 m from the release point only. The f i r s t r e p l i c a t i o n had 4 traps placed i n each cardinal d i r e c t i o n , and the second and t h i r d r e p l i c a t i o n s has 2 traps at 45° i n t e r v a l s as shown here. 78 a r e a . Two d i f f e r e n t p l o t d e s i g n s were used. The f i r s t d e s i g n had f o u r t r a p s i n each o f t h e f o u r c a r d i n a l d i r e c t i o n s , 500 in from t h e r e l e a s e p o i n t . The second d e s i g n , w i t h two r e p l i c a t i o n s , had p a i r s o f t r a p s p l a c e d a t 45° i n t e r v a l s around t h e c i r c l e , a l s o a t a 500 m d i s t a n c e . F o r p u r p o s e s o f p r e s e n t a t i o n , t h e two d e s i g n s were combined as one e x p e r i m e n t . The h y p o t h e s i s t e s t e d f o r t h i s d i s t a n c e was t h a t d i s p e r s a l would be n o n - d i r e c t i o n a l o r downwind, b u t no t upwind, because b e e t l e s would n o t p e r c e i v e t h e pheromones 500 m from t h e r e l e a s e p o i n t . Trypodendron l i n e a t u m were c o l l e c t e d d u r i n g t h e i r mass f l i g h t p e r i o d o f May and June, 1986, u s i n g l i n e a t i n - b a i t e d m u l t i p l e f u n n e l t r a p s , p l a c e d a d j a c e n t t o a l a r g e l o g boom s t o r a g e a r e a a t t h e mouth o f t h e n o r t h arm o f t h e F r a s e r R i v e r i n Vancouver, B.C. B e e t l e s were c o l l e c t e d d a i l y and p l a c e d i n 1 L p l a s t i c c o n t a i n e r s w i t h s l i g h t l y m o i s t e n e d c l o t h t o w e l s a t 4°C, t o m i n i m i z e t h e i r a c t i v i t y and p r e s e r v e them f o r e x p e r i m e n t a l use. T h i s s t o r a g e t e c h n i q u e was found t o be s a t i s f a c t o r y , as m o r t a l i t y was g e n e r a l l y low f o r t h e f i r s t 50 days. B e e t l e s were marked w i t h f l u o r e s c e n t powder ( F e r r o I n d u s t r i a l P r o d u c t s L t d . , S u r r e y , B.C.) u s i n g t h e vacuum d u s t e r t e c h n i q u e d e s c r i b e d by L i n t o n e t a l . (1987). Marked b e e t l e s were p l a c e d i n a c o o l e r f o r t h e p e r i o d f o l l o w i n g a p p l i c a t i o n o f t h e powder, and remained t h e r e u n t i l t h e i r r e l e a s e i n t h e f o r e s t , 2 - 3 h l a t e r . A b r i e f e x p e r i m e n t was r u n p r i o r t o t h i s s t u d y , t o l o o k a t t h e e f f e c t m a r k i n g 79 T. l i n e a t u m w i t h f l u o r e s c e n t powder may have on m o r t a l i t y and s h o r t d i s t a n c e f l i g h t r e s p o n s e t o s e m i o c h e m i c a l s i n a wind t u n n e l (Appendix 3 ) . No e f f e c t o f m a r k i n g was o b s e r v e d f o r up t o one week f o l l o w i n g i t s a p p l i c a t i o n . Hence, t h e m a r k i n g t e c h n i q u e was c o n s i d e r e d s a t i s f a c t o r y f o r use i n t h i s s t u d y . The r e l e a s e - r e c a p t u r e p r o c e d u r e was t h e same f o r a l l t h r e e e x p e r i m e n t s . On f l i g h t days, b e e t l e s were r e l e a s e d from a shady s i t e under t h e canopy u s u a l l y a t 12 00 h PDT o r when a i r t e m p e r a t u r e r o s e above 15°C, t h e low t e m p e r a t u r e f l i g h t t h r e s h o l d f o r T. l i n e a t u m (Chapman and K i n g h o r n 1958). B e e t l e s , r a n g i n g i n number from 1750 t o 3650, were p p l a c e d on a 1 m wooden t r a y , w i t h a s m a l l frame r i d g e , 0.2 m from t h e edge, e f f e c t i v e l y g i v i n g t h e b e e t l e s two r i d g e s from w h i c h t o t a k e - o f f . The t r a y was s e t 0.5 m above t h e ground. An a b s o r b e n t paper, l i n e d on one s i d e w i t h c e l l o p h a n e , was p l a c e d u n d e r neath t h e t r a y t o c a t c h any b e e t l e s t h a t f e l l o f f . S i m i l a r paper was used as a b e d d i n g f o r t h e i n n e r s q uare o f t h e t r a y , where t h e b e e t l e s were p l a c e d . An e x p e r i m e n t was t e r m i n a t e d 4 - 6 h f o l l o w i n g b e e t l e r e l e a s e , when t e m p e r a t u r e s d e c l i n e d t oward 15°C. The b e e t l e s were t h e n c o l l e c t e d from a l l t h e t r a p s i n t h e p l o t . I n an e f f o r t t o see how many marked b e e t l e s f l e w f o r more t h a n 1 day w i t h i n t h e p l o t , t r a p s were checked l a t e i n t h e a f t e r n o o n o f t h e second day p o s t - r e l e a s e p r o v i d e d t e m p e r a t u r e s had exceeded t h e f l i g h t t h r e s h o l d o f 15°C. 80 The b e e t l e s from t h e t r a p s were b r o u g h t back t o t h e l a b o r a t o r y f o r c o u n t i n g . An u l t r a - v i o l e t lamp was used t o a i d i n i d e n t i f y i n g t h e marked specimens. Unmarked b e e t l e s would sometimes have a l i t t l e powder on them from c o n t a c t w i t h t h e marked b e e t l e s , y e t t h e o r i g i n a l l y marked ones had t h e powder somewhat impregnated on t o t h e i r e x o s k e l e t o n , e n a b l i n g easy d i s t i n c t i o n between marked and unmarked b e e t l e s . V a r i o u s weather measurements were made w h i l e t h e e x p e r i m e n t s were i n p r o g r e s s . Wind d i r e c t i o n and wind speed d a t a were c o l l e c t e d a t 10 min i n t e r v a l s , w i t h t h e use o f a bee-smoker p l a c e d o v e r a c i r c u l a r g r i d 1 m i n r a d i u s and b r o k e n up i n t o 16 i n t e r v a l s o f 22.5° (360° c i r c l e ) . The spout o f t h e smoker was o r i e n t a t e d v e r t i c a l l y a t 0.5 m above t h e f o r e s t f l o o r . Wind speed was measured by t i m i n g t h e movement o f a p u f f o f smoke f o r a d i s t a n c e o f 1 m. Wind p a r a m e t e r s were measured 10 m e a s t o f t h e r e l e a s e t r a y . Temperature measurements were a l s o made e v e r y 10 m i n u t e s w i t h a mercury thermometer s e t i n t h e shade. R e l a t i v e h u m i d i t y was measured h o u r l y w i t h a s l i n g p s y c h r o m e t e r . B a r o m e t r i c p r e s s u r e d a t a were o b t a i n e d from t h e U n i v e r s i t y o f B r i t i s h Columbia C l i m a t o l o g i c a l S t a t i o n l o c a t e d c a . 1 km from t h e s t u d y s i t e . B e e t l e c a t c h e s and wind d i r e c t i o n d a t a a r e p r e s e n t e d i n t h e form o f c i r c u l a r h i s t o g r a m s . The s t a t i s t i c a l p r o c e d u r e d e s c r i b e d by B a t s c h e l e t (1981), i s summarized as f o l l o w s . Mean a n g l e s (0^) o f b e e t l e c a t c h from t h e r e l e a s e p o i n t and 81 mean wind d i r e c t i o n (# w) were d e t e r m i n e d u s i n g a r e c t a n g u l a r c o o r d i n a t e system w i t h x and y axes and o r i g i n 0. The mean v e c t o r l e n g t h (r) was c a l c u l a t e d f o r each mean a n g l e o f b e e t l e f l i g h t and wind d i r e c t i o n . I n unimodal d i s t r i b u t i o n s , r s e r v e s as a measure o f c o n c e n t r a t i o n o f t h e sample p o i n t s around t h e mean a n g l e , and t h u s i n d i r e c t l y as a measure o f d i s p e r s i o n . The v a l u e s o f r range from 0 t o 1. As r i n c r e a s e s , t h e d i s p e r s i o n o f t h e d a t a around t h e mean a n g l e d e c r e a s e s . S i n c e t h e s e d a t a were grouped i n t o a n g u l a r i n t e r v a l s , i t was n e c e s s a r y t o a p p l y a c o r r e c t i o n f a c t o r t o r , because w i t h o u t i t , r would be b i a s e d and have a tendency t o be t o o s m a l l . Use o f r a l o n e as a c o m p a r a t i v e s t a t i s t i c i s c a u t i o n e d because t h e s i z e o f a p o p u l a t i o n t e n d s t o a f f e c t r i n v e r s e l y . The d a t a f o r each c i r c u l a r d i s t r i b u t i o n were s u b j e c t e d t o t h e R a y l e i g h t e s t f o r randomness. The p r o b a b i l i t y o f o b t a i n i n g a v e c t o r l e n g t h g r e a t e r t h a n t h e measured r i s g i v e n by -nr p = e (M.A. Stephens p e r s o n a l c o m m u n i c a t i o n ) 4 . where n e q u a l s t h e number o f o b s e r v a t i o n s . I n c i r c u l a r d i s t r i b u t i o n s where t h i s p r o b a b i l i t y (p) was < 0.05, t h e d i s t r i b u t i o n s were c o n s i d e r e d d i r e c t e d and t h e mean a n g l e (Jd) c o n s i d e r e d s i g n i f i c a n t . To d e t e r m i n e i f b e e t l e f l i g h t d i r e c t i o n d i f f e r e d from wind d i r e c t i o n , a one-sample t e s t , a nalogous t o a one-sample 4 Dr. M.A. Stephens, P r o f e s s o r o f Mathematics and S t a t i s t i c s , Simon F r a s e r U n i v e r s i t y , Burnaby, B.C. 82 t - t e s t f o r d a t a on a l i n e a r s c a l e (Zar 1984), was c o n d u c t e d u s i n g wind d i r e c t i o n as a p r e - a s s i g n e d a n g l e t o be t e s t e d a g a i n s t t h e b e e t l e f l i g h t d i r e c t i o n a n g l e s . A + d - v a l u e was d e t e r m i n e d f o r each t e s t a n g l e , s e r v i n g as 95% c o n f i d e n c e i n t e r v a l s ( B a t s c h e l e t 1981). I f t h e p r e - a s s i g n e d a n g l e f e l l w i t h i n t h e c o n f i d e n c e l i m i t s o f t h e f l i g h t a n g l e , t h e n t h e a n g l e s were n o t c o n s i d e r e d t o be s i g n i f i c a n t l y d i f f e r e n t . To d e t e r m i n e d i f f e r e n c e s i n t h e number o f b e e t l e s caught a t t h e t h r e e d i s t a n c e s i n ex p e r i m e n t 1, a two-way a n a l y s i s o f v a r i a n c e (ANOVA) was r u n (SAS 1985). A t e s t between b e e t l e c a t c h e s a t 100 m between e x p e r i m e n t s 1 and 2 was r u n u s i n g a one-way ANOVA (SAS 1985). 5.3 R e s u l t s and D i s c u s s i o n Weather Of t h e weather measurements made d u r i n g t h e s t u d y , a s i d e from t e m p e r a t u r e , o n l y wind d i r e c t i o n seemed t o have a major i n f l u e n c e on b e e t l e f l i g h t . Wind speed was l i g h t a v e r a g i n g 1.4 - 1.9 km/h d a y - 1 . Even on v e r y windy days o u t s i d e t h e f o r e s t , t h e wind speed a t 0.5 m above t h e f o r e s t f l o o r remained r e l a t i v e l y l i g h t . T h i s phenomenon i s p r e s e n t e d i n d e t a i l by Pedg l e y (1982). W e i g h t i n g i n d i v i d u a l w i nd d i r e c t i o n s by wind speed had a m i n i m a l e f f e c t on t h e c o m p u t a t i o n o f t h e average wind d i r e c t i o n (jdw) , as t h e ranges o f wind speed o b s e r v e d w i t h i n r e p l i c a t e s were v e r y 83 s m a l l . The range o f s t a n d a r d d e v i a t i o n s f o r w ind speed o v e r t h e 13 r e l e a s e s was 0.4 - 0.6 km/h. I n a d d i t i o n , c o mparisons between wind d i r e c t i o n o v e r t h e f i r s t h our f o l l o w i n g b e e t l e r e l e a s e and t h e o v e r a l l f l i g h t p e r i o d were no t p e r c e p t i v e l y d i f f e r e n t . B a r o m e t r i c p r e s s u r e g e n e r a l l y d e c r e a s e d d u r i n g each day t h e s t u d y was r u n . R e l a t i v e h u m i d i t y c o n s i s t e n t l y ranged between 70 - 90% on f l i g h t d ays. F i n a l l y , s k y c o n d i t i o n s were a l m o s t always sunny, w h i c h was n e c e s s a r y f o r t h e t e m p e r a t u r e i n t h e f o r e s t t o r i s e above 15°C. Temperature n e v e r exceeded 21°C, w e l l below t h e h i g h t e m p e r a t u r e t h r e s h o l d o f 3 0°C r e p o r t e d by R u d i n s k y and Daterman (1964). E x p e r i m e n t 1 The p e r c e n t a g e r e c a p t u r e , based on t h e number o f b e e t l e s t h a t f l e w s u c c e s s f u l l y from t h e t r a y , a veraged 24.9% ( T a b l e 5 ) . The averages were f a i r l y c o n s i s t e n t f o r t h e s i x r e p l i c a t i o n s . L e s s t h a n 10% o f t h e r e c a p t u r e d b e e t l e s were c o l l e c t e d d u r i n g t h e 24 h t h a t f o l l o w e d t h e f i r s t b e e t l e c o l l e c t i o n . The o v e r a l l h i g h r a t e o f r e c a p t u r e may be p a r t i a l l y a t t r i b u t e d t o t h e absence o f competing h o s t m a t e r i a l i n t h e s t u d y a r e a . T h i s can be c o n t r a s t e d w i t h Shore and McLean (1988), where s t i c k y vane t r a p s caught o n l y 6% o f t h e marked b e e t l e s , i n a d r y l a n d s o r t c o n t a i n i n g a c o n s i d e r a b l e s u p p l y o f s u s c e p t i b l e h o s t m a t e r i a l . D i r e c t i o n a l d a t a a r e p r e s e n t e d i n d i v i d u a l l y as c i r c u l a r h i s t o g r a m s ( F i g . 1 7 ) . W i t h i n each c i r c l e i s a d o t t e d l i n e Table 5. Numbers of marked T.lineatum released and recaptured in the f l i g h t d i r e c tion study, on the University Endowment Lands, Vancouver, B.C., 1986. No. Marked Beetles Recaptured No. Beetles No. Beetles % of No. Exp. Rep. Released That Flew N That Flew 1 1 1 750 1 470 417 28.4 2 2 400 1 919 411 20.4 3 2 000 1 543 383 24.8 4 2 000 1 877 643 34.2 5 2 000 1 880 379 20.2 6 2 000 1 846 394 21.3 Totals 12 150 10 535 2 627 24.9 2 1 2 000 1 796 207 11.5 2 1 950 1 584 199 12.5 3 2 000 1 691 223 13.2 4 2 000 1 709 214 12.5 Totals 7 950 6 780 843 12.4 3 1 2 500 1 985 35 1.8 2 3 650 2 732 51 1.8 3 3 550 2 497 24 1.0 Totals 9 700 7 214 110 1.5 N CIRCULAR TRAPLINES WIND DIRECTION REP 5m 25m 100m F i g . 17 D i r e c t i o n a l d a t a based on f r e q u e n c y d i s t r i b u t i o n s o f T. l i n e a t u m f l i g h t and wind o b s e r v a t i o n s f o r exp e r i m e n t 1 i n t h e U n i v e r s i t y Endowment Lands s t u d y . 86 r e p r e s e n t i n g t h e mean a n g l e . F o r b e e t l e s , i t i s t h e i r mean d i r e c t i o n o f f l i g h t . A l t h o u g h i t i s n o t known whether b e e t l e s f l e w d i r e c t l y t o t h e t r a p s o r n o t , f o r p u r p o s e s o f t h i s p r e s e n t a t i o n , b e e t l e f l i g h t d i r e c t i o n r e p r e s e n t s t h e r e s u l t a n t d i r e c t i o n o f b e e t l e c a t c h . F o r wind, i t i s i t s mean d i r e c t i o n . The numbers i n t h e c i r c l e r e p r e s e n t t h e r -v a l u e . S i g n i f i c a n t r - v a l u e s a r e n o t e d w i t h a s t a r . The d o t t e d l i n e s and a s s o c i a t e d v a l u e s o u t s i d e t h e c i r c l e match f r e q u e n c y w i t h b a r l e n g t h . I n r e p l i c a t e s 1 - 3 ( F i g . 1 7 ) , wind d i r e c t i o n was w e s t e r l y t o s o u t h w e s t e r l y and b e e t l e f l i g h t d i r e c t i o n was upwind, t o t h e west, a t a l l t h r e e d i s t a n c e s . However, b e e t l e f l i g h t d i r e c t i o n a t 100 m i n r e p l i c a t e 3 was n o t s i g n i f i c a n t . When t h e wind d i r e c t i o n changed t o t h e s o u t h e a s t i n r e p l i c a t e 4, f l i g h t d i r e c t i o n a t 5, 25, and 100 m, was t o t h e s o u t h e a s t , n o t s i g n i f i c a n t , and t o t h e n o r t h e a s t , r e s p e c t i v e l y . I n r e p l i c a t e 5, n e i t h e r w ind d i r e c t i o n n o r b e e t l e f l i g h t d i r e c t i o n a t 5 and 25 m were s i g n i f i c a n t . I n t h e l a s t r e p l i c a t e , w i nd d i r e c t i o n was s o u t h e a s t e r l y , as was b e e t l e f l i g h t d i r e c t i o n a t 5 and 25 m, b u t n o t a t 100 m. The one-sample t e s t f o r mean b e e t l e f l i g h t d i r e c t i o n was used f o r comparisons w i t h t h e wind d i r e c t i o n . F o r t h e f i v e r e p l i c a t i o n s w i t h a s i g n i f i c a n t wind d i r e c t i o n , b e e t l e f l i g h t d i r e c t i o n was s i g n i f i c a n t l y d i f f e r e n t a t 5 m once, a t 25 m f o u r t i m e s , and a t 100 m t h r e e t i m e s ( T a b l e 6 ) . However, a t 25 m, t h e b e e t l e f l i g h t d i r e c t i o n was c l o s e t o Table 6. Relationship of f l i g h t d i r e c tion of T. Iineatum to wind dire c t i o n , using a one-sample test for the mean directional f l i g h t angle, on the University Endowment Lands, Vancouver, B.C., 1986. Flight Wind Distance Direction Direction Exp. Rep. (m) \ ± d 1 \ v s . ^ 3 5 259 + 7 NS' 1 1 25 284 + 9 260 * 100 261 + 18 NS 5 286 + 10 NS 1 2 25 251 + 14 281 * 100 264 + 18 NS 5 256 + 22 * 1 3 25 242 + 16 218 * 100 NS 5 105 + 25 NS 1 4 25 NS 128 100 48 + 16 * 5 NS 1 5 25 142 + 45 NS 100 322 + 37 5 151 + 28 NS 1 6 25 149 + 28 155 NS 100 14 + 50 * 2 1 100 NS NS 2 2 100 12 + 23 275 * 2 3 100 236 + 29 186 * 2 4 100 8 + 41 235 * 3 1 500 90 + 35 94 NS 3 2 500 65 ± 10 117 * 3 3 500 67 + 25 95 * J&k + d = mean f l i g h t direction of beetles (rounded to the nearest deg.) + 95% C.L.; NS = not s i g n i f i c a n t . J?W = mean wind direction for experiments 1 and 2; and mean wind direction - 180° for experiment 3; NS = not s i g n i f i c a n t . Significance levels indicated: * = p < 0.05; NS = not s i g n i f i c a n t . 88 t h e w ind d i r e c t i o n on a l l o c c a s i o n s , whereas a t 100 m, when b e e t l e f l i g h t d i r e c t i o n was not s i g n i f i c a n t , i t t e n d e d t o be v e r y d i f f e r e n t from wind d i r e c t i o n . These d a t a i n d i c a t e upwind f l i g h t a t 5 m, and re d u c e d d i r e c t i o n a l i t y a t i n c r e a s i n g d i s t a n c e s from t h e r e l e a s e p o i n t . We must a l s o c o n s i d e r t h a t a l t h o u g h wind was s i g n i f i c a n t l y d i r e c t e d i n 5 o f t h e 6 r e p l i c a t i o n s , b e e t l e c a t c h e s d i d o c c u r i n a l l d i r e c t i o n s . T h i s s u g g e s t s t h a t some b e e t l e s were n o t r e s p o n d i n g i m m e d i a t e l y t o t h e pheromones and f l e w n o n - d i r e c t i o n a l l y w i t h r e s p e c t t o wind u n t i l t h e y p e r c e i v e o r respond t o t h e pheromone plumes. The numbers o f b e e t l e s caught a t 5, 25, and 100 m from t h e r e l e a s e p o i n t d i d n o t d i f f e r s i g n i f i c a n t l y d e s p i t e i n c r e a s e d i n t e r t r a p s p a c i n g s o f 8, 20, and 32 m, r e s p e c t i v e l y ( T a b l e 7 ) . T h i s s u g g e s t s t h a t a l t h o u g h some b e e t l e s a r e i m m e d i a t e l y a t t r a c t e d t o o l f a c t o r y s t i m u l i , o t h e r s p r e f e r r e d t o f l y b e f o r e r e s p o n d i n g o r t h e y may not have e n c o u n t e r e d t h e plume u n t i l r e a c h i n g t h e o u t e r t r a p s . I t must be n o t e d t h a t a l t h o u g h f l i g h t e x e r c i s e has been documented as a n e c e s s a r y p r e r e q u i s i t e f o r T. l i n e a t u m t o respond t o s e m i o c h e m i c a l s d u r i n g s p r i n g d i s p e r s a l (Graham 1959; B e n n e t t and Borden 1971), t h e b e e t l e s used i n t h i s s t u d y had p r e v i o u s l y f l o w n and responded t o pheromone-baited t r a p s . Thus i t i s p o s s i b l e t h a t t h e s e b e e t l e s responded more i m m e d i a t e l y t o t h e pheromone t h a n b e e t l e s j u s t emerging from t h e f o r e s t d u f f on t h e i r f i r s t f l i g h t m i ght have done. T h i s was assumed n o t t o be c r i t i c a l i n i n t e r p r e t i n g t h e 89 d a t a , because a wind t u n n e l e x p e r i m e n t d e s c r i b e d i n Appendix 2, shows m i n i m a l d i f f e r e n c e s between f l o w n and u n f l o w n T. l i n e a t u m i n t h e response t o s e m i o c h e m i c a l - b a i t e d f u n n e l t r a p s . E x p e r i m e n t 2 I n c i d e n c e o f r e c a p t u r e dropped by 50% when t h e 5 and 25 m t r a p s were removed from t h e p l o t ( T a b l e 5 ) . L e s s t h a n 8% o f t h e r e c a p t u r e d b e e t l e s were c o l l e c t e d d u r i n g t h e 24 h f o l l o w i n g t h e f i r s t c o l l e c t i o n . However, more b e e t l e s were caught a t 100 m i n t h i s e x periment t h a n i n t h e p r e v i o u s one (Ta b l e 7 ) . W i t h o u t t h e p r e s e n c e o f o l f a c t o r y s t i m u l i n e a r t h e r e l e a s e s i t e , a h i g h e r p e r c e n t a g e o f b e e t l e s f l e w f a r t h e r t o g e t t o a t t r a c t i v e m a t e r i a l . D i r e c t i o n a l d a t a show t h a t wind d i r e c t i o n and b e e t l e f l i g h t d i r e c t i o n were b o t h random i n r e p l i c a t e 1 ( F i g . 1 8 ) . I n r e p l i c a t e s 2, 3, and 4, wind d i r e c t i o n and b e e t l e f l i g h t d i r e c t i o n were a l l s i g n i f i c a n t , however t h e mean f l i g h t d i r e c t i o n s were n e i t h e r upwind n o r downwind. They were s i g n i f i c a n t l y d i f f e r e n t from each o t h e r i n a l l t h r e e c a s e s ( T a b l e 6 ) . D e s p i t e s t a t i s t i c a l l y s i g n i f i c a n t b e e t l e f l i g h t d i r e c t i o n s , c l o s e e x a m i n a t i o n o f t h e c i r c u l a r h i s t o g r a m s i n F i g . 18 shows t h a t t h e b e e t l e s were caught i n a l l d i r e c t i o n s f o r each o f t h e f o u r r e p l i c a t i o n s . I n g e n e r a l , t h e r e was no c o r r e l a t i o n between b e e t l e f l i g h t d i r e c t i o n and w i n d d i r e c t i o n . The random movement o f b e e t l e s may be e x p l a i n e d by t h e absence o f pheromones n e a r t h e r e l e a s e s i t e and t h e 90 T a b l e 7. P e r c e n t o f T. l i n e a t u m caught a t v a r i e d d i s t a n c e s i n t h e m a r k - r e c a p t u r e s t u d y , e x p e r i m e n t s 1 and 2, U n i v e r s i t y Endowment Lands, Vancouver, B.C., 1986. No. o f D i s t a n c e % R e l e a s e d B e e t l e s Exp. R e p l i c a t e s (m) R e c a p t u r e d + S.E. 5 8.1 + 1.5 a 2 1 6 25 9.7 + 0.8 a 100 7.2 + 0.7 a B 3 2 4 100 12.4 + 0.4 A Data s u b j e c t e d t o a r c s i n e t r a n s f o r m a t i o n . D i f f e r e n t l o w e r c a s e l e t t e r s r e p r e s e n t s i g n i f i c a n t d i f f e r e n c e s w i t h i n an e x p e r i m e n t (SNK; p < 0.05). D i f f e r e n t c a p i t a l l e t t e r s r e p r e s e n t s i g n i f i c a n t d i f f e r e n c e s i n b e e t l e c a t c h a t 100 m between e x p e r i m e n t s (SNK; p < 0.05). 91 CIRCULAR TRAPLINES WIND DIRECTION EXP REP 100m 500m F i g . 18 D i r e c t i o n a l d a t a based on f r e q u e n c y d i s t r i b u t i o n s o f T. l i n e a t u m f l i g h t and wind o b s e r v a t i o n s f o r e x p e r i m e n t s 2 and 3 i n t h e U n i v e r s i t y Endowments Land s t u d y . Dashed l i n e i n c i r c l e s r e p r e s e n t s mean d i r e c t i o n . Numbers i n c i r c l e s a r e t h e r - v a l u e s . Those marked w i t h a s t a r a r e s i g n i f i c a n t (p < 0.05). The d o t t e d l i n e s and a s s o c i a t e d v a l u e s o u t s i d e t h e c i r c l e s a r e t h e s c a l e s f o r t h e f r e q u e n c y b a r s . 92 c o n s i s t e n t p r e s e n c e o f low wind speeds, w h i c h a v e r a g e d 1.5 km/h t h r o u g h o u t t h i s e x p e r i m e n t . T h i s c o n t r a s t s somewhat w i t h Gara (1963), who i n v e s t i g a t e d I p s c o n f u s u s (Lec.) d i s p e r s a l , a l s o i n t h e absence o f o l f a c t o r y s t i m u l i n e a r h i s r e l e a s e s i t e . A t a f o r e s t s i t e t h a t e x p e r i e n c e d t w i c e t h e w i nd speed as o u r s , he d e t e r m i n e d t h a t t h e weaker f l y i n g I . c o n f u s u s g e n e r a l l y f l e w downwind u n t i l r e s p o n d i n g t o o l f a c t o r y s t i m u l i , whereupon t h e y o r i e n t e d upwind. D i f f e r e n c e s between t h e d i r e c t i o n a l d a t a f o r e x p e r i m e n t s 1 and 2 a t 100 m may be due t o t h e i n f l u e n c e o f t h e c l o s e - r a n g e t r a p s i n e x p e r i m e n t 1. W i t h upwind f l i g h t r e c o r d e d a t 5 and 25 m, we can assume t h a t b e e t l e s f l y i n g f u r t h e r d i s t a n c e s may have a l s o been i n f l u e n c e d i n i t i a l l y t o f l y upwind by t h e i r d e t e c t i o n o f t h e t r a p s c l o s e r t o t h e r e l e a s e p o i n t . As a r e s u l t , upwind f l i g h t was r e c o r d e d i n r e p l i c a t e s 1 and 2 o f e xperiment 1, a l t h o u g h t h e o t h e r r e p l i c a t e s d i d n o t e x h i b i t t h e same r e s p o n s e s . E x p e r i m e n t 3 The p r o p o r t i o n o f b e e t l e s r e c a p t u r e d (1.5%) was g r e a t l y r e d u c e d when t r a p s were p l a c e d 500 m from t h e r e l e a s e p o i n t ( T a b l e 5 ) . L e s s t h a n 10% o f t h e r e c a p t u r e d b e e t l e s were c o l l e c t e d d u r i n g t h e 24 h f o l l o w i n g t h e f i r s t c o l l e c t i o n . A l t h o u g h t h e d i s t a n c e p r o b a b l y p l a y e d a major r o l e i n t h e low numbers r e c a p t u r e d , a n o t h e r cause may have been t h e d i s t a n c e s between t h e t r a p s a l o n g t h e p e r i m e t e r o f t h e p l o t , 93 (800 m i n r e p l i c a t e 1 and 400 m i n r e p l i c a t e s 2 and 3 ) , w h i c h l e f t l a r g e a r e a s o f open space. The d i r e c t i o n a l d a t a f o r t h e b e e t l e s t h a t were caught showed s t r o n g e v i d e n c e o f downwind movement. I n r e p l i c a t e 1, t h e wind was b l o w i n g from t h e west and t h e b e e t l e s were caught t o t h e e a s t o f t h e r e l e a s e p o i n t ( F i g . 1 8 ) . I n r e p l i c a t e 2, d i r e c t i o n o f wind came from t h e n o r t h w e s t w h i l e b e e t l e c a t c h was t o t h e n o r t h e a s t . Wind d i r e c t i o n i n r e p l i c a t e 3, was from t h e west and b e e t l e c a t c h was t o t h e n o r t h e a s t . B e e t l e c a t c h d i r e c t i o n a t 500 m i n r e p l i c a t e 1 was n o t s i g n i f i c a n t l y d i f f e r e n t from wind d i r e c t i o n , y e t was i n r e p l i c a t e s 2 and 3 (Table 6 ) . However i n t h e l a t t e r two r e p l i c a t i o n s , f l i g h t and wind d i r e c t i o n s were f a i r l y c l o s e . These d a t a s u g g e s t t h a t b e e t l e s caught a t t h e 500 m t r a p s i n one day w i t h i n t h e homogeneous f o r e s t s e t t i n g , were ones t h a t were f l y i n g w i t h t h e wind. I t i s assumed from e x p e r i m e n t 2 d a t a , t h a t b e e t l e d i s p e r s a l p a t t e r n w i t h i n t h e f o r e s t , where l i g h t wind speeds p r e d o m i n a t e , i s n o n - d i r e c t e d w i t h r e s p e c t t o wind w i t h o u t t h e p r e s e n c e o f o l f a c t o r y s t i m u l i n e a r t h e r e l e a s e s i t e . B e e t l e s f l y i n g c r o s s w i n d o r upwind p r o b a b l y c o u l d n o t make i t as f a r as t h e ones c a r r i e d o r h e l p e d by t h e wind. I n c o n c l u s i o n , e x p e r i m e n t 1 showed t h a t s p r i n g d i s p e r s i n g Tj, l i n e a t u m f l y upwind toward p r o x i m a t e o l f a c t o r y s t i m u l i ( i e . 0 - 25 m) w i t h i n t h e f o r e s t . Upwind f l i g h t a t 25 and even 100 m may have been i n f l u e n c e d by t h e 5 m t r a p s . C a t c h e s o f b e e t l e s i n a l l t r a p s , however, s u g g e s t e d t h a t 94 some b e e t l e s d i d not p e r c e i v e o r r e s pond a t t a k e - o f f t o o l f a c t o r y s t i m u l i and f l e w i n an unknown manner u n t i l t h e y d i d r e s p o n d . A t 100 m, t h e p a t t e r n o f t r a p p e d b e e t l e s appeared more random and i n c o n s i s t e n t i n r e l a t i o n t o w ind d i r e c t i o n a t t h e r e l e a s e p o i n t . I n e x p e r i m e n t 2, where no o l f a c t o r y s t i m u l i were p r e s e n t near t h e r e l e a s e s i t e , numbers o f b e e t l e s caught a t 100 m were n o t r e l a t e d t o wind d i r e c t i o n . A t 500 m, t h e p a t t e r n o f b e e t l e r e c a p t u r e s u g g e s t e d t h a t t h e few r e c a p t u r e d were a s s i s t e d i n t h e i r f l i g h t by t h e wind. N o n - d i r e c t i o n a l w ind movement c o r r e s p o n d e d w i t h n o n - d i r e c t i o n a l f l i g h t b o t h t i m e s t h a t i t o c c u r r e d i n t h e s t u d y . Graham (1959) f i r s t n o t i c e d t h a t p h y s i c a l a c t i v i t y , s uch as f l i g h t , was needed f o r s p r i n g emerging T. l i n e a t u m b e f o r e t h e y would respond t o o l f a c t o r y s t i m u l i . He s u g g e s t e d t h a t t h e i n i t i a l d o m i n a t i o n o f p o s i t i v e p h o t o t a c t i c b e h a v i o r s e r v e d as a mechanism t h a t f o r c e d t h e i n s e c t s t o d i s p e r s e away from t h e emergence s i t e s t o f i n d s u s c e p t i b l e h o s t m a t e r i a l . T h i s b e h a v i o r has a l s o been n o t e d f o r o t h e r s c o l y t i d s (Henson 1962; A t k i n s 1966a; Choudhury and Kennedy 1980). These r e s u l t s a l l o w e d me t o b u i l d on Graham's h y p o t h e s i s . I n a c l o s e d canopy f o r e s t a t w i nd speeds a v e r a g i n g l e s s t h a n 2 km/h, T. l i n e a t u m appeared t o d i s p e r s e i n a d i f f u s e p a t t e r n when pheromone s o u r c e s were no c l o s e r t h a n 100 m away from t h e i r p o i n t o f r e l e a s e . W i t h pheromone d i s t a n c e s s e t 500 m from t h e i r p o i n t o f r e l e a s e , t h e f u r t h e s t f l y i n g b e e t l e s were i n f l u e n c e d by t h e d i r e c t i o n 95 o f p r e v a i l i n g winds i n t h e form o f downwind d i s p l a c e m e n t . These o b s e r v a t i o n s s u g g e s t t h a t i n n a t u r a l f o r e s t s e t t i n g s , where wind speed i s c o n s i s t e n t l y low, and s u i t a b l e h o s t m a t e r i a l such as w i n d f a l l s , were w i d e l y s c a t t e r e d , b e e t l e s w i l l f l y i n a n o n - d i r e c t i o n a l manner w i t h r e s p e c t t o wind. Upon p e r c e p t i o n o f t h e pheromones as f a r as 25 m away, p h y s i o l o g i c a l l y r e a d y T. l i n e a t u m r e s p o n d by f l y i n g upwind, s u g g e s t i n g chemo-anemotactic b e h a v i o r . T h e i r i n i t i a l f l i g h t p a t t e r n as w e l l as r e s p o n s e t o t h e a g g r e g a t i o n pheromone, s e r v e s t o m i n i m i z e t h e i r e x p e n d i t u r e o f energy, w h i l e m a x i m i z i n g t h e i r chance o f s u c c e s s f u l l y f i n d i n g and c o l o n i z i n g a n o r m a l l y s c a r c e r e s o u r c e . 96 CHAPTER 6: DISPERSAL OF TRYPODENDRON LINEATUM (OLIVIER) WITHIN A VALLEY SETTING 6.1 I n t r o d u c t i o n E x a m i n a t i o n o f t h e movement o f s p r i n g f l y i n g Trypodendron l i n e a t u m ( O l i v i e r ) i n a second-growth f o r e s t (Chapter 5) s e r v e d as an i m p o r t a n t i n i t i a l s t u d y o f t h e d i s p e r s a l p a t t e r n o f t h i s s p e c i e s . U s i n g t h e s e r e s u l t s , I d e s i g n e d a second s e t o f s t u d i e s t h a t examined p o p u l a t i o n d i s p e r s a l i n a more t y p i c a l s e t t i n g . C o n s e q u e n t l y , t h e s t u d y was moved t o a v a l l e y , p h y s i o g r a p h i c a l l y c h a r a c t e r i s t i c o f t h e f o r e s t s a l o n g t h e c o a s t o f B r i t i s h C olumbia ( P o j a r 1983). T h i s v a l l e y had a l t e r n a t e p a t c h e s o f o l d - g r o w t h and r e g e n e r a t i o n f o r e s t s i t e s t h a t p r o v i d e d a good s i m u l a t i o n o f commercial f o r e s t s i n t h i s r e g i o n . As wind was shown t o be a v e r y i m p o r t a n t f a c t o r w i t h r e s p e c t t o d i s p e r s a l d i r e c t i o n by T. l i n e a t u m ( C h a p t e r s 2 and 5 ) , t h i s v a l l e y p r o v i d e d a u n i q u e o p p o r t u n i t y f o r s t u d y i n g p o p u l a t i o n movement, because o f t h e c h a r a c t e r i s t i c d i u r n a l u p - v a l l e y w i nd p a t t e r n s t h a t a r e e x p e c t e d d u r i n g t h e summer ( B a r r y 1981). The o b j e c t i v e s o f t h e s t u d y r e p o r t e d i n t h i s c h a p t e r were; 1) make m e t e o r o l o g i c a l comparisons between an open and f o r e s t e d s i t e w i t h i n t h e v a l l e y ; 2) t e s t i f s h o r t d i s t a n c e f l i g h t (5 - 50 m) o f T. l i n e a t u m i n an o l d - g r o w t h f o r e s t i s upwind; 3) a s c e r t a i n l o n g d i s t a n c e f l i g h t c a p a b i l i t i e s o f 97 t h e b e e t l e s ; and 4) s u r v e y t h e i r d i s p e r s a l r a t e ( d i s t a n c e / t i m e ) . 6.2 M a t e r i a l s and Methods S i t e P arameters M a r k - r e c a p t u r e e x p e r i m e n t s were c a r r i e d o ut i n t h e Cedar Creek V a l l e y , o f t h e C o q u i t l a m Lake Watershed D i s t r i c t , i n t h e l o w e r c o a s t a l m a i n l a n d o f B r i t i s h Columbia. I t was a t y p i c a l U-shaped v a l l e y w i t h an u p - v a l l e y a z i m u t h r a n g i n g from 48 t o 68° ( F i g . 1 9 ) . A 3 km a r e a s e r v e d as our w o r k i n g a r e a . W i t h i n t h e s t u d y a r e a , t h e v a l l e y bottom e l e v a t i o n dropped from 600 t o 500 m. The peaks on b o t h s i d e s o f t h e v a l l e y r e a c h e d 1200 m i n e l e v a t i o n . The f o r e s t c o v e r was p r e d o m i n a n t l y over-mature w e s t e r n r e d c e d a r , Thuja  p l i c a t a Donn. and w e s t e r n hemlock, Tsuga h e t e r o p h y l l a (Raf.) S a r g . As a r e s u l t o f s t a n d breakdown, u n d e r s t o r y v e g e t a t i o n was v e r y dense, c o n s i s t i n g m o s t l y o f s a p l i n g s t a g e a m a b i l i s f i r , A b i e s a m a b i l i s (Dougl.) F o r b e s . C l e a r c u t a r e a s 35 t o 50 ha i n s i z e , r e g e n e r a t e d 8 t o 10 y e a r s ago, o c c u r i n a p a t c h - l i k e d i s t r i b u t i o n a l o n g t h e l o w e r s l o p e s o f t h e v a l l e y ( F i g . 1 9 ) . The e x p e r i m e n t a l work was c o n c e n t r a t e d a l o n g t h e n o r t h s i d e o f t h e c r e e k . Moving up t h e v a l l e y , a l o n g t h e r o a d , t h e f i r s t o p e n i n g on t h e n o r t h s i d e spans 700 m i n l e n g t h . I t i s f o l l o w e d by 700 m o f o l d growth, t h e n a n o t h e r 700 m o p e n i n g , a 200 m o l d growth p a t c h , and t h e n a 900 m o p e n i n g , EXPERIMENTS 2 ® RELEASE SITES(A.B) i WEATHER STATION(C) O L D G R O W T H ROAD REGENERATION CREEK F i g . 19. A p e r s p e c t i v e v i e w o f Cedar Creek V a l l e y , C o q u i t l a m Lake Watershed, B r i t i s h Columbia. F i v e m a r k - r e c a p t u r e e x p e r i m e n t s f o r s t u d y i n g T. l i n e a t u m d i s p e r s a l were c a r r i e d o u t i n t h i s v a l l e y from June-August, 1987. The u p - v a l l e y a z i m u t h r a n g e s from 48 t o 68°. The r o a d moving from t h e so u t h w e s t t o t h e n o r t h e a s t c o v e r s a d i s t a n c e o f 4 km i n t h i s v i e w . S l o p e peaks r e a c h 1200 m i n e l e v a t i o n , w h i l e c r e e k e l e v a t i o n averages 550 m. 00 where t h e r o a d t h e n c r o s s e s t h e c r e e k on t o t h e s o u t h e r n s i d e o f t h e v a l l e y i n t o a n o t h e r l a r g e o p e n i n g ( F i g . 1 9 ) . The most r e c e n t l o g g i n g (1986/1987) o c c u r r e d 1 km s o u t h e a s t o f t h e d o w n - v a l l e y edge o f our w o r k i n g a r e a , and t h u s was no t c o n s i d e r e d a competing s o u r c e o f a t t r a c t i o n f o r t h e marked amb r o s i a b e e t l e s . However, r e c e n t windthrow was p r e s e n t a l o n g t h e e a s t e r n edges o f t h e o p e n i n g s , and i n a s u r v e y made b e f o r e c o n d u c t i n g t h e e x p e r i m e n t s , a m b r o s i a b e e t l e a t t a c k s on t h e down t r e e s were n o t e d . T h i s w i l l be d i s c u s s e d l a t e r . M u l t i p l e - f u n n e l t r a p s b a i t e d w i t h l i n e a t i n , t h e a g g r e g a t i o n pheromone o f T. l i n e a t u m , were used t o c a p t u r e b e e t l e s f o l l o w i n g t h e i r r e l e a s e . Two Hereon® c o n t r o l l e d r e l e a s e d i s p e n s e r s , w i t h a t o t a l r e l e a s e r a t e o f 340 ug/24 b h16-, were p l a c e d i n each t r a p . R e l e a s e s i t e s and t r a p placement were i n s t a l l e d a c c o r d i n g t o e x p e r i m e n t a l o b j e c t i v e s . F i v e e x p e r i m e n t s were conducted t o t e s t v a r i o u s h y p o t h e s e s c o n c e r n i n g T. l i n e a t u m r e s p o n s e and f l i g h t t o ward pheromone-baited t r a p s i n f o r e s t e d and open s i t e s . E x p e r i m e n t 1 B e e t l e r e c a p t u r e d i r e c t i o n was e v a l u a t e d a t d i s t a n c e s o f 25 and 50 m from t h e r e l e a s e p o i n t w i t h r e s p e c t t o wind, i n a dense o l d - g r o w t h f o r e s t s e t t i n g . I n C h a p t e r 5, a l t h o u g h upwind f l i g h t was ob s e r v e d a t 25 m, t h e 5 m t r a p s i n p l a c e may have had an i n f l u e n c e on t h e 25 m c a t c h e s . T h i s e x p e r i m e n t c o r r e c t s f o r t h a t , w i t h 25 m as t h e c l o s e s t 100 s o u r c e o f a t t r a c t i o n . The two c i r c u l a r t r a p l i n e s were s e t up a t r e l e a s e s i t e A ( F i g . 19). A t 25 m, 8 t r a p s were p l a c e d a t a n g u l a r i n t e r v a l s o f 45°, s t a r t i n g a t an a z i m u t h o f 22.5°. A t 50 m, 16 t r a p s were p l a c e d a t a n g u l a r i n t e r v a l s o f 22.5°, s t a r t i n g a t an a z i m u t h o f 0°. Three r e l e a s e s were r u n u s i n g t h i s d e s i g n . E x p e r i m e n t 2 The low p r o p o r t i o n o f marked b e e t l e s r e c a p t u r e d i n e x p e r i m e n t 1 prompted experiment 2, w h i c h was d e s i g n e d t o maximize b e e t l e c a t c h . The 25 and 50 m t r a p l i n e s were k e p t i n p l a c e , w i t h an a d d i t i o n a l 4 t r a p s p l a c e d 5 m from r e l e a s e p o i n t A, a t a n g u l a r i n t e r v a l s o f 90°, s t a r t i n g a t an a z i m u t h o f 0°. Meanwhile, a f i r s t a t t e m pt a t l o n g d i s t a n c e f l i g h t r e c a p t u r e was made w i t h t r a p l i n e s o f 10 t r a p s each, p l a c e d 700 m west ( t r a p l i n e W) and 700 m e a s t ( t r a p l i n e E-B) o f r e l e a s e s i t e A ( F i g . 19). F o r b o t h t h e s e l i n e s , 4 t r a p s were p l a c e d above t h e r o a d and 6 below i t , w i t h each t r a p a p p r o x i m a t e l y 2 0 - 35 m a p a r t . Three r e l e a s e s were r u n u s i n g t h i s d e s i g n . E x p e r i m e n t 3 F o r e x p e r i m e n t s 3, 4, and 5, t h e r e l e a s e s i t e (B, F i g . 19) was moved t o t h e edge o f t h e f o r e s t , n e a r t h e r o a d and a r e g e n e r a t i o n o p e n i n g . T h i s s i t e was chosen t o b e t t e r s i m u l a t e a b e e t l e emergence s i t e , s i n c e f a l l d i s p e r s i n g b e e t l e s f a v o r f o r e s t edges f o r o v e r w i n t e r i n g (Dyer and 101 K i n g h o r n 1961; L i n d g r e n and Borden 1983). T h i s s i t e was a l s o l e s s dense v e g e t a t i v e l y t h a n s i t e A, p o s s i b l y e n h a n c i n g t h e r e c a p t u r e o f marked b e e t l e s . I n e x p e r i m e n t 3, i n c r e a s e d emphasis was p l a c e d on a s s e s s i n g l o n g d i s t a n c e f l i g h t c a p a b i l i t i e s , i n r e l a t i o n t o t h e c h a r a c t e r i s t i c u p - v a l l e y w i n d s , w h i c h o c c u r r e d t h r o u g h o u t t h e summer. A c i r c u l a r t r a p l i n e was d e p l o y e d 2 5 m from t h e r e l e a s e p o i n t B, u s i n g t h e same a z i m u t h s e t t i n g s as i n e x p e r i m e n t s 1 and 2. L i n e a r t r a p l i n e s were d e p l o y e d a t t h e west edge, c e n t e r , and e a s t edge o f t h e a d j a c e n t o p e n i n g , 50 m ( t r a p l i n e E-A), 375 m ( t r a p l i n e E-B), and 700 m ( t r a p l i n e E-C), e a s t o f r e l e a s e s i t e B, r e s p e c t i v e l y . T r a p l i n e W from e x p e r i m e n t 2 was a l s o used, and was 1.0 km west o f r e l e a s e s i t e B. Four r e l e a s e s u t i l i z i n g t h i s d e s i g n were c a r r i e d o u t . Ex p e r i m e n t 4 B e e t l e f l i g h t d i s p e r s a l o u t s i d e t h e f o r e s t was t h e f o c u s o f t h i s e x p e r i m e n t . T r a p l i n e s W, E-A, and E-C were used, as w e l l as t h r e e a d d i t i o n a l ones p l a c e d f u r t h e r up-v a l l e y . Two t r a p l i n e s (E-D and E-E) were s e t up 1.1 and 1.9 km e a s t o f r e l e a s e s i t e B, r e s p e c t i v e l y ( F i g . 1 9 ) . L i n e E-F was e s t a b l i s h e d on t h e s o u t h s i d e o f t h e v a l l e y c a . 1.3 km from r e l e a s e s i t e B. Four t r a p s were p l a c e d above t h e r o a d and 2 below i t f o r l i n e s E-D and E-E. Three t r a p s were p l a c e d above and below t h e r o a d f o r l i n e E-F. Only two r e l e a s e s o f t h i s d e s i g n were r u n i n t h i s e x p e r i m e n t . 102 E x p e r i m e n t 5 T h i s e x p e r i m e n t had t h e same d e s i g n as e x p e r i m e n t 4 e x c e p t t h a t t r a p l i n e E-A was removed. I t h o u g h t t h a t by removing t h e s e t r a p s , l o n g d i s t a n c e f l i g h t would be b e t t e r f a c i l i t a t e d . U n f o r t u n a t e l y o n l y one r e l e a s e was c a r r i e d o u t due t o t h e l a c k o f b e e t l e s . B e e t l e P r e p a r a t i o n A d u l t T. l i n e a t u m used i n t h i s s t u d y were c o l l e c t e d d u r i n g t h e i r mass f l i g h t p e r i o d o f May and June, 1987, u s i n g l i n e a t i n - b a i t e d m u l t i p l e f u n n e l t r a p s p l a c e d a d j a c e n t t o a l a r g e l o g boom s t o r a g e a r e a a t t h e mouth o f t h e n o r t h arm o f t h e F r a s e r R i v e r , i n Vancouver, B.C. B e e t l e c o l l e c t i o n and s t o r a g e p r o c e d u r e s were s i m i l a r t o t h o s e f i r s t d i s c u s s e d i n Ch a p t e r 2. The p r o c e d u r e s used f o r mark i n g b e e t l e s w i t h f l u o r e s c e n t powder were d e s c r i b e d i n Ch a p t e r 5. Four c o l o r e d powders were used t h r o u g h o u t t h e summer, w i t h a d i f f e r e n t c o l o r used f o r each s u c c e s s i v e r e l e a s e . R e l e a s e - R e c a p t u r e The r e l e a s e - r e c a p t u r e p r o c e d u r e s used f o r a l l 5 e x p e r i m e n t s were v e r y s i m i l a r . B e e t l e s were u s u a l l y r e l e a s e d between 1200 and 1300 h PDT o r when t h e t e m p e r a t u r e r o s e above 15°C, t h e low t e m p e r a t u r e f l i g h t t h r e s h o l d f o r T. l i n e a t u m . Between 2000 t o 7000 b e e t l e s were p l a c e d on a 1 103 m wooden t r a y , w i t h a s m a l l frame r i d g e 0.2 m from t h e edge, e f f e c t i v e l y g i v i n g t h e b e e t l e s two r i d g e s from w h i c h t o t a k e - o f f . The t r a y was s e t 0.5 m above t h e ground. An a b s o r b e n t paper, l i n e d on one s i d e w i t h c e l l o p h a n e , was p l a c e d u n d e r n e a t h t h e t r a y t o c a t c h any b e e t l e s t h a t f e l l o f f . S i m i l a r paper was used as a b e d d i n g f o r t h e i n n e r s q u are o f t h e t r a y , where t h e b e e t l e s were p l a c e d . I n e x p e r i m e n t s 1 - 3 , two b e e t l e c o l l e c t i o n s were made from t h e c i r c u l a r t r a p l i n e s d u r i n g t h e day o f r e l e a s e . F o r a l l o f t h e r e l e a s e s b u t t h e f i r s t one, c o l l e c t i o n s were made 2 hours a f t e r r e l e a s e and a t t h e end o f t h e day, u s u a l l y 4 ho u r s a f t e r r e l e a s e . The f i r s t r e l e a s e was 6 h o u r s l o n g w i t h t h e c o l l e c t i o n i n t e r v a l o c c u r r i n g a t 3 h o u r s . I n t h e e x p e r i m e n t s t h a t u t i l i z e d l i n e a r t r a p l i n e s (2 - 5 ) , b e e t l e s were c o l l e c t e d a t t h e end o f t h e r e l e a s e day. F o r a l l e x p e r i m e n t s , b e e t l e s were c o l l e c t e d a g a i n p r i o r t o t h e n e x t r e l e a s e (Day 2 c a t c h e s ) . Day 2 t r a p p i n g p e r i o d s ranged from 2 t o 12 days. The b e e t l e s caught i n t h e t r a p s were b r o u g h t back t o t h e l a b f o r c o u n t i n g , and e x a m i n a t i o n under an u l t r a - v i o l e t lamp t o i d e n t i f y marked specimens. The number o f marked males and fe m a l e s and unmarked b e e t l e s were t a l l i e d . The number o f r e l e a s e d b e e t l e s n o t f l y i n g was a l s o r e c o r d e d . 104 Weather Data Weather was m o n i t o r e d a t t h e r e l e a s e s i t e s as w e l l as i n an a d j a c e n t r e g e n e r a t i o n o p e n i n g . A t t h e open s i t e (C) ( F i g . 1 9 ) , a p o r t a b l e weather s t a t i o n was s e t up t o r e c o r d a i r t e m p e r a t u r e , r e l a t i v e h u m i d i t y , w ind d i r e c t i o n , and wind speed c o n t i n u o u s l y t h r o u g h o u t t h e summer. The s t a t i o n c o n s i s t e d o f a Campbell S c i e n t i f i c (C.S.) model 201 c o m b i n a t i o n t e m p e r a t u r e (-35 t o 50°C ) / r e l a t i v e h u m i d i t y (10 t o 95%) probe, f i t t e d w i t h a g i l l r a d i a t i o n s h i e l d ; a Weathermeasure model 2 02 0 m i c r o response wind vane ( ± 2 ° a c c u r a c y ; low t h r e s h o l d o f 0.8 km/h); and a model 2030 m i c r o r e s p o n s e anemometer (+ 0.24 km/h a c c u r a c y ; low t h r e s h o l d o f 0.8 km/h). These d a t a c o l l e c t i n g d e v i c e s were l a t c h e d on t o a c r o s s a r m , p l a c e d 2 m above t h e ground on a s t a i n l e s s s t e e l tower. The d a t a were c o l l e c t e d i n a C.S. CR21 m i c r o l o g g e r . The d a t a from t h e m i c r o l o g g e r were s t o r e d on a C.S. RC 235 c a s s e t t e r e c o r d e r . Measurement i n t e r v a l s were s e t a t 10 min t o c o i n c i d e w i t h manual weather measurements made a t t h e r e l e a s e s i t e . Data were r e a d from t h e c a s s e t t e t a p e s u s i n g a C.S. C 2 0 c a s s e t t e i n t e r f a c e i n t o U n i v e r s i t y o f B r i t i s h C olumbia computer main frame f i l e s . Problems w i t h t h i s system on d i f f e r e n t o c c a s i o n s r e s u l t e d i n no weather d a t a from t h e c l e a r i n g f o r 3 o f t h e 13 m a r k - r e c a p t u r e r e l e a s e s . A t t h e r e l e a s e s i t e , measurements were c a r r i e d o u t o n l y on t h e day b e e t l e s were r e l e a s e d . Wind d i r e c t i o n and speed d a t a were c o l l e c t e d a t 10 min i n t e r v a l s , w i t h t h e use o f a 105 bee-smoker p l a c e d o v e r a c i r c u l a r g r i d 1 m i n r a d i u s and b r o k e n up i n t o 16 a n g u l a r i n t e r v a l s o f 22.5° (360° c i r c l e ) . The s p o u t o f t h e smoker was o r i e n t e d v e r t i c a l l y , 0.5 m above t h e f o r e s t f l o o r . Wind speed was measured by t i m i n g t h e movement o f smoke f o r a d i s t a n c e o f 1 m. Wind p a r a m e t e r s were measured 10 m e a s t o f t h e r e l e a s e t r a y i n e x p e r i m e n t s 1 and 2, and 10 m n o r t h o f t h e r e l e a s e t r a y f o r e x p e r i m e n t s 3 t o 5. Temperature measurements were t a k e n e v e r y 10 min from a mercury thermometer s e t i n t h e shade. R e l a t i v e h u m i d i t y was measured h o u r l y w i t h a s l i n g p s y c h r o m e t e r . Data A n a l y s i s The c i r c u l a r t r a p l i n e s i n e x p e r i m e n t s 1 t o 3 and t h e wind measurements i n a l l t h e e x p e r i m e n t s a l l o w f o r t h e p r e s e n t a t i o n o f t h e b e e t l e c a t c h e s and wind d i r e c t i o n i n t h e form o f c i r c u l a r h i s t o g r a m s . The s t a t i s t i c a l p r o c e d u r e used i s d e s c r i b e d i n d e t a i l by B a t s c h e l e t (1981), and i s summarized i n Ch a p t e r 5. To t e s t whether b e e t l e f l i g h t d i r e c t i o n d i f f e r e d from wind d i r e c t i o n , a one-sample t e s t , analogous t o a one-sample t - t e s t f o r d a t a on a l i n e a r s c a l e (Zar 1984), was co n d u c t e d u s i n g wind d i r e c t i o n as a p r e - a s s i g n e d a n g l e t o be t e s t e d a g a i n s t t h e b e e t l e f l i g h t d i r e c t i o n a n g l e s . A + d - v a l u e was d e t e r m i n e d f o r each t e s t a n g l e , s e r v i n g as 95% c o n f i d e n c e i n t e r v a l s ( B a t s c h e l e t 1981). I f t h e p r e - a s s i g n e d a n g l e f e l l w i t h i n t h e c o n f i d e n c e l i m i t s o f t h e f l i g h t a n g l e , t h e a n g l e s were n o t c o n s i d e r e d t o be s i g n i f i c a n t l y d i f f e r e n t . 106 Frequency d i s t r i b u t i o n comparisons were made w i t h i n t e r v a l 1 and 2 d a t a a l o n g w i t h f u l l day ( t o t a l ) d a t a , however s i n c e d i f f e r e n c e s between i n d i v i d u a l i n t e r v a l s and t o t a l d a t a f o r a day were m i n i m a l , w i t h r e s p e c t t o d i r e c t i o n , t h e t o t a l d a t a c omparisons o n l y a r e p r e s e n t e d . A two-way a n a l y s i s o f v a r i a n c e (ANOVA) was r u n (SAS 1985) t o t e s t f o r d i f f e r e n c e s i n t h e number o f b e e t l e s caught a t t h e d i f f e r e n t d i s t a n c e s f o r e x p e r i m e n t s 1 and 2. A t e s t f o r b e e t l e c a t c h e s a t 25 m between e x p e r i m e n t s 1, 2, and 3 was r u n u s i n g a one-way ANOVA (SAS 1985). 6.3 R e s u l t s and D i s c u s s i o n Weather The h i g h p r e s s u r e system t h a t u s u a l l y s e t t l e s i n t o t h e west c o a s t d u r i n g J u l y and August, and b r i n g s i n warm s t a b l e a i r masses, d i d n o t come u n t i l t h e end o f August i n 1987. As a r e s u l t , t h e weather a l o n g t h e c o a s t a l mountains v a r i e d from day t o day, making i t d i f f i c u l t t o complete as many b e e t l e r e l e a s e s as d e s i r e d ; however, 13 were completed. D u r i n g b e e t l e r e l e a s e days, t h e m e t e o r o l o g i c a l measurements and o b s e r v a t i o n s made w i t h i n f o r e s t e d and open s e t t i n g s a l l o w e d f o r between s i t e c o m p a r i s o n s . S i n c e b e e t l e s were r e l e a s e d o n l y on warm days, t h e m e t e o r o l o g i c a l c omparisons between t h e d i f f e r e n t s e t t i n g s a r e o n l y a p p r o p r i a t e f o r d r y days i n w h i c h t h e t e m p e r a t u r e i n t h e f o r e s t r o s e above 15°C f o r a t l e a s t a c o u p l e o f h o u r s . F o r 107 t h e s e days, some i n t e r e s t i n g p a t t e r n s a r e a p p a r e n t . The t e m p e r a t u r e s i n s i t e A and B ( l e s s dense) averaged 3 and 2°C l e s s , r e s p e c t i v e l y , t h a n i n t h e open s e t t i n g , r e g a r d l e s s o f t h e s p e c i f i c average t e m p e r a t u r e s f o r t h e day. As e x p e c t e d , r e l a t i v e h u m i d i t y (R.H.) was h i g h e r under t h e canopy t h a n i n t h e open. F o r e s t s i t e s A and B averaged 2 0 and 10% h i g h e r R.H., r e s p e c t i v e l y , t h a n t h e open s i t e ( C ) . W h i l e R.H. d e c r e a s e d d u r i n g t h e day a t t h e open s i t e , i t d e c r e a s e d on a l l 6 measured o c c a s i o n s a t s i t e A and on 3 o f 5 o c c a s i o n s a t s i t e B. Wind i n t h e open s i t e was s i g n i f i c a n t l y d i r e c t e d up-v a l l e y on a l l measured o c c a s i o n s ( T a b l e 8 ) . The range o f mean d i r e c t i o n s t h e wind blew from was 198 - 236°, w h i l e t h e v a l l e y d i r e c t i o n toward Lake C o q u i t l a m a t t h a t s i t e i s 228°. Average wind speed d i f f e r e d l i t t l e between a l l measured days, r a n g i n g from 5.0 - 6.3 km/h ( T a b l e 8 ) . The v a r i a t i o n i n w i nd speed o b s e r v e d among r e l e a s e days was f a i r l y s m a l l , w i t h t h e range o f s t a n d a r d d e v i a t i o n s c a l c u l a t e d t o be 1.1 -1.6 km/h. I n c o n t r a s t , a t s i t e A, a l t h o u g h v a r i a b l e t h r o u g h o u t t h e day, average wind d i r e c t i o n t e n d e d t o be downslope toward t h e c r e e k ( T a b l e 8 ) . S i g n i f i c a n t (non-random) wind d i r e c t i o n was n o t e d on 3 o f 6 t e s t days. However t h i s m ight be somewhat d e c e i v i n g and w i l l be d i s c u s s e d i n a l a t e r s e c t i o n . Wind speed was l i g h t , a v e r a g i n g 1.1 - 1.3 km/h, s l i g h t l y l e s s t h a n t h a t a t t h e r e l e a s e s i t e i n t h e second-growth, even-aged f o r e s t r e p o r t e d i n C h a p t e r 5, and c a . 0.2 Table 8. Comparison of wind di r e c t i o n and speed between forested and open sites within the Cedar Creek valley (up-valley azimuth of 48 to 68°) in the Coquitlam Lake watershed, B.C., 1987. Forest Open Site X Uindspeed X Windspeed Exp. Rel. Julian S i t e N ~9 2 r 3 + S.O. N 0 r + S.D. K c Fw c Date (Obs.) (km/h) (Obs.) (km/h) 1 1 169 A 34 4 0.305 1.37 + 0.46 33 236 0.723* 6.05 + 1.44 2 175 24 333 0.417* 1.39 + 0.37 24 209 0.842* ... 4 3 177 26 355 0.426* 1.33 + 0.58 26 226 0.703* 5.94 + 1.26 2 1 181 A 27 334 0.281 1.12 + 0.60 27 219 0.802* 5.33 + 1.41 2 191 26 350 0.428* 1.35 + 0.56 -- ... ... ... 3 195 27 262 0.195 1.08 + 0.59 27 211 0.726* 6.32 + 1.30 3 1 202 B 25 220 0.763* 1.40 + 0.70 24 198 0.831* 6.12 + 1.26 2 205 7 201 0.515 0.65 + 0.46 -- ... ... ... 3 210 27 277 0.724* 1.23 + 0.46 -- ... ... ... 4 216 26 251 0.724* 1.88 + 0.75 25 234 0.672* 6.01 + 1.49 4 1 218 B 26 241 0.706* 1.43 + 0.36 26 207 0.810* 5.04 + 1.13 2 230 24 228 0.730* 1.42 + 0.56 24 201 0.768* 6.24 + 1.57 5 1 233 B 24 231 0.660* 1.54 + 0.60 24 206 0.800* 5.51 + 1.44 I S i t e A was dense and far from any openings; s i t e B was less dense and within 25 m of the road and 35 m from the clearcut. 2 — 0 represents mean wind d i r e c t i o n . 3 w — Stars represent a s i g n i f i c a n t 0 (Rayleigh's Test for Randomness; p < 0.05). 4 w -No data. 109 t i m e s t h e speed o b s e r v e d i n t h e open s i t e i n t h i s s t u d y . The range o f s t a n d a r d d e v i a t i o n s f o r t h e 6 r e l e a s e s was 0.4 - 0.6 km/h, v e r y s i m i l a r t o t h e second-growth f o r e s t s i t e . A t s i t e B, w h i c h was c l o s e t o t h e r o a d and t o a n o t h e r l a r g e o p e n i n g , and i n a d d i t i o n had a l e s s dense u n d e r s t o r y t h a n s i t e A, wind c o n s i s t e n t l y blew from down- t o u p - v a l l e y ( T a b l e 8 ) . Average wind d i r e c t i o n f o r 7 r e l e a s e days ranged from 201 - 277°, and on 6 o f t h e s e days, t h e d i r e c t i o n was s i g n i f i c a n t . Wind speed was s i m i l a r t o t h a t a t s i t e A and averaged from 0.7 - 1.9 km/h, w i t h s t a n d a r d d e v i a t i o n s o f 0.4 - 0.8 km/h. W e i g h t i n g i n d i v i d u a l wind d i r e c t i o n s by wind speed had a m i n i m a l e f f e c t on t h e c o m p u t a t i o n o f mean wind d i r e c t i o n (j? w) i n t h e f i r s t m a r k - r e c a p t u r e s t u d y (Chapter 5 ) . Thus i t was assumed t o be a s i m i l a r c a s e f o r r e l e a s e s i t e s A and B i n t h e v a l l e y , s i n c e t h e w i n d speed and i t s v a r i a t i o n were so s i m i l a r a t a l l t h r e e s i t e s . E x p e r i m e n t 1 B e e t l e r e c a p t u r e averaged 6.4, 8.3, and 5.8% f o r r e l e a s e s 1, 2, and 3, r e s p e c t i v e l y ( T a b l e 9 ) , c o n s i d e r a b l y l e s s t h a n from t h e f i r s t two e x p e r i m e n t s r e p o r t e d i n C h a p t e r 5. I n t h e f i r s t 2 r e l e a s e s , c a . 80% o f t h e b e e t l e s a t b o t h 25 and 50 m were r e c a p t u r e d d u r i n g t i m e i n t e r v a l 1 ( T a b l e 10) . T h i s changed f o r r e l e a s e 3, where c a . 1/2 o f t h e b e e t l e s were caught d u r i n g t h e f i r s t i n t e r v a l . Of a l l b e e t l e s r e c a p t u r e d , a p p r o x i m a t e l y 12% were r e c a p t u r e d d u r i n g t h e n e x t c o l l e c t i o n day f o l l o w i n g t h e day o f r e l e a s e . T h i s T a b l e 9. F l i g h t and r e c a p t u r e s u c c e s s o f T. l i n e a t u m i n t h e m a r k - r e c a p t u r e f l i g h t s t u d y a t t h e C o q u i t l a m Lake Watershed, B.C. (Day 1 and Day 2 d a t a p o o l e d ) , 1987. Captured B e e t l e T o t a l s  Marked Unmarked Exp. R e l . No. % That % of Flown R e l e a s e d Flew NO. B e e t l e s No. 1 1 2 000 81.5 105 6.4 3 569 2 2 340 81.5 158 8.3 2 490 3 2 175 86.0 109 5.8 2 092 T o t a l s 6 515 83.0 372 6.9 8 151 2 1 2 400 88.3 179 8.4 1 238 2 3 990 53.6 234 10.9 4 110 3 2 000 85.2 187 10.9 847 T o t a l s 8 390 71.1 600 10.1 6 195 3 1 2 400 48.8 83 7.1 1 820 2 5 000 76.8 501 13.1 828 3 5 000 61.3 192 6.3 1 322 4 4 600 67.8 178 5.3 287 T o t a l s 17 000 65.8 954 8.5 4 057 4 1 4 000 73.9 85 2.9 1 120 2 7 000 53.5 92 2.5 46 T o t a l s 11 000 60.9 177 2.6 1 166 5 1 4 900 81.6 17 0.4 191 Table 10. The number of marked T. Iineatum recaptured with respect to time i n a dense old growth forest i n the Coquitlam Lake Watershed, B.C., 1987. No. Beetles Recaptured Exp. Rel. Release Time Traplines Site I n t e r v a l 1 5 m 25 m 50 m N 5? N % N 1 A 1 31 77.5 52 80.0 2 2 5.0 3 4.6 Day 2 7 17.5 10 15.4 2 A 1 50 80.6 79 82.3 2 8 12.9 15 15.6 Day 2 4 6.5 2 2.1 3 A 1 29 46.0 24 51.1 2 24 38.1 16 34.0 Day 2 10 15.9 7 14.9 2 1 A 1 57 74.0 15 51.7 21 39.6 2 16 20.8 14 48.3 20 37.7 Day 2 14 18.2 0 0.0 12 22.6 2 A 1 50 54.3 32 45.1 34 48.6 2 30 32.6 21 29.6 24 34.3 Day 2 12 13.0 18 25.4 12 17.1 3 A 1 32 52.5 33 53.2 32 50.0 2 12 19.7 17 27.4 20 31.3 Day 2 17 27.9 12 19.4 12 18.8 3 1 B 1 34 45.3 2 5 6.7 Day 2 36 48.0 2 B 1 340 80.9 2 -- --Day 2 80 19.1 3 B 1 95 53.4 2 14 7.9 Day 2 69 38.8 4 B 1 124 75.2 2 16 9.7 Day 2 25 15.2 Time intervals: 1 = 1st half of day 1 ; 2 = f i n a l Day 2 = period u n t i l next c o l l e c t i o n (2 - 12 days Represents % of beetles caught at that distance. half of l a t e r ) . period; 112 i n d i c a t e s t h a t a t s h o r t d i s t a n c e s , t h e m a j o r i t y o f T. l i n e a t u m t h a t respond t o nearby o l f a c t o r y s t i m u l i ( i e . 25 -50 m) do so w i t h i n t h e f i r s t few hou r s o f r e l e a s e . The o v e r a l l number o f b e e t l e s r e c a p t u r e d d i d n o t d i f f e r s i g n i f i c a n t l y between 25 and 50 m, w i t h i n t e r t r a p s p a c i n g s o f 20 m f o r b o t h t r a p l i n e s ( T a b l e 1 1 ) . T h i s i s c o n s i s t e n t w i t h t h e d a t a from Chapter 5, where an e q u a l number o f b e e t l e s were c a p t u r e d a t d i s t a n c e s o f 5 t o 100 m from t h e r e l e a s e s i t e , i n d i c a t i n g t h a t w h i l e some b e e t l e s a r e a t t r a c t e d t o o l f a c t o r y s t i m u l i i m m e d i a t e l y , o t h e r s f l y f a r t h e r . D i r e c t i o n a l d a t a show t h a t f o r a l l 3 r e l e a s e s , t h e mean wind d i r e c t i o n ( 0 W ) a t S i t e A was from t h e n o r t h . R - v a l u e s were s i g n i f i c a n t f o r t h e l a t t e r 2 r e l e a s e s ( F i g . 2 0 ) . B e e t l e f l i g h t d i r e c t i o n (0^) a ^ 2 5 m w a s n o n - d i r e c t i o n a l f o r a l l 3 r e l e a s e s , w h i l e a t 50 m, 0^ was s i g n i f i c a n t l y d i r e c t e d e a s t t o s o u t h e a s t f o r a l l 3 r e l e a s e s . A one-sample t e s t was used t o compare 0^ a t 50 m t o £TW i n t h e f o r e s t f o r r e l e a s e s 2 and 3. F o r b o t h r e l e a s e s , 0-^ d i f f e r e d s i g n i f i c a n t l y w i t h # w ( T a b l e 12). Thus # b a t b o t h 25 and 50 m was n o t upwind. However, upon c l o s e e x a m i n a t i o n o f w ind d i r e c t i o n h i s t o g r a m s ( F i g . 20) f o r r e l e a s e s 2 and 3, i t was n o t i c e d t h a t t h e h i g h e s t f r e q u e n c y o f o b s e r v a t i o n s o c c u r r e d + 45° o f t h e mean a n g l e , w i t h few o b s e r v a t i o n s o c c u r r i n g a t t h e mean. T h e r e f o r e , d e s p i t e a s i g n i f i c a n t r -v a l u e , w ind d i r e c t i o n was q u i t e v a r i a b l e , and must be c o n s i d e r e d when i n t e r p r e t i n g t h e s e r e s u l t s . 113 T a b l e 11. Mean p e r c e n t o f T. l i n e a t u m r e c a p t u r e d a t v a r i e d d i s t a n c e s i n t h e Cedar Creek V a l l e y , C o q u i t l a m Lake Watershed, B.C., 1987. Mean % o f R e l e a s e No. o f D i s t a n c e Flown B e e t l e s Exp. S i t e R e l e a s e s (m) R e c a p t u r e d + S.E. 1 A 3 25 3.1 + 0.3 a 2 B 3 50 3.9 + 0.8 a 2 A 3 5 4.0 + 0.2 a 25 2.7 + 0.6 a B 50 3.1 + 0.4 a 3 B 4 25 7.1 + 1.3 A 1 Data s u b j e c t e d t o a r c s i n e t r a n s f o r m a t i o n . D i s t a n c e s w i t h i n each ex p e r i m e n t f o l l o w e d by d i f f e r e n t l o w e r c a s e l e t t e r s a r e s i g n i f i c a n t l y d i f f e r e n t (SNK; p < 0.05). D i f f e r e n t c a p i t a l l e t t e r s , w i t h r e s p e c t t o t h e number o f b e e t l e s caught a t 25 m f o r t h e 3 e x p e r i m e n t s , r e p r e s e n t s s i g n i f i c a n t d i f f e r e n c e s (SNK; p < 0.05). 114 T a b l e 12. Comparison o f T. l i n e a t u m f l i g h t and w i n d d i r e c t i o n i n an o l d growth f o r e s t i n t h e C o q u i t l a m Lake Watershed, B.C., 1987. E x p . 1 R e l . D i s t a n c e 0 B + d 2 0 W J S i g n i f i c a n c e (m) (p < 0.05) 1 25 50 138 NS ± 38 NS 2 25 NS 333 50 107 + 39 * 3 25 NS 355 50 103 + 37 * 1 5 25 50 91 ± 18 NS NS NS 2 5 25 2 ± 28 NS 350 NS 50 13 ± 32 NS 3 5 94 ± 21 25 56 + 39 NS 50 61 ± 41 1 25 205 + 36 220 NS 2 25 193 + 17 NS 3 25 272 ± 15 277 NS 4 25 234 ± 15 251 * Exps. 1 and 2 c a r r i e d o u t a t r e l e a s e s i t e A and exp 3 a t r e l e a s e s i t e B. 0 B ± d = mean f l i g h t d i r e c t i o n o f b e e t l e s ( t o t h e n e a r e s t degree) + 95% C.L. 0 W = mean wind d i r e c t i o n a t t h e r e l e a s e s i t e . D i f f e r e n c e between 0 B and 0^: * = s i g n i f i c a n t ; NS = n o t s i g n i f i c a n t . 115 EXP REL 1 1 2 1 3 1 CIRCULAR TRAPLINES 5 m 25 m 50 m WIND DIRECTION FOREST CLEAR CUT F i g . 20. D i r e c t i o n a l d a t a based on f r e q u e n c y d i s t r i b u t i o n s o f T. l i n e a t u m f l i g h t and wind o b s e r v a t i o n s f o r e x p e r i m e n t s 1 - 3 , i n t h e Cedar Creek V a l l e y , C o q u i t l a m Lake Watershed, B.C., 1987. L i n e i n c i r c l e s r e p r e s e n t mean d i r e c t i o n . Numbers i n c i r c l e s r e p r e s e n t s t h e r - v a l u e s . Those marked w i t h a s t a r a r e s i g n i f i c a n t (p < 0.05). The number and a s s o c i a t e d l i n e o u t s i d e t h e c i r c l e s are t h e s c a l e s f o r t h e f r e q u e n c y b a r s . 116 E x p e r i m e n t 2 B e e t l e r e c a p t u r e i n c r e a s e d from an o v e r a l l a verage o f 6.9% i n t h e f i r s t e x p e r i m e n t t o 10.1% i n t h e second e x p e r i m e n t ( T a b l e 9 ) , and must be a t t r i b u t e d t o t h e a d d i t i o n o f t h e f o u r t r a p s a t 5 m. B e e t l e r e c a p t u r e w i t h r e s p e c t t o t i m e was f a i r l y c o n s i s t e n t f o r a l l 3 r e l e a s e s . A p p r o x i m a t e l y 1/2 o f t h e b e e t l e s were r e c a p t u r e d d u r i n g t h e f i r s t h a l f o f t h e f i r s t day (time i n t e r v a l 1 ) , w i t h t h e r e m a i n i n g c a t c h b e i n g s p l i t between t h e second h a l f o f day 1 ( t i m e i n t e r v a l 2) and day 2 ( T a b l e 1 0 ) . The o v e r a l l number o f b e e t l e s r e c a p t u r e d d i d n o t d i f f e r s i g n i f i c a n t l y between 5, 25, and 50 m, w i t h i n t e r t r a p s p a c i n g s o f 8, 20, and 2 0 m, r e s p e c t i v e l y ( T a b l e 11). Only 1 b e e t l e was caught i n each o f t h e t r a p l i n e s W and E-B. Both c a t c h e s o c c u r r e d d u r i n g t h e day o f r e l e a s e . A s i m i l a r w i nd p a t t e r n was o b s e r v e d a t t h e r e l e a s e s i t e when compared t o t h e p r e v i o u s e x p e r i m e n t . However, o n l y d u r i n g t h e second r e l e a s e was wind s i g n i f i c a n t l y d i r e c t e d (from t h e n o r t h ) ( F i g . 2 0 ) . B e e t l e f l i g h t d i r e c t i o n was s i g n i f i c a n t l y d i r e c t e d f o r a l l t h r e e r e l e a s e s a t 5 m, t h e t h i r d r e l e a s e a t 25 m, and t h e second and t h i r d r e l e a s e s a t 50 m. A one-sample t e s t f o r comparing 0^ t o Fw was made f o r t h e second r e l e a s e d a t a s i n c e wind was s i g n i f i c a n t l y d i r e c t e d f o r t h i s r e l e a s e o n l y . I n t h i s c a s e , 0 b was not s i g n i f i c a n t l y d i f f e r e n t from 0^ a t 5 and 50 m ( T a b l e 1 2 ) . Thus, upwind f l i g h t i s i n d i c a t e d by t h e s e d a t a . 117 N e v e r t h e l e s s , i t s h o u l d be n o t i c e d t h a t t h e f r e q u e n c y o f w i nd d i r e c t i o n a g a i n s u r r ounded t h e mean w i t h o u t showing many o b s e r v a t i o n s a t t h e mean ( F i g . 2 0 ) . I n g e n e r a l i t can be s a i d t h a t a t r e l e a s e s i t e A, w ind d i r e c t i o n was e x t r e m e l y v a r i a b l e , and w h i l e t h e 10 minute i n t e r v a l s may not have been f i n e enough t o d e t e c t i t i n a l l c a s e s , w ind s h i f t s were c o n s t a n t l y o c c u r r i n g . Under such c o n d i t i o n s , b e e t l e f l i g h t d i r e c t i o n would be e x p e c t e d t o be as v a r i a b l e as t h e wind i t s e l f , as t h e s e d a t a seem t o i n d i c a t e . E x p e r i m e n t 3 B e e t l e r e c a p t u r e remained low a t r e l e a s e s i t e B and may be p a r t i a l l y a t t r i b u t e d t o t h e reduced number o f t r a p s p l a c e d n e a r t h e r e l e a s e s i t e . P e r c e n t r e c a p t u r e a veraged 8 .5 t h r o u g h f o u r r e l e a s e s , y e t t h i s was l a r g e l y b o l s t e r e d by r e l e a s e 2 d a t a ( T a b l e 9 ) . The second r e l e a s e day s t a n d s out because 90 min f o l l o w i n g b e e t l e r e l e a s e , a major r a i n s t o r m o c c u r r e d . Wind speed measurements showed an average o f o n l y 0.7 km/h p r i o r t o t h e s t o r m ; t h e l o w e s t measured o v e r t h e summer. D e s p i t e o n l y a 90 minute f l i g h t t i m e , t h e number o f b e e t l e s r e c a p t u r e d d u r i n g t h i s r e l e a s e f a r s u r p a s s e d t h e t o t a l s from a l l t h e o t h e r r e l e a s e s . V i t e e t a l . (1964) found t h a t Dendroctonus f r o n t a l i s Zimmerman a c t i v i t y was u n u s u a l l y h i g h on c l o u d y days p r i o r t o t h e o n s e t o f a r a i n s t o r m . Edson (1978) has s i m i l a r r e s u l t s w i t h t h e mountain p i n e b e e t l e , Dendroctonus ponderosae Hopk. These 118 a u t h o r s s u g g e s t e d t h a t c o n d i t i o n s w i t h i n t h e canopy would be more s t a b l e , t r a p p i n g t h e pheromones i n t h e f o r e s t . T h i s e x p l a n a t i o n may n o t be s u f f i c i e n t , however. Of t h e t o t a l b e e t l e s caught i n t h e t r a p l i n e s p l a c e d i n t h e open s e t t i n g s d u r i n g t h e f o u r r e l e a s e s ( F i g . 21A), b o t h o f t h e b e e t l e s c aught i n t r a p l i n e W, as w e l l as 80% o f t h e b e e t l e s caught on t h e e a s t e r n t r a p l i n e s (E-A, E-B, and E-C) were r e c o v e r e d a f t e r t h e f i r s t 90 minute p e r i o d . U n f o r t u n a t e l y t h e weather s t a t i o n m a l f u n c t i o n e d a t t h i s t i m e . Y e t s i n c e w i n d p a r a m e t e r s a t s i t e B c o r r e l a t e w e l l w i t h open s i t e w i n d p a r a m e t e r s ( T a b l e 8 ) , i t can be assumed t h a t w ind speed was g r e a t l y r e d u c e d i n t h e open. A t low w ind speeds, T. l i n e a t u m p r e f e r t o f l y (Rudinsky and Daterman 19 64) and r e s p o n d b e s t t o s e m i o c h e m i c a l s (Chapter 2 ) . The c o m b i n a t i o n o f i n c r e a s e d f l i g h t c a p a b i l i t i e s a l o n g w i t h a more a p p a r e n t o r c o n s t a n t pheromone plume may e x p l a i n t h e i n c r e a s e i n r e s p o n s e . O t h e r e x p l a n a t i o n s f o r t h e i n c r e a s e d r e s p o n s e may i n c l u d e changes i n a t m o s p h e r i c p r e s s u r e (Haufe 1954) o r a t m o s p h e r i c e l e c t r i c i t y (Edwards 1960), t y p i c a l o f m e t e o r o l o g i c a l c o n d i t i o n s p r i o r t o a r a i n s t o r m . However t h e s e f a c t o r s were n e i t h e r measured n o r t e s t e d . B e e t l e r e c a p t u r e d u r i n g t i m e i n t e r v a l 1 ranged from 45 t o 81% o f a l l b e e t l e s r e c a p t u r e d ( T a b l e 1 0 ) . I n t e r v a l 2 c a t c h e s were r e l a t i v e l y low, r a n g i n g from 7 t o 10%, w h i l e day 2 c a t c h e s i n c r e a s e d i n i m p o r t a n c e r e l a t i v e t o t h e two p r e v i o u s e x p e r i m e n t s , w h i c h v a r i e d from 15 t o 48%. The mean % o f b e e t l e s r e c a p t u r e d a t 25 m was s i g n i f i c a n t l y h i g h e r 119 t h a n i n e x p e r i m e n t s 1 and 2 (T a b l e 1 1 ) . However, h i g h c a t c h e s f o r t h e second r e l e a s e were p r o b a b l y most r e s p o n s i b l e f o r t h i s . N e v e r t h e l e s s , i n c r e a s e d c a t c h e s a t 25 m may have been p a r t l y due t o t h e r e d u c t i o n i n t h e amount o f u n d e r s t o r y v e g e t a t i o n , as w e l l as t h e absence o f t h e t r a p s a t 5 m used i n ex p e r i m e n t 2. D i r e c t i o n a l d a t a a t r e l e a s e s i t e B showed t h a t w i n d d i r e c t i o n was c o n s i s t e n t l y s o u t h w e s t e r l y and s i g n i f i c a n t l y d i r e c t e d f o r r e l e a s e s 1, 3, and 4 ( F i g . 2 0 ) . F o r t h e s e 3 r e l e a s e s , b e e t l e f l i g h t was s i g n i f i c a n t l y d i r e c t e d . One-sample comparisons showed t h a t 0 b d i d n o t d i f f e r from 0 W d u r i n g r e l e a s e s 1 and 3, w h i l e d u r i n g r e l e a s e 4, t h e d i f f e r e n c e was o n l y by 2° (T a b l e 1 2 ) . I t i s a p p a r e n t t h a t when wind i s s t r o n g l y d i r e c t e d as i t was a t s i t e B, t h e n b e e t l e f l i g h t d i r e c t i o n i s d e f i n i t e l y upwind a t 25 m. These r e s u l t s s u p p o r t t h o s e from C h a p t e r 5, where upwind f l i g h t a t 25 m may have been i n f l u e n c e d by t h e 5 m t r a p s . D u r i n g t h i s e x p e r i m e n t , T. l i n e a t u m showed i t s a b i l i t y t o f l y l o n g d i s t a n c e s , even when s u r r o u n d e d by t r a p s a t c l o s e range. On t h e days o f r e l e a s e , 2 b e e t l e s were r e c a p t u r e d 1 km d o w n - v a l l e y ( l i n e W) a c r o s s a l a r g e f o r e s t s e t t i n g ( F i g . 21A). A s i d e from t h e 25 m t r a p s , t h e h i g h e s t number o f b e e t l e s (52) were r e c a p t u r e d a t t h e n e x t c l o s e s t t r a p l i n e , E-A. Only 6 were caught i n l i n e E-B t r a p s , l o c a t e d i n t h e m i d d l e o f t h e 700 m l o n g r e g e n e r a t i o n o p e n i n g . Ten b e e t l e s were caught 325 m f u r t h e r u p - v a l l e y i n l i n e E-C t r a p s . T h i s p a t t e r n h e l d f o r day 2 c a p t u r e s as 120 A TOTAL -2 52 6 10 MALE - 1 40 2 6 FEMALE =1 12 4 4 WEST E-A E-B E-C T R A P L I N E S ® Beetle release site with 8 traps placed in a circular trapline 25 m from the point Number caught represents beetles caught in these traps. 21. Recapture of marked T. Lineatum i n pheromone traps, for experiment 3, i n the Cedar Creek Valley, Coquitlam Lake Uatershed, B.C., 1987. Traps were placed c i r c u l a r l y around the release point and i n traplines 50, 375, and 700 m east and 1000 m west of i t . These catches are based on 4 releases where a to t a l of 11 196 beetles flew. A) The t o t a l number of beetles recaptured on the day of release; B) The number of beetles recaptured during the next c o l l e c t i o n date (2 - 10 days l a t e r ) . 121 w e l l . Four b e e t l e s were caught i n l i n e W ( F i g . 21B). L i n e E-A t r a p s a g a i n r e c a p t u r e d t h e most b e e t l e s w i t h a s h a r p d e c r e a s e i n l i n e E-B, and t h e n an i n c r e a s e i n l i n e E-C. Three p o i n t s s h o u l d be n o t e d : 1) T. l i n e a t u m f l e w down-v a l l e y 1 km, d e s p i t e s t r o n g u p - v a l l e y winds t h r o u g h o u t t h e day. A l t h o u g h l o n g d i s t a n c e f l i g h t was e x p e c t e d t o be s t r i c t l y downwind (Chapter 5 ) , i t now appears t h a t b e e t l e s a r e c a p a b l e f l y i n g a g a i n s t t h e wind a t d i s t a n c e s as g r e a t as 1 km. ; 2) a t d i s t a n c e s > 375 m, as many o r more T. l i n e a t u m were caught i n day 2 p e r i o d s t h a n d u r i n g t h e day o f r e l e a s e ; and 3) fewer b e e t l e s were caught i n t h e m i d d l e o f t h e r e g e n e r a t i o n o p e n i n g t h a n a t t h e edge 325 m f u r t h e r away ( l i n e E-C t r a p s ) . The l a t t e r p o i n t may be e x p l a i n e d by t h e e f f e c t o f a l a r g e o pening on wind d i r e c t i o n . Kimmins (1987) e x p l a i n e d t h a t l a m i n a r a i r f l o w o v e r a c o o l f o r e s t canopy w i l l become t u r b u l e n t upon c o n t a c t w i t h a l a r g e a r e a where t h e s u r f a c e t e m p e r a t u r e s a r e warmer and more v a r i a b l e . T h i s t u r b u l e n c e causes a i r t o move upwards i n t h e m i d d l e o f t h e op e n i n g . A i r movement w i l l t h e n become i n c r e a s i n g l y l a m i n a r as i t approaches t h e n e x t f o r e s t canopy. T h i s p a t t e r n o f a i r movement was a l s o o b s e r v e d a t our work s i t e when numerous l i g h t w e i g h t seeds o f 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., were o b s e r v e d i n t h e a i r , and t h e i r d i s p e r s a l by t h e wind was m o n i t o r e d o v e r t h e open s e t t i n g i n whi c h t r a p l i n e s E-A, E-B, and E-C were d e p l o y e d . I n t h e m i d d l e o f t h i s s e t t i n g , seeds 10 - 30 m above t h e ground were moving i n a l l d i r e c t i o n s , whereas seeds o b s e r v e d 122 f u r t h e r a l o n g t h e v a l l e y , near t h e n e x t f o r e s t canopy (near l i n e E-C) were b e i n g t a k e n u p - v a l l e y i n t h e s t r o n g winds. E x p e r i m e n t 4 I n t h i s e x p e r i m e n t , t h e number o f b e e t l e s r e c a p t u r e d d e c r e a s e d c o n s i d e r a b l y w i t h t h e removal o f t h e 25 m t r a p s from e x p e r i m e n t 3 ( T a b l e 9 ) . However, more b e e t l e s were r e c a p t u r e d i n t h e 50 m t r a p s (E-A) i n e x p e r i m e n t 4 (79 b e e t l e s / r e l e a s e ) ( F i g . 22) t h a n i n e x p e r i m e n t 3 (20 b e e t l e s / r e l e a s e ) ( F i g . 2 1 ) . T h i s s u p p o r t s d a t a from C h a p t e r 5, where t h e f r e q u e n c y o f l o n g d i s t a n c e f l i g h t i n c r e a s e d when a t t r a c t a n t s p l a c e d c l o s e t o t h e r e l e a s e s i t e were removed. S i x t y p e r c e n t o f t h e b e e t l e s r e c a p t u r e d i n t h e 50 m t r a p s were caught d u r i n g t h e day o f r e l e a s e ( F i g . 2 2 ) . A t t h e l o n g e r d i s t a n c e s , a l l b e e t l e s were r e c a p t u r e d d u r i n g t h e day 2 p e r i o d s . A l t h o u g h t h r e e b e e t l e s were r e c a p t u r e d (2 i n l i n e E-D t r a p s and 1 i n l i n e E-F t r a p s ) d u r i n g t h e day o f r e l e a s e , t h e y were from e x p e r i m e n t 3. Day 2 p e r i o d s were 3 and 5 days, r e s p e c t i v e l y , f o r r e l e a s e s 1 and 2. O v e r a l l , 1 b e e t l e was caught 1 km d o w n - v a l l e y (W), 8 were caught 700 m u p - v a l l e y (E-C), 9 were caught 1.1 km u p - v a l l e y (E-D), 1 was caught 1.9 km u p - v a l l e y ( E - E ) , and 1 was caught a c r o s s t h e v a l l e y ( E - F ) , 1.3 km away from t h e s i t e . 123 A T O T A L M A L E F E M A L E = 0 93 0 0 0 0 = 0 73 0 0 0 0 = 0 20 0 0 0 0 T R A P L I N E S ® Beetle Release Site Fig. 22. Recapture of marked T. Lineatum in pheromone traps for experiment 4, i n the Cedar Creek Valley, Coquitlam Lake Watershed, B.C., 1987. Traplines were set up 50, 700, 1100, 1900 m east and 1000 m west of the release point. Additionally, l i n e E-F was placed on the other side of the va l l e y , ca. 1300 m east of the release point. These catches are based on 2 releases where a t o t a l of 6 698 beetles flew. A) The number of beetles recaptured on the day of release; B) The number of beetles recaptured during the next c o l l e c t i o n date ( 3 - 5 days l a t e r ) . 124 E x p e r i m e n t 5 Only 17 b e e t l e s o r 0.4% o f t h e b e e t l e s t h a t f l e w ( T a b l e 9) were r e c a p t u r e d f o l l o w i n g t h e one r e l e a s e . Removal o f t r a p l i n e E-A was a l a r g e r e a s o n f o r t h i s . D u r i n g t h e two r e l e a s e s from e x p e r i m e n t 4, a t o t a l o f 19 b e e t l e s were r e c a p t u r e d i n l i n e s W, E-C, E-D, E-E, and E-F, whereas i n t h e o n l y r e l e a s e i n t h i s e x p e r i m e n t , 17 b e e t l e s were r e c a p t u r e d i n t h o s e same t r a p l i n e s . Thus, a g a i n more b e e t l e s were f l y i n g a l o n g e r d i s t a n c e . On t h e day o f r e l e a s e , 2 b e e t l e s were t r a p p e d on l i n e E-C and 1 made i t t o l i n e E-D. The r e s t o f t h e b e e t l e s , 1 i n l i n e W, 6 i n E-C, 5 i n E-D, 1 i n E-E, and 1 i n E-F, were caught d u r i n g t h e day 2 p e r i o d . P o p u l a t i o n D i s p e r s a l I n e x p e r i m e n t 4, t h e t o t a l number o f b e e t l e s r e c a p t u r e d a t d i s t a n c e s > 700 m u p - v a l l e y was 19, and a t d i s t a n c e s o f > 1 km, t h e number caught was 14. T h i s i s 10.6 and 7.8%, r e s p e c t i v e l y , o f t h e 180 marked b e e t l e s r e c a p t u r e d d u r i n g t h e two r e l e a s e s ( T a b l e 9 ) . I f a p o p u l a t i o n e s t i m a t e c o u l d be made f o r t h e a r e a , t h e number o f b e e t l e s f l y i n g t h o s e d i s t a n c e s c o u l d be e s t i m a t e d . T h e r e f o r e p o p u l a t i o n e s t i m a t e s f o r t h e f o r e s t e d p a t c h c o n t a i n i n g r e l e a s e s i t e s A and B ( F i g . 19) were made based on t h e m a r k - r e c a p t u r e d a t a from e x p e r i m e n t s 1 and 2. These e x p e r i m e n t s were used because t h e y were c l o s e s t t o t h e peak emergence t i m e o f T. 125 l i n e a t u m . and t r a p arrangement was d e s i g n e d t o c a p t u r e as many b e e t l e s w i t h i n as c l o s e an a r e a as p o s s i b l e . Data from t r a p l i n e s W and E-B i n experiment 2 were n o t used i n making t h e e s t i m a t e s , because t h e y extended f a y beyond t h e f o r e s t e d r e l e a s e a r e a . The L i n c o l n i n d e x f o r p o p u l a t i o n e s t i m a t i o n was used: N = a n / r where N = p o p u l a t i o n e s t i m a t e a = # o f marked b e e t l e s t h a t f l e w r = # o f marked b e e t l e s r e c a p t u r e d n = t o t a l # o f b e e t l e s c a p t u r e d The v a r i a n c e was d e t e r m i n e d as s 2 = a 2 n ( n - r ) / r 3 and 75% c o n f i d e n c e l i m i t s (C.L.) as N + 2s (Southwood 1978). I t i s assumed ( a l t h o u g h n o t t r u l y known) t h a t t h o s e b e e t l e s r e c a p t u r e d a t r e l e a s e s i t e A come from an a r e a e q u i v a l e n t t o t h e average f l i g h t range o f t h e b e e t l e s up t o t h e p o i n t o f a t t a c k . T o t a l p o p u l a t i o n e s t i m a t e s from such an a r e a f o r t h e 3 r e l e a s e s i n exp e r i m e n t 1 a r e 118 000 + 12 000 b e e t l e s , and f o r t h e 3 r e l e a s e s i n t h e second e x p e r i m e n t , 28 000 + 2 000 b e e t l e s . T h i s i s a t o t a l o f c a . 146 000 b e e t l e s . I t i s a c o n s e r v a t i v e p o p u l a t i o n e s t i m a t e , s i n c e c a l c u l a t i o n s were made on b e e t l e s t h a t had f l o w n o v e r a 1 month p e r i o d t h a t p r o b a b l y f o l l o w e d t h e e a r l i e r peak emergence days o f A p r i l and May. Based on t h e assu m p t i o n t h a t uncaught b e e t l e s behaved s i m i l a r l y t o caught b e e t l e s , w i t h r e s p e c t t o d i s t a n c e , we can e s t i m a t e t h a t 15 400 b e e t l e s can f l y t o an a t t r a c t i v e h o s t 700 m from t h e i r emergence s i t e . I t can a l s o be s a i d t h a t 11 400 b e e t l e s can f l y t o an a t t r a c t i v e 126 h o s t a t l e a s t 1 km away. Such h i g h numbers s h o u l d be a c o n c e r n t o t h o s e i n v o l v e d i n h a r v e s t p l a n n i n g o p e r a t i o n s . I n c o n c l u s i o n , o b s e r v e d m e t e o r o l o g i c a l d i f f e r e n c e s between t h e open s i t e and f o r e s t e d s i t e s A and B o c c u r r e d as e x p e c t e d . F o r e s t c o v e r r e d u c e s s u n l i g h t r e s u l t i n g i n a d e c r e a s e i n t e m p e r a t u r e w i t h an i n c r e a s e i n r e l a t i v e h u m i d i t y . D e s p i t e s t r o n g , c o n s i s t e n t down- t o u p - v a l l e y winds d u r i n g sunny days, a t t h e open s i t e , w i nd d i r e c t i o n a t r e l e a s e s i t e A was q u i t e v a r i a b l e , w h i l e a t s i t e B, w i t h l e s s v e g e t a t i o n and c l o s e r t o t h e v a l l e y i n f l u e n c e ( v i a t h e r o a d ) , wind d i r e c t i o n was s t r o n g l y d i r e c t e d from down- t o u p - v a l l e y . T. l i n e a t u m f l i g h t d i r e c t i o n a t s i t e A was m o s t l y v a r i a b l e as was t h e wind d i r e c t i o n . The dense u n d e r s t o r y may have had a n e g a t i v e e f f e c t on t h e number o f b e e t l e s r e c a p t u r e d (Shore and McLean 1984), e i t h e r d i r e c t l y o r i n d i r e c t l y due t o i t s i n f l u e n c e on t h e wind w h i c h i n t u r n i n f l u e n c e s t h e pheromone plume. A n o t h e r f a c t o r t h a t may have had an i n f l u e n c e on b e e t l e r e c a p t u r e was t h e p r e s e n c e o f s u s c e p t i b l e windthrown m a t e r i a l a t t h e e a s t e r n edges o f t h e r e g e n e r a t i o n s i t e s . No sample was made t o see whether marked b e e t l e s were a t t r a c t e d t o t h e s e s i t e s . E x p e r i m e n t 2 d a t a showed t h a t few b e e t l e s , emerging from f o r e s t s f a r away from any o p e n i n g s , would t r a v e l l o n g d i s t a n c e s t o a t t r a c t i v e m a t e r i a l . However t h i s s h o u l d be t e s t e d f u r t h e r w i t h o u t a t t r a c t a n t s p r e s e n t n e a r t h e r e l e a s e s i t e . 127 A t r e l e a s e s i t e B, where wind d i r e c t i o n was g e n e r a l l y u n i d i r e c t i o n a l , b e e t l e r e c a p t u r e was upwind a t a d i s t a n c e o f 25 m. T. l i n e a t u m showed i t s a b i l i t y t o f l y r e l a t i v e l y l o n g d i s t a n c e s i n t h i s e x p e r i m e n t w i t h most b e e t l e s caught i n s h o r t - d i s t a n c e t r a p s . More b e e t l e s were c a p t u r e d a t a f a r t h e r d i s t a n c e , a t t h e edge o f t h e o p e n i n g t h a n a t a c l o s e r d i s t a n c e i n t h e m i d d l e o f i t , i n d i c a t i n g t h e s t r o n g e f f e c t o f wind t u r b u l e n c e i n an open s e t t i n g on b e e t l e r e s p o n s e t o a t t r a c t i v e m a t e r i a l . A s h i f t i n response t i m e was n o t e d i n t h i s s t u d y . I n e x p e r i m e n t s 1 - 3 , t h e m a j o r i t y o f b e e t l e s c a p t u r e d a t 50 m o r l e s s o c c u r r e d w i t h i n t h e f i r s t 2 h o u r s o f r e l e a s e , y e t a t d i s t a n c e s r a n g i n g from 350 - 700 m, as many b e e t l e s were caught d u r i n g t h e days f o l l o w i n g t h e r e l e a s e as were caught d u r i n g t h e day o f r e l e a s e . I n e x p e r i m e n t s 4 and 5 most r e c a p t u r e d b e e t l e s a t d i s t a n c e s > 1 km from t h e r e l e a s e s i t e were caught a f t e r Day 1. T h i s c o u l d have been due t o t h e i n c r e a s e d t r a v e l d i s t a n c e , t h e weakened s t a t e o f t h e s t o r e d i n s e c t s , o r a c o m b i n a t i o n o f b o t h . Only 21 o f t h e b e e t l e s r e l e a s e d i n e x p e r i m e n t s 4 and 5 were c a p t u r e d u p - v a l l e y a t d i s t a n c e s > 1 km, w i t h 3 b e e t l e s r e c a p t u r e d as f a r as 1.9 km. However, based on t h e p o p u l a t i o n d i s p e r s a l e s t i m a t e s above, a s u b s t a n t i a l number o f b e e t l e s a r e c a p a b l e o f f l y i n g a t l e a s t 1 km from s i t e s w i t h l a r g e numbers o f o v e r w i n t e r i n g b e e t l e s . F u r n i s s and F u r n i s s (1972) have r e p o r t e d f i n d i n g T. l i n e a t u m f a r from h o s t m a t e r i a l , n e a r mountain t o p s a l o n g t h e west c o a s t o f 128 t h e U n i t e d S t a t e s . Dyer (1961) r e p o r t e d f i n d i n g a marked b e e t l e 4 km from i t s r e l e a s e s i t e , one y e a r l a t e r . T h i s v a l l e y s t u d y r e p r e s e n t s more s u b s t a n t i a l e v i d e n c e o f t h e a b i l i t y o f T. l i n e a t u m t o f l y o r be t a k e n by t h e wind c o n s i d e r a b l e d i s t a n c e s . Trap placement d u r i n g t h i s s t u d y was c o n c e n t r a t e d up-v a l l e y from t h e r e l e a s e s i t e s i n an u n b a l a n c e d d e s i g n , based on e v i d e n c e from C h a p t e r 5. B e e t l e r e c a p t u r e 1 km down-v a l l e y from t h e r e l e a s e s i t e , a g a i n s t t h e wind, may w a r r a n t a r e - e v a l u a t i o n o f t h e h y p o t h e s i s t h a t l o n g d i s t a n c e f l i g h t by T. l i n e a t u m i s p r i m a r i l y downwind. T h i s would r e q u i r e a more b a l a n c e d d e s i g n i n f u t u r e s t u d i e s t o b e t t e r c h a r a c t e r i z e f l i g h t p a t t e r n s o f T. l i n e a t u m i n t h e absence o f a t t r a c t a n t s c l o s e t o t h e b e e t l e r e l e a s e s i t e s . 129 CHAPTER 7 : POPULATION MOVEMENT PATTERNS OF TRYPODENDRON LINEATUM WITH RESPECT TO WIND AND VEGETATION WITHIN A VALLEY SETTING. 7 . l I n t r o d u c t i o n B e e t l e c a t c h d a t a from t h e f i r s t m a r k - r e c a p t u r e s t u d y , h e l d i n t h e U n i v e r s i t y Endowment Lands (Chap. 5 ) , s u g g e s t e d t h a t l o n g d i s t a n c e b e e t l e f l i g h t would be c o n s i s t e n t l y downwind from a r e l e a s e p o i n t o r emergence s i t e . T h i s l e d t o p l o t d e s i g n s f o r r e c a p t u r i n g b e e t l e s t h a t f a v o r e d downwind f l i g h t ( u p - v a l l e y i n most cases) by T. l i n e a t u m i n t h e f i r s t v a l l e y s t u d y (Chap. 6 ) . The one t r a p l i n e t h a t was p l a c e d 1 km upwind (down-valley) from t h e r e l e a s e p o i n t , r e c a p t u r e d 8 b e e t l e s , w h i c h when compared t o t h e numbers o f b e e t l e s r e c a p t u r e d downwind, was u n e x p e c t e d l y h i g h . Thus, i n t h i s c h a p t e r a second v a l l e y s t u d y i s r e p o r t e d i n w h i c h t h e o b j e c t i v e s were: 1) compare T. l i n e a t u m r e c a p t u r e p a t t e r n t o wind d i r e c t i o n w i t h o u t any t r a p p i n g b i a s and w i t h o u t a t t r a c t a n t s n e a r t h e r e l e a s e s i t e s t h a t c o u l d p o t e n t i a l l y i n f l u e n c e b e e t l e f l i g h t d i r e c t i o n ; 2) as a f o l l o w - u p t o t h e f i r s t v a l l e y s t u d y , where T. l i n e a t u m r e c a p t u r e t e n d e d t o be l o w e s t i n t h e c e n t e r o f open s e t t i n g s , b e e t l e r e c a p t u r e p a t t e r n s between open and f o r e s t s e t t i n g s were compared; and 3) an attempt was made t o t e s t whether T. l i n e a t u m t e n d t o d i s p e r s e a l o n g c e r t a i n r o u t e s i n t h e f o r e s t , i e . , t h e operi a r e a , o r a l o n g f o r e s t edges such as r i g h t s - o f - w a y o r r o a d s . 130 7.2 M a t e r i a l s and Methods S i t e P a rameters Two m a r k - r e c a p t u r e e x p e r i m e n t s were c a r r i e d o u t i n t h e Cedar Creek V a l l e y o f t h e C o q u i t l a m Lake Watershed, i n t h e l o w e r c o a s t a l m a i n l a n d o f B r i t i s h C olumbia, from June t o J u l y , 1988. A d e t a i l e d d e s c r i p t i o n o f t h e w o r k i n g s i t e i s g i v e n i n C h a p t e r 6. F o r b o t h e x p e r i m e n t s , b e e t l e s were r e l e a s e d s i m u l t a n e o u s l y from two s i t e s on t h e n o r t h s i d e o f Cedar Creek ( F i g . 2 3 ) . S i t e A was l o c a t e d on t h e d o w n - v a l l e y edge o f a f o r e s t s e t t i n g , and s i t e B was l o c a t e d 650 m u p - v a l l e y a t t h e o p p o s i n g edge o f t h e same f o r e s t s e t t i n g . T h i s was t h o u g h t t o be s u p e r i o r t o t h e one r e l e a s e s i t e used i n t h e f i r s t v a l l e y s t u d y (Chapter 6 ) , because i t a l l o w e d f o r s e v e r a l h y p o t h e s e s o f T. l i n e a t u m d i s p e r s a l t o be t e s t e d w i t h o u t b i a s i n g t r a p placement up- o r d o w n - v a l l e y . A l s o i t a l l o w e d a g a i n f o r t h e r e l e a s e o f b e e t l e s from t h e f o r e s t edges, w h i c h i s where most T. l i n e a t u m a r e known t o o v e r w i n t e r (Dyer and K i n g h o r n 1961). B e e t l e s r e l e a s e d from s i t e A a r e h e r e a f t e r d e s i g n a t e d as p o p u l a t i o n A, w h i l e b e e t l e s r e l e a s e d from s i t e B a r e d e s i g n a t e d as p o p u l a t i o n B. Groups o f f o u r s e m i o c h e m i c a l - b a i t e d f u n n e l t r a p s were used as s o u r c e s o f a t t r a c t i o n f o r r e c a p t u r i n g marked b e e t l e s . F u n n e l t r a p s w i t h i n each group were s e t up a t t h e c o r n e r s o f a square p l o t , 10 x 10 m i n a r e a , i n an e f f o r t t o 131 F i g . 23 Contour map o f t h e Cedar Creek V a l l e y i n t h e C o q u i t l a m Lake Watershed, w i t h t h e d e s i g n s o f t h e f i r s t (A) and second (B) m a r k - r e c a p t u r e e x p e r i m e n t s , c a r r i e d o ut i n 1988. D o t t e d p o l y g o n s r e p r e s e n t r e g e n e r a t i o n o penings s u r r o u n d e d by o l d - g r o w t h f o r e s t . The u p - v a l l e y a z i m u t h = 48 - 68°. 1321 produce a l a r g e r a c t i v e space o f s e m i o c h e m i c a l plume t h a n p r o d u c e d by one t r a p , t o i n c r e a s e t h e l e v e l o f a t t r a c t i o n o f T. l i n e a t u m t o t h e d e s i g n a t e d s i t e s i n w h i c h t r a p s were d e p l o y e d ( L i n d g r e n e t a l . 1983). A l l f o u r t r a p s w i t h i n each group were b a i t e d w i t h l i n e a t i n , t h e a g g r e g a t i o n pheromone o f T. l i n e a t u m . Two Hereon® c o n t r o l l e d r e l e a s e d i s p e n s e r s w i t h a t o t a l r e l e a s e r a t e o f 340 ug/24 were p l a c e d i n each t r a p . A l t h o u g h e t h a n o l , t h e h o s t a t t r a c t a n t f o r T. l i n e a t u m , i s n o t c o n s i d e r e d e f f e c t i v e i n f u n n e l t r a p s (Chapter 2, Salom and McLean 1988), i t was i n c l u d e d i n two o f t h e t r a p s f o r each group. T h i s was due i n l a r g e p a r t t o a new, and p o t e n t i a l l y b e t t e r d i s p e n s e r , i n t h e form o f a 2.5 by 41 cm p o l y - v i n y l c h l o r i d e bag, f i l l e d w i t h 15 ml o f 95% e t h a n o l . F o r each t r a p b a i t e d w i t h e t h a n o l , one d i s p e n s e r was used, r e s u l t i n g i n a r e l e a s e r a t e o f 30 mg/24 h o v e r a p e r i o d o f 4 5 days'2-. E x p e r i m e n t 1 T h i s e x p e r i m e n t , which was r e p l i c a t e d f o u r t i m e s , was d e s i g n e d t o e v a l u a t e l o n g d i s t a n c e T. l i n e a t u m f l i g h t w i t h i n t h e v a l l e y i n t h e absence o f a t t r a c t a n t s n e a r t h e b e e t l e r e l e a s e s i t e s . Groups o f f u n n e l t r a p s were s e t up i n a s y m m e t r i c a l d e s i g n up, down, and a c r o s s t h e v a l l e y , w i t h r e s p e c t t o r e l e a s e s i t e s A and B ( F i g . 23A). No f u n n e l t r a p group was s e t any c l o s e r t o a r e l e a s e s i t e t h a n 200 m. The f i r s t h y p o t h e s i s t e s t e d was t h a t i n t h e absence o f a t t r a c t a n t s n e a r t h e r e l e a s e s i t e , most o f t h e r e l e a s e d 133 b e e t l e s would be r e c a p t u r e d downwind. The p l o t d e s i g n a l l o w e d f o r two ways t o t e s t t h i s ( F i g . 23A). F i r s t , t h e number o f p o p u l a t i o n A b e e t l e s r e c a p t u r e d i n t h e seven t r a p groups (7 - 1 3 ) , s e t u p - v a l l e y from s i t e A, c o u l d be compared t o t h e number o f p o p u l a t i o n B b e e t l e s r e c a p t u r e d i n an e q u a l number o f t r a p groups (1 - 7 ) , s e t d o w n - v a l l e y from s i t e B. S e c o n d l y , t h e number o f p o p u l a t i o n A b e e t l e s r e c a p t u r e d i n t h e f i v e t r a p groups (1 - 5 ) , s e t d o w n - v a l l e y from s i t e A, w i t h t h e number o f p o p u l a t i o n B b e e t l e s r e c a p t u r e d i n an e q u a l number o f t r a p groups (9 - 1 3 ) , s e t u p - v a l l e y from s i t e B. The s y m m e t r i c a l placement o f t r a p groups r e l a t i v e t o s i t e s A and B r e s u l t e d i n a m i r r o r image t h a t made t h e s e comparisons v a l i d . A second h y p o t h e s i s t e s t e d was t h a t b e e t l e r e c a p t u r e numbers would be d i f f e r e n t i n an open s e t t i n g when compared t o a f o r e s t e d one, d e s p i t e t h e u n e q u a l number o f t r a p groups r e p r e s e n t i n g b o t h t r e a t m e n t s . Four groups ( 1 , 2, 12, and 13) were s e t w i t h i n t h e f o r e s t and n i n e groups (3 - 11) were s e t up i n t h r e e s e p a r a t e r e g e n e r a t i o n o p e n i n g s ; one u p - v a l l e y , one down-v a l l e y , and one a c r o s s t h e v a l l e y , from b o t h r e l e a s e s i t e s ( F i g . 23A). Ex p e r i m e n t 2 T h i s e x p e r i m e n t , which was a l s o r e p l i c a t e d f o u r t i m e s , was d e s i g n e d t o t e s t what t y p e o f f l y i n g r o u t e s a r e p r e f e r r e d by d i s p e r s i n g T. l i n e a t u m . One approach was t o compare t h e number o f up- ( p o p u l a t i o n A) o r d o w n - v a l l e y 134 ( p o p u l a t i o n B) b e e t l e s t r a p p e d i n t h e f o r e s t , 30 m above t h e r o a d ( 8 ) , on t h e r o a d ( 7 ) , and i n t h e f o r e s t , 100 m below t h e r o a d (6) ( F i g . 23B). T h i s i s based on t h e h y p o t h e s i s t h a t d i s p e r s i n g b e e t l e s use r i g h t s - o f - w a y r o a d s as " f l y w a y s " , making c u t t i m b e r a l o n g t h e s e p a t h s e s p e c i a l l y s u s c e p t i b l e t o b e e t l e a t t a c k (J.A. McLean, p e r s o n a l communication)! 5-. A second approach examined t h e r o u t e t a k e n by b e e t l e s f l y i n g toward t r a p s f a r t h e s t from t h e r e l e a s e s i t e s ( 1 , 2, 11, and 12). B e e t l e c a t c h e s were compared between open t r a p groups (3 and 10) and f o r e s t t r a p groups (4 and 9 ) , p l a c e d i n between t h e r e l e a s e s i t e s and t h e l o n g d i s t a n c e t r a p groups ( F i g . 23B). As i n t h e f i r s t e x p e r i m e n t , l o n g d i s t a n c e f l i g h t d e t e r m i n a t i o n s , w i t h r e s p e c t t o wind were examined, w i t h o u t t h e p r e s e n c e o f s e m i o c h e m i c a l s near t h e b e e t l e r e l e a s e s i t e s . A g a i n t r a p groups were p l a c e d s y m m e t r i c a l l y up- and d o w n - v a l l e y from r e l e a s e s i t e s A and B, and a l l o w e d f o r two t e s t s . F i r s t , t h e number o f p o p u l a t i o n A b e e t l e s r e c a p t u r e d i n t h e e i g h t t r a p groups (5 - 1 2 ) , s e t u p - v a l l e y from s i t e A, c o u l d be compared t o t h e number o f p o p u l a t i o n B b e e t l e s r e c a p t u r e d i n an e q u a l number o f t r a p groups (1 - 8 ) , s e t d o w n - v a l l e y from s i t e B. S e c o n d l y , t h e number o f p o p u l a t i o n A b e e t l e s r e c a p t u r e d i n t h e f o u r t r a p groups (1 - 4 ) , d o w n - v a l l e y from s i t e A, was compared t o t h e number o f p o p u l a t i o n B b e e t l e s r e c a p t u r e d i n an e q u a l number o f t r a p groups (9 - 1 2 ) , up-v a l l e y from s i t e B. |_5_ Dr. J.A. McLean, P r o f e s s o r , Department o f F o r e s t S c i e n c e s , U n i v e r s i t y o f B r i t i s h Columbia, Vancouver, B.C. 135 Mark-Recapture A d u l t T. l i n e a t u m used i n t h i s s t u d y were c o l l e c t e d d u r i n g t h e i r mass f l i g h t p e r i o d o f A p r i l and May, 1988. B e e t l e s were c a p t u r e d u s i n g l i n e a t i n - and e t h a n o l - b a i t e d f u n n e l t r a p s , p l a c e d a d j a c e n t t o a l a r g e l o g boom s t o r a g e a r e a a t t h e mouth o f t h e n o r t h arm o f t h e F r a s e r R i v e r i n Vancouver, B.C. B e e t l e c o l l e c t i o n and s t o r a g e p r o c e d u r e s were s i m i l a r t o t h o s e f i r s t d i s c u s s e d i n C h a p t e r 2. The p r o c e d u r e s used f o r mar k i n g b e e t l e s w i t h f l u o r e s c e n t powder were d e s c r i b e d i n C h a p t e r 5. T h i s i n c l u d e d a d e s c r i p t i o n o f a b r i e f wind t u n n e l s t u d y showing no v a r i a t i o n i n r e s p o n s e t o s e m i o c h e m i c a l s and no i n c r e a s e i n m o r t a l i t y one week f o l l o w i n g m a r k i n g (Appendix 3 ) . F o r t h e 1988 s t u d y , f o u r c o l o r e d powders were used t h r o u g h o u t t h e s t u d y , w i t h d i f f e r e n t c o l o r s used f o r p o p u l a t i o n A and B f o r each r e l e a s e . The r e l e a s e - r e c a p t u r e p r o c e d u r e s f o r b o t h e x p e r i m e n t s were v e r y s i m i l a r . On f l i g h t days, b e e t l e s were u s u a l l y r e l e a s e d from 1130 and 13 3 0 h PDT, o r when t h e t e m p e r a t u r e r o s e above 15°C, t h e low t e m p e r a t u r e f l i g h t t h r e s h o l d f o r T. l i n e a t u m (Chapman and K i n g h o r n 1958). F o r each day o f r e l e a s e , 8000 b e e t l e s were r e l e a s e d s i m u l t a n e o u s l y from b o t h s i t e s A (4000) and B (4000) . A t each s i t e , t h e b e e t l e s were p l a c e d on a 1 m wooden t r a y , w i t h a s m a l l frame r i d g e 0.2 m from t h e edge, e f f e c t i v e l y g i v i n g t h e b e e t l e s two r i d g e s from w h i c h t o t a k e - o f f . The t r a p s were s e t 0.5 m above t h e 136 ground. An a b s o r b e n t paper, l i n e d on one s i d e w i t h c e l l o p h a n e , was p l a c e d u n d e r neath t h e t r a y t o c a t c h any b e e t l e s t h a t f e l l o f f . S i m i l a r paper was used as a b e d d i n g from t h e i n n e r square o f t h e t r a y , where t h e b e e t l e s were p l a c e d . C o l l e c t i o n s from a l l t r a p s were g e n e r a l l y i n i t i a t e d 4 -5 h f o l l o w i n g b e e t l e r e l e a s e . I n a c o n t i n u i n g e f f o r t t o see how many marked b e e t l e s f l e w f o r more t h a n 1 day w i t h i n t h e v a l l e y , c o l l e c t i o n s were a g a i n made d u r i n g t h e morning p r i o r t o t h e n e x t r e l e a s e . The b e e t l e s caught i n t h e t r a p s were b r o u g h t back t o t h e l a b f o r c o u n t i n g , and an u l t r a v i o l e t lamp was used t o i d e n t i f y marked specimens. The number o f marked males and fem a l e s were t a l l i e d , as w e l l as t h e number o f unmarked b e e t l e s c o l l e c t e d . The number o f r e l e a s e d b e e t l e s n o t f l y i n g was a l s o r e c o r d e d . Weather Parameters M e t e o r o l o g i c a l measurements were made a t t h e r e l e a s e s i t e s as w e l l as i n an a d j a c e n t o p e n i n g . A t t h e open s i t e ( F i g . 2 3 ) , a p o r t a b l e weather s t a t i o n (W.S.) was s e t up t o r e c o r d a i r t e m p e r a t u r e , r e l a t i v e h u m i d i t y , w i n d d i r e c t i o n , and wind speed c o n t i n u o u s l y t h r o u g h o u t t h e summer. D e t a i l s o f t h e weather equipment were d e s c r i b e d i n C h a p t e r 6. I n an e f f o r t t o i n c r e a s e t h e p r e c i s i o n o f t h e s e d a t a , 5 min measurement i n t e r v a l s were used, compared w i t h t h e 10 min 137 i n t e r v a l s used i n t h e f i r s t two m a r k - r e c a p t u r e s t u d i e s ( C h a p t e r s 5 and 6 ) . A t each r e l e a s e s i t e , measurements were c a r r i e d o u t o n l y w h i l e t h e e x p e r i m e n t s were b e i n g a c t u a l l y r u n . Temperature measurements were made e v e r y 5 min w i t h a mercury thermometer and r e l a t i v e h u m i d i t y was measured h o u r l y w i t h a s l i n g p s ychrometer. Wind d i r e c t i o n and speed d a t a were c o l l e c t e d a t 5 min i n t e r v a l s , w i t h t h e use o f a bee-smoker p l a c e d o v e r a c i r c u l a r g r i d 1 m i n r a d i u s and br o k e n up i n t o 16 a n g u l a r i n t e r v a l s o f 22.5° (360° c i r c l e ) . The s p o u t o f t h e smoker was o r i e n t e d v e r t i c a l l y , 0.5 m above t h e f o r e s t f l o o r . Wind speed was measured by t i m i n g t h e movement o f a p u f f o f smoke f o r a d i s t a n c e o f 1 m. The c i r c u l a r g r i d was p l a c e d 10 m n o r t h o f t h e r e l e a s e t r a y a t b o t h s i t e s A and B. Data A n a l y s i s The wind measurements from a l l t h r e e s i t e s a l l o w e d f o r t h e p r e s e n t a t i o n o f wind d i r e c t i o n i n t h e form o f c i r c u l a r h i s t o g r a m s . The s t a t i s t i c a l p r o c e d u r e s used a r e d e s c r i b e d by B a t s c h e l e t (1981), and a r e summarized i n C h a p t e r 5. F o r t h e wind d a t a i n t h i s s t u d y , mean wind d i r e c t i o n s (0W) were d e t e r m i n e d f o r t h e f i r s t h a l f , second h a l f , and o v e r a l l measurement p e r i o d o f a r e l e a s e day, f o r a l l e i g h t r e l e a s e days. N o n - p a r a m e t r i c s t a t i s t i c s were r e q u i r e d f o r a n a l y z i n g c omparisons between t h e number o f marked T. l i n e a t u m 138 r e c a p t u r e d i n t r a p groups, based on t h e h y p o t h e s e s t e s t e d . Comparisons between up- and d o w n - v a l l e y movement o f p o p u l a t i o n s A and B were made f o r b o t h e x p e r i m e n t s o n l y when wind d i r e c t i o n t h r o u g h o u t t h e measurement p e r i o d o f a r e l e a s e day was r e l a t i v e l y s i m i l a r between t h e weather s t a t i o n and s i t e s A and B. S i m i l a r i t i e s i n wind d i r e c t i o n t e n d e d t o o c c u r when wind blew from down- t o u p - v a l l e y . On t h e s e o c c a s i o n s , t h e c r i t e r i a used t o a l l o w f o r b e e t l e f l i g h t d i r e c t i o n comparisons r e q u i r e d t h a t wind d i r e c t i o n was + 45° t h e v a l l e y a z i m u t h from t h e s i t e measured. The u p - v a l l e y a z i m u t h f o r t h e weather s t a t i o n and s i t e A was c a . 48°, and f o r s i t e B c a . 68°. The number o f p o p u l a t i o n A b e e t l e s r e c a p t u r e d u p - v a l l e y were compared t o t h e number o f p o p u l a t i o n B b e e t l e s r e c a p t u r e d d o w n - v a l l e y . A l s o c o m p a r i s o n s were made f o r each p o p u l a t i o n i n t h e o p p o s i t e d i r e c t i o n s , r e s p e c t i v e l y , w i t h t h e d a t a a n a l y z e d u s i n g t h e Mann-Whitney U t e s t (Zar 1984). The same s t a t i s t i c a l t e s t was used f o r comparing t h e number o f marked b e e t l e s r e c a p t u r e d i n t h e f o r e s t e d and open s e t t i n g s . A c o m p a r i s o n among t h r e e p o s s i b l e r o u t e s t a k e n by T. l i n e a t u m up- and d o w n - v a l l e y i n a f o r e s t f o r e x p e r i m e n t 2 was a n a l y z e d u s i n g t h e K r u s k a l - W a l l i s , a n a l y s i s o f v a r i a n c e by r a n k t e s t (Zar 1984). F o r l o n g e r d i s t a n c e f l i g h t , c o m parisons between an open and a f o r e s t e d r o u t e were made u s i n g t h e Mann-Whitney U t e s t . 139 7.3 R e s u l t s and D i s c u s s i o n Weather S i m u l t a n e o u s weather measurements a t t h r e e d i f f e r e n t s i t e s i n t h e Cedar Creek V a l l e y , p r o v i d e d an o p p o r t u n i t y f o r more p r e c i s e d o c u m e n t a t i o n o f weather p a t t e r n s t h a n was p r e v i o u s l y o b t a i n e d (Chapter 6 ) . I n 1987, t h e f i r s t e x p e r i m e n t a l season i n t h e v a l l e y , weather d a t a were c o l l e c t e d s i m u l t a n e o u s l y a t t h e open s i t e and one f o r e s t e d s i t e a t a t i m e ( o f t h e two f o r e s t e d s i t e s u s e d ) . There were n o t a b l e d i f f e r e n c e s i n wind d i r e c t i o n , wind speed, t e m p e r a t u r e , and r e l a t i v e h u m i d i t y between t h e two f o r e s t s e t t i n g s d u r i n g t h e summer. However, i t was v e r y d i f f i c u l t t o know what changes i f any were o c c u r r i n g i n o t h e r p a r t s o f t h e f o r e s t a t t h e same t i m e , and i f so, how t h e y would a f f e c t T. 1ineatum f l i g h t . The s t u d y r e p o r t e d i n t h i s c h a p t e r , n o t o n l y a l l o w s f o r t h e observance o f s i m u l t a n e o u s weather p a t t e r n d i f f e r e n c e s i n t h e f o r e s t , b u t a l s o t h e i r e f f e c t on b e e t l e f l i g h t , s i n c e marked b e e t l e s were s i m u l t a n e o u s l y r e l e a s e d from b o t h s i t e s . As i n t h e p r e v i o u s two m a r k - r e c a p t u r e s t u d i e s , m e t e o r o l o g i c a l measurements were made o n l y on days t h a t b e e t l e s were r e l e a s e d . These days were warm and sunny w i t h t e m p e r a t u r e s r i s i n g above 15°C under t h e f o r e s t canopy. T h e r e f o r e , m e t e o r o l o g i c a l comparisons between t h e s i t e s a p p l i e s o n l y f o r t h e s e s p e c i f i c c o n d i t i o n s . 140 Both r e l e a s e s i t e s A and B, l o c a t e d 25 m below a r o a d and 650 m a p a r t from each o t h e r ( F i g . 23) were s i m i l a r i n v e g e t a t i o n c o v e r and d e n s i t y . From e i g h t measurement days t h r o u g h t h e e x p e r i m e n t a l p e r i o d o f June and J u l y , 1988, t e m p e r a t u r e s between t h e two s i t e s , n e v e r d i f f e r e d from each o t h e r by more t h a n 1°C (Table 13). I n c o n t r a s t , t h e t e m p e r a t u r e i n t h e open s i t e was g e n e r a l l y 2 - 4°C warmer t h a n t h e f o r e s t e d s i t e s . T h i s i s s i m i l a r t o t h e r e s u l t s i n C h a p t e r 6. R e l a t i v e h u m i d i t y was a l s o s i m i l a r a t b o t h s i t e s and was u s u a l l y h i g h e r t h a n t h e open s i t e ( T a b l e 1 3 ) . A g a i n t h i s i s c o n s i s t e n t w i t h t h e p r e v i o u s y e a r ' s s t u d y ( C h a p t e r 6) . Wind speed was v e r y s i m i l a r i n b o t h f o r e s t e d s i t e s . A t s i t e A, t h e average wind speed ranged from 1.3 - 2.3 km/h, w i t h s t a n d a r d d e v i a t i o n s r a n g i n g from 0.4 - 0.8 km/h ( T a b l e 1 3 ) . A t s i t e B, t h e average w i n d speed ranged between 1.2 -2.0 km/h, w i t h s t a n d a r d d e v i a t i o n s r a n g i n g from 0.5 - 0.8 km/h. The average wind speeds and r e s p e c t i v e v a r i a t i o n s a r e v i r t u a l l y t h e same as t h o s e measured i n 1987 from t h e f o r e s t e d s i t e s i n t h e v a l l e y (Chapter 6 ) , and from t h e p r e v i o u s y e a r i n t h e second-growth f o r e s t ( C h a p t e r 5 ) . T h i s i n d i c a t e s t h a t i f t h e w e i g h t i n g o f i n d i v i d u a l w ind d i r e c t i o n s by wind speed had a m i n i m a l e f f e c t on t h e c o m p u t a t i o n o f average wind d i r e c t i o n from t h e f i r s t mark-r e c a p t u r e s t u d y (Chapter 5 ) , i t s h o u l d n o t be i m p o r t a n t i n t h i s s t u d y . As a r e s u l t , t h e w e i g h t i n g o f wind d i r e c t i o n by wind speed was n o t c a l c u l a t e d . 141 Table 13. Weather measurements made between 1130 and 1700 h PDT, from the Cedar Creek Valley in the Coquitlam Lake Watershed i n B r i t i s h Columbia during 1988. Exp. Rel. Julian S i t e X S.D. Temperature Relative Date Wind speed (km/h) (°C) Humidity (%) (km/h) 1 1 165 Open 4.41 A 1.25 B 1.55 2 170 Open 6.10 A 1.91 B 1.46 3 177 Open 4.40 A 1.47 B 1.22 4 195 Open 6.77 A 2.03 B 1.57 2 1 204 Open 5.04 A 1.68 B 2.01 2 207 Open 6.97 A 2.15 B 1.92 3 211 Open 5.80 A 1.93 B 1.63 4 215 Open 6.90 A 2.32 B 1.88 1.74 19.6 40.9 0.39 16.8 61.8 0.45 16.3 53.8 1.86 28.0 23.9 0.62 24.4 42.8 0.48 23.4 43.0 1.44 20.4 39.4 0.38 18.2 52.8 0.47 18.0 56.5 1.45 23.5 39.9 0.46 20.0 49.6 0.70 20.9 49.4 1.81 27.8 24.1 0.76 23.9 42.4 0.55 24.3 39.2 1.36 22.7 67.2 0.43 21.1 65.0 0.80 20.6 64.6 1.92 28.8 35.2 0.77 25.7 46.8 0.60 26.0 47.0 1.51 23.4 51.6 0.76 21.4 . 58.3 0.78 20.8 57.6 142 The w ind d i r e c t i o n d a t a a r e p r e s e n t e d i n d i v i d u a l l y as c i r c u l a r h i s t o g r a m s ( F i g s . 24 - 2 7 ) . The b l a c k b a r s r a d i a t i n g out o f t h e c i r c l e c i r c u m f e r e n c e r e p r e s e n t t h e f r e q u e n c y o f measured v a l u e s from t h a t p a r t i c u l a r d i r e c t i o n . W i t h i n each c i r c l e , t h e d o t t e d l i n e r e p r e s e n t s t h e 0 W and v e c t o r l e n g t h f o r t h e f i r s t 2 h o r f i r s t h a l f o f t h e m e a s u r i n g p e r i o d f o r t h a t day. The dashed l i n e i s t h e same f o r t h e l a t t e r 2 h o r t h e l a t t e r h a l f o f t h e m e a s u r i n g p e r i o d f o r t h a t day. The s o l i d l i n e r e p r e s e n t s t h e o v e r a l l jzfw and v e c t o r l e n g t h f o r t h e m e asuring p e r i o d . I f t h e l i n e s a r e p r e s e n t , t h e n t h e d i r e c t i o n s a r e c o n s i d e r e d s i g n i f i c a n t based on t h e R a y l e i g h t e s t f o r randomness ( B a t s c h e l e t 1981). F o r a l l m e a s u r i n g p e r i o d s , a t each s i t e , j 0 w was s i g n i f i c a n t . As i n t h e f i r s t v a l l e y s t u d y , average wind d i r e c t i o n a t t h e open s i t e was s i g n i f i c a n t l y d i r e c t e d u p - v a l l e y on a l l t e s t days ( F i g s . 24 - 2 7 ) . U p - v a l l e y winds were a l s o v e r y c o n s i s t e n t a t r e l e a s e s i t e A. The o n l y d i f f e r e n c e was a l a r g e amount o f v a r i a b i l i t y d u r i n g t h e f i r s t r e l e a s e o f e x p e r i m e n t 2 ( F i g . 26A). F o r a l l b u t t h e f i r s t r e l e a s e o f b o t h e x p e r i m e n t s , u p - v a l l e y w ind p r e d o m i n a t e d a l s o a t f o r e s t e d s i t e B. I n e xperiment 1, r e l e a s e 1, w ind blew from up- t o d o w n - v a l l e y w i t h a J0W o f 27°, d u r i n g t h e f i r s t h a l f o f t h e m e asuring day ( F i g . 24A). I n e x p e r i m e n t 2, r e l e a s e 1, t h e o v e r a l l 0W f o r t h e day was 10° ( F i g . 24B). I n t h e f i r s t v a l l e y s t u d y i n 1987, a l l measurements from s i t e B (same i n b o t h s t u d i e s ) showed 0W blew u p - v a l l e y . T h i s i n d i c a t e s t h e u p - v a l l e y winds t h a t t y p i c a l l y o c c u r d i u r n a l l y 143 A. Experiment 1/Release 1 1 2 3 4 5 ( 7 I 9 10 1 1 1 2 1 3 FUNNEL TRAP GROUPS F i g . 24. Wind d i r e c t i o n and T. l i n e a t u m f l i g h t r e c a p t u r e f o r e x p e r i m e n t 1, r e l e a s e s 1 (A) and 2 (B) , r e p r e s e n t e d by c i r c u l a r and xy a x i s h i s t o g r a m s , r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n a l o n g t h e l e n g t h o f t h e v a l l e y . The x - a x i s r e p r e s e n t s t h e v a l l e y a z i m u t h . L i n e s i n t h e c i r c l e s a r e t h e mean d i r e c t i o n f o r t h e f i r s t 2 h ( d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g t h e r - v a l u e f o r t h e v e c t o r . 144 < U W cc in -t w ea Cs. O cc w CQ 20 -16 -12 -8 -4 -0 20 -16 -12 8 4 0 Tt A. Experiment 1/Release 3 e c s > o c B. Experiment 1/Release 4 ^ lb 11 12 13 MALE FEMALE Site A SlttB I 1 I 1 I 1 ' I ' I 3? •o TTT TTT 1 1 1 1 1 " i 1 1 • i • i 9 10 11 12 13 B s-D CD FUNNEL TRAP GROUPS F i g . 25. Wind d i r e c t i o n and T. l i n e a t u m f l i g h t r e c a p t u r e f o r e x p e r i m e n t 1, r e l e a s e s 3 (A) and 4 ( B ) , r e p r e s e n t e d by c i r c u l a r and xy a x i s h i s t o g r a m s , r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n a l o n g t h e l e n g t h o f t h e v a l l e y . The x - a x i s r e p r e s e n t s t h e v a l l e y a z i m u t h . L i n e s i n t h e c i r c l e s a r e t h e mean d i r e c t i o n f o r t h e f i r s t 2 h ( d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g t h e r - v a l u e f o r t h e v e c t o r . 145 Q CO W -fa O W D Z 40 -( 30 20 10 0 40 30 20 10 0 40 -30 20 -I 10 0 40 - | 30 20 -10 -0 1 2 i 1 i 1 2 A. Experiment 2/ReIease 1 Weather s l t t A | Station • „ n PI _ H R SlteB 3 4 5 6 7 8 B. Experiment 2/Release 2 9 10 11 12 MALE FEMALE I 1 'I' 1 I S i u B • I ' I 3 4 ,,B,B 5 6 7 8 FUNNEL TRAP GROUPS f n r T 9 10 r 1 r 1 11 12 o D BB ? •o e o B BB F i g . 26. Wind d i r e c t i o n and T. l i n e a t u m f l i g h t r e c a p t u r e f o r e x p e r i m e n t 2, r e l e a s e s 1 (A) and 2 ( B ) , r e p r e s e n t e d by c i r c u l a r and xy a x i s h i s t o g r a m s , r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n a l o n g t h e l e n g t h o f t h e v a l l e y . The x - a x i s r e p r e s e n t s t h e v a l l e y a z i m u t h . L i n e s i n t h e c i r c l e s a r e t h e mean d i r e c t i o n f o r t h e f i r s t 2 h ( d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g t h e r - v a l u e f o r t h e v e c t o r . 146 A. Experiment 2/Release 3 40 -30 20 10 Weather Station 40 30 20 10 i i ' T T • V • T • 1 2 3 4 5 6 7 8 B. Experiment 2/Release 4 T fU. 9 10 11 12 MALE FEMALE 40 -30 20 -10 -Weather SUUOD I Site A I SlteB 40 30 20 10 0 ' I ' " l " • T ' " I 5 6 7 8 ' I ' i I ' i ' 9 10 11 12 1 2 • I I 3 4 FUNNEL TRAP GROUPS 27. Wind d i r e c t i o n and T. l i n e a t u m f l i g h t r e c a p t u r e f o r e x p e r i m e n t 2, r e l e a s e s 3 (A) and 4 ( B ) , r e p r e s e n t e d by c i r c u l a r and xy a x i s h i s t o g r a m s , r e s p e c t i v e l y , p l a c e d r e l a t i v e t o t h e i r l o c a t i o n a l o n g t h e l e n g t h o f t h e v a l l e y . The x - a x i s r e p r e s e n t s t h e v a l l e y a z i m u t h . L i n e s i n t h e c i r c l e s a r e t h e mean d i r e c t i o n f o r t h e f i r s t 2 h ( d o t t e d ) , f i n a l 2 h (dashed), and a l l day ( s o l i d ) , f o l l o w i n g b e e t l e r e l e a s e , w i t h l i n e l e n g t h s i g n i f y i n g t h e r - v a l u e f o r t h e v e c t o r . 147 t h r o u g h o u t t h e summer, on warm, c l o u d l e s s days, i n mountainous t e r r a i n ( B a r r y 1981), i s n o t a b s o l u t e . However, based on two y e a r s o f measurements, we found t h i s phenomenon t o be c o n s i s t e n t . E x p e r i m e n t 1 The p r o p o r t i o n o f marked b e e t l e s r e c a p t u r e d t h r o u g h t h e f o u r r e l e a s e s averaged c a . 1% (Table 1 4 ) . The % r e c a p t u r e among r e l e a s e s were s i m i l a r , r a n g i n g from 0.8 - 1.3%. Over 11% o f t h e r e c a p t u r e d b e e t l e s were caught d u r i n g t h e second c o l l e c t i o n d a t e o f t h e r e s p e c t i v e r e l e a s e s , w h i c h i s a much l o w e r v a l u e t h a n was r e c o r d e d i n 1987. The number o f unmarked b e e t l e s c a p t u r e d was i n i t i a l l y v e r y h i g h , w i t h a d e c r e a s i n g t r e n d as t h e summer p r o g r e s s e d , r e s u l t i n g i n a t o t a l o f 148 000 unmarked b e e t l e s c a p t u r e d . B e e t l e r e c a p t u r e p a t t e r n i s f i r s t r e l a t e d t o wind d i r e c t i o n . D u r i n g t h e f i r s t r e l e a s e , w h i l e w i n d blew up-v a l l e y from t h e weather s t a t i o n and s i t e A, i t blew down-v a l l e y from s i t e B, e s p e c i a l l y d u r i n g t h e f i r s t 2 h f o l l o w i n g b e e t l e r e l e a s e ( F i g . 2 4 ) . The b e e t l e r e c a p t u r e d a t a s u p p o r t s t h e h y p o t h e s i s o f downwind movement, e s p e c i a l l y f o r p o p u l a t i o n B, where o v e r 80% were r e c a p t u r e d downwind ( d o w n - v a l l e y ) , i n t r a p groups 1 - 7 ( F i g . 24A). P o p u l a t i o n A b e e t l e s were r e c a p t u r e d up- and d o w n - v a l l e y , w h i c h may have r e s u l t e d from t h e o p p o s i n g J2f w's a t s i t e s A and B. 148 Table 14. Flight and recapture success of Trypodendron lineatum in the mark-recapture study conducted at Coquitlam Lake Watershed i n B r i t i s h Columbia during 1988. Captured Beetles  Marked Unmarked No. No. % of Flown Exp. Release Population Released Flying % Flown Total No. Beetles No. 1 1 A 4 000 3 166 79.2 16 0.5 B 4 000 3 250 81.3 46 1.4 Total 8 000 6 416 80.2 62 1.0 66 178 2 A 4 000 3 486 87.2 24 0.7 B 4 000 3 463 86.6 29 0.8 Total 8 000 6 949 86.9 53, 0.8 43 537 3 A 4 000 3 544 88.6 22 0.6 B 4 000 3 428 85.7 71 2.1 Total 8 000 6 972 87.2 93 1.3 27 226 4 A 4 000 2 898 72.5 19 0.7 B 4 000 2 608 65.2 35 1.3 Total 8 000 5 506 68.8 54 1.0 11 025 Population A 16 000 13 094 81.8 81 0.6 Totals B 16 000 12 749 79.7 181 1.4 Total 32 000 25 843 80.1 262 1.0 147 966 2 1 A 4 000 2 790 69.8 84 3.0 B 4 000 3 188 79.7 64 2.0 Total 8 000 5 978 74.7 148 2.5 24 519 2 A 4 000 3 370 84.3 99 2.9 B 4 000 2 790 69.8 28 1.0 Total 8 000 6 160 77.0 127 2.1 5 209 3 A 4 000 2 790 69.8 101 3.6 B 4 000 2 442 61.1 57 2.3 Total 8 000 5 232 65.4 158 3.0 3 306 4 A 4 000 2 906 72.7 121 4.2 B 4 000 2 442 61.1 68 2.8 Total 8 000 5 348 66.9 189 3.5 2 276 Population A 16 000 11 856 74.1 405 3.4 Totals B 16 000 10 862 67.9 217 2.0 Total 32 000 22 718 71.0 622 2.7 35 310 149 F o r r e l e a s e s 2 - 4 , ^ w was p r e d o m i n a n t l y down- t o up-v a l l e y a t a l l t h r e e measurement s i t e s . I n most c a s e s , b e e t l e f l i g h t d i r e c t i o n f o r b o t h p o p u l a t i o n s A and B were downwind ( F i g s . 24B and 25A and B ) . P o p u l a t i o n B were r e c a p t u r e d u p - v a l l e y (downwind) f o r a l l t h r e e r e l e a s e s , w h i l e p o p u l a t i o n A b e e t l e s were r e c a p t u r e d s i m i l a r l y i n two o f t h e t h r e e r e l e a s e s . The n o t a b l e e x c e p t i o n i s f o r r e l e a s e 2, where p o p u l a t i o n A b e e t l e s were r e c a p t u r e d d o w n - v a l l e y , a p p a r e n t l y upwind. The r e a s o n b e h i n d t h i s d e v i a t i o n i n p a t t e r n cannot be r e a d i l y e x p l a i n e d . O v e r a l l a n a l y s i s o f up- and d o w n - v a l l e y b e e t l e r e c a p t u r e p a t t e r n s show t h a t when wi n d was s i g n i f i c a n t l y d i r e c t e d u p - v a l l e y a t a l l t h r e e measurement s i t e s , a s i g n i f i c a n t l y h i g h e r number o f p o p u l a t i o n A and B b e e t l e s were r e c a p t u r e d u p - v a l l e y (downwind) from t h e r e l e a s e s i t e s ( T a b l e 1 5 ) . Data from t h e 1987 v a l l e y s t u d y s u g g e s t e d t h a t b e e t l e s e i t h e r a v o i d e d f l y i n g o r c o u l d n o t s u c c e s s f u l l y o r i e n t t h e m s e l v e s t o s e m i o c h e m i c a l - b a i t e d t r a p s i n t h e c e n t e r o f an open s e t t i n g , where wind p a t t e r n s t e n d t o be q u i t e u n s e t t l e d ( F i g . 2 1 ) . T h i s n o t i o n i s extended t o compare t h e number o f b e e t l e s r e c a p t u r e d i n t h e f o r e s t s e t t i n g , where wind speeds a r e low, and t h e open s e t t i n g , where wind speeds a r e c o n s i d e r a b l y h i g h e r ( T a b l e s 8 and 13). W i t h i n t h e p l o t d e s i g n used f o r t h i s e x p e r i m e n t , a l l f o u r t r a p groups i n t h e f o r e s t ( 1 , 2, 11, and 12) were f u r t h e r away from t h e b e e t l e r e l e a s e s i t e s t h a n a l l t h e t r a p groups i n t h e open (3 - 11) ( F i g . 23A). Y e t t h e mean number o f b e e t l e s Table 15. Comparison between up and down-valley movement of adult Trypodendron Iineatum during the mark-recapture study conducted i n the Cedar Creek Valley within the Coquitlam Lake Watershed in B r i t i s h Columbia during 1988. X Number of Harked Beetles Caught/Release ^ Exp. Trap Population Trap Population Locations A S.D. Locations B S.D. 1 Up-valley 10.3 5.1 Down-valley 2.3 2.5 * 3 (7,8,9,10,11,12,13)2 (1,2,3,4,5,7) 1 Down-valley 7.7 9.1 Up-valley 37.0 19.3 (1,2,3,4,5) (9,10,11.12,13) 2 Up-valley 87.0 11.5 Down-valley 13.3 7.4 (5,6,7,8.9,10,11,12) (1,2,3,4,5,6,7) 2 Down-valley 0.7 1.2 Up-valley 20.7 9.9 * (1.2,3,4) (9,10,11,12) Data are from releases 2, 3, and 4 for both experiments, where up-valley wind prevailed at a l l s i t e s during the release days. Numbers i n parentheses represent trap groups from Figure 23. Significant differences within a row are indicated by a star (Mann-Whitney U test; p < 0.10); the most sensitive probability t h i s test can provide given a sample size of 3. 151 r e c a p t u r e d / r e l e a s e i n t h e f o r e s t e d t r a p groups was s i g n i f i c a n t l y g r e a t e r t h a n t h o s e r e c a p t u r e d i n t h e open t r a p groups ( U 4 4 = 16; p < 0.05), w i t h a r e c a p t u r e r a t i o o f 3:1 ( f o r e s t : o p e n ) . I t i s a p p a r e n t t h a t w ind speed i s p r o b a b l y one o f t h e p r i m a r y r e a s o n s f o r t h i s , as i n d i c a t e d by t h e d a t a i n C h a p t e r s 2 and 6. One o t h e r c o n s i d e r a t i o n i s t h e a b i l i t y o f T. l i n e a t u m t o d i s p e r s e from one s i d e o f t h e v a l l e y t o t h e o t h e r . Trap groups 2, 6, 7, 8, and 13 were s e t a c r o s s t h e c r e e k from b o t h r e l e a s e s i t e s ( F i g . 23A). Throughout t h e f o u r r e l e a s e s , 38% o f t h e 262 t o t a l r e c a p t u r e d b e e t l e s were c o l l e c t e d from t h e s e t r a p s w h i c h r e p r e s e n t e d 38% o f a l l t h e t r a p s d e p l o y e d i n t h e v a l l e y . The d a t a show t h a t b e e t l e s a r e c a p a b l e o f f l y i n g a c r o s s v a l l e y s , w i t h no a p p a r e n t b a r r i e r p r e s e n t e d by t h e c r e e k . E x p e r i m e n t 2 A new p l o t d e s i g n ( F i g . 23B), w i t h t h e o b j e c t i v e s o f n o t a l l o w i n g t h e p r e s e n c e o f any t r a p group w i t h i n 200 m o f t h e r e l e a s e s i t e s , w h i l e p r o v i d i n g i n s i g h t i n t o t h e f l i g h t r o u t e s t a k e n by T. l i n e a t u m . r e s u l t e d i n an i n c r e a s e up t o 2.7% o f t h e p r o p o r t i o n o f marked b e e t l e s r e c a p t u r e d ( T a b l e 1 4 ) . T h i s can be a t t r i b u t e d t o t r a p groups 6, 7, and 8, p l a c e d i n between b o t h b e e t l e r e l e a s e s i t e s , w h i c h caught 431/583 (74%) o f t h e marked b e e t l e s r e c a p t u r e d d u r i n g t h i s e x p e r i m e n t ( F i g s . 26 and 2 7 ) . Over 26% o f t h e r e c a p t u r e d b e e t l e s were c o l l e c t e d d u r i n g t h e second c o l l e c t i o n d a t e f o r 152 t h e r e s p e c t i v e r e l e a s e s . I t i s p o s s i b l e t h a t t h e h i g h e r p r o p o r t i o n o f b e e t l e s r e c a p t u r e d d u r i n g t h i s p e r i o d , when compared t o e x p e r i m e n t 1 d a t a , may have been a r e s u l t o f t h e i n c r e a s i n g l y n e g a t i v e e f f e c t s s t o r a g e has on t h e b e e t l e s t h r o u g h t i m e , r e s u l t i n g i n b e e t l e s b e i n g s l o w e r d i s p e r s e r s . N i j h o l t (1967) found t h a t t h e f a t c o n t e n t o f T. l i n e a t u m d e c r e a s e d as s t o r a g e t i m e i n c r e a s e d . The p r o p o r t i o n o f r e l e a s e d b e e t l e s f l y i n g a l s o d e c r e a s e d w i t h t i m e ( T a b l e 1 4 ) , as f u r t h e r e v i d e n c e t o t h e weaker c o n d i t i o n o f t h e i n s e c t s . I n s u p p o r t o f t h i s h y p o t h e s i s , t h e p r o p o r t i o n r e c a p t u r e d d u r i n g t h e second c o l l e c t i o n s t e n d e d t o i n c r e a s e w i t h t i m e ; 13, 15, 31, and 22% f o r r e l e a s e 1 t h r o u g h 4, r e s p e c t i v e l y . The number o f unmarked b e e t l e s r e c a p t u r e d d u r i n g t h e e x p e r i m e n t dropped t o 35 000 b e e t l e s ( T a b l e 1 4 ) , as was e x p e c t e d from o t h e r s e a s o n a l c a t c h r e c o r d s (Chapman and N i j h o l t 1980; R u d i n s k y and Daterman 1964; Shore e t a l . 1986). B e e t l e r e c a p t u r e b e e t l e p a t t e r n s may a g a i n be r e l a t e d t o wind. F o r a l l b u t t h e f i r s t r e l e a s e , # w blew p r e d o m i n a n t l y down- t o u p - v a l l e y a t a l l t h r e e m e a s u r i n g s i t e s ( F i g s . 26 and 2 7 ) . D u r i n g t h e f i r s t r e l e a s e , t h e same # w was o b s e r v e d a t t h e weather s t a t i o n and s i t e A ( F i g . 2 6A). But a g a i n , as i n e x p e r i m e n t 1, r e l e a s e 1, J0 W a t s i t e B came from t h e o p p o s i t e d i r e c t i o n , u p - v a l l e y . P o p u l a t i o n A b e e t l e s were r e c a p t u r e d p r e d o m i n a n t l y u p - v a l l e y (downwind) i n t r a p groups 6, 7, and 8. I n c o n t r a s t , an e q u a l number o f p o p u l a t i o n B b e e t l e s were caught up- and d o w n - v a l l e y . T h i s 153 p a t t e r n may have o c c u r r e d due t o t h e o p p o s i n g #w''s measured from s i t e s A and B. I n r e l e a s e s 2 - 4 , a l m o s t a l l p o p u l a t i o n A b e e t l e s were r e c a p t u r e d u p - v a l l e y (downwind) and a h i g h e r p r o p o r t i o n o f r e c a p t u r e d p o p u l a t i o n B b e e t l e s were c o l l e c t e d u p - v a l l e y t h a n d o w n - v a l l e y ( F i g s 26B and 27A and B ) . D e s p i t e s t r o n g u p - v a l l e y w i n d s , some p o p u l a t i o n B b e e t l e s were r e c a p t u r e d i n t r a p s 6, 7, and 8. One e x p l a n a t i o n f o r t h i s may be t h a t g i v e n a c h o i c e , some b e e t l e s may have p r e f e r r e d t o f l y upwind under g e n t l e r wind c o n d i t i o n s i n t h e f o r e s t , r a t h e r t h a n f l y downwind under h i g h e r w ind speed c o n d i t i o n s i n t h e open. A n a l y s i s o f up-and d o w n - v a l l e y b e e t l e r e c a p t u r e p a t t e r n s show t h a t when wind was s i g n i f i c a n t l y d i r e c t e d u p - v a l l e y , a t a l l t h r e e measurement s i t e s , a s i g n i f i c a n t l y h i g h e r number o f p o p u l a t i o n A and B b e e t l e s were r e c a p t u r e d u p - v a l l e y (downwind) from t h e i r r e l e a s e s i t e s ( T a b l e 1 5 ) . I n an e f f o r t t o see i f b e e t l e s used a r o a d as a f l y w a y , a c o m p a r i s o n among r e c a p t u r e d b e e t l e numbers i n t r a p groups 6, 7, and 8 f o r u p - v a l l e y f l y i n g p o p u l a t i o n A and down-v a l l e y f l y i n g p o p u l a t i o n B b e e t l e s was made ( F i g . 23B). No s t a t i s t i c a l d i f f e r e n c e s between t r a p groups were found f o r b o t h r e c a p t u r e d p o p u l a t i o n s A and B ( T a b l e 1 6 ) . A s l i g h t l y h i g h e r number o f b e e t l e s from b o t h p o p u l a t i o n s were r e c a p t u r e d i n t r a p group 7 (on t h e r o a d ) , y e t t h i s may have been due t o t h e r e l e a s e s i t e s b e i n g c l o s e r t o t h i s t r a p group t h a n t h e o t h e r two. The number o f u p - v a l l e y A b e e t l e s r e c a p t u r e d i n t r a p groups 6, 7, and 8 was c o n s i d e r a b l y T a b l e 16. Movement o f T. l i n e a t u m t h r o u g h a f o r e s t , f l y i n g u p - v a l l e y ( p o p u l a t i o n A) and d o w n - v a l l e y ( p o p u l a t i o n B) d u r i n g f o u r s e p a r a t e r e l e a s e s i n E x p e r i m e n t 2, w i t h i n t h e C o q u i t l a m Lake Watershed i n B r i t i s h C olumbia d u r i n g 1 9 8 8 . 1 / 2 Trap G r o u p s 3 X Number o f P o p u l a t i o n A B e e t l e s Caught/ R e l e a s e S.D. X Number o f P o p u l a t i o n B B e e t l e s Caught/ R e l e a s e S. D. 30 m Above The Road (8) 29.8 6.2 4.0 2. 7 On The Road (7) 37.0 12.7 9.5 6. 4 100 m Below The Road (6) 22 .8 10.2 4.3 1. 9 1 B e e t l e r e c a p t u r e d a t a a r e from one r e l e a s e p e r i o d t o t h e n e x t . No s i g n i f i c a n t d i f f e r e n c e s were found between t r a p groups f o r b o t h p o p u l a t i o n s A and B ( K r u s k a l - W a l l i s t e s t ; p > 0.05). Number i n p a r e n t h e s e s r e p r e s e n t s t r a p group number from F i g u r e 23B. 155 h i g h e r t h a n t h e number o f d o w n - v a l l e y B b e e t l e s , a d d i n g s u p p o r t t o t h e downwind f l i g h t h y p o t h e s i s ( T a b l e 1 6 ) . I n f l i g h t r o u t e comparisons between t r a p s groups i n t h e f o r e s t e d and open s i t e s , t h e r e s u l t s t h r o u g h t h e f o u r r e l e a s e s show t h a t T. l i n e a t u m were r e c a p t u r e d i n h i g h e r numbers i n f o r e s t e d t r a p groups 4 and 9 t h a n i n t h e non-f o r e s t e d groups 3 and 10 ( T a b l e 1 7 ) . U n f o r t u n a t e l y , t h i s does n o t n e c e s s a r i l y i n d i c a t e b e e t l e f l i g h t r o u t e , as i t may j u s t i n d i c a t e , as i n e x p e r i m e n t 1, t h a t T. l i n e a t u m a r e more e a s i l y r e c a p t u r e d i n t h e f o r e s t t h a n i n t h e open s e t t i n g . I n c o n c l u s i o n , i t was found t h a t w i t h o u t t h e p r e s e n c e o f c l o s e - r a n g e a t t r a c t a n t s , T. l i n e a t u m g e n e r a l l y f l y downwind i n t h e i r s e a r c h f o r h o s t m a t e r i a l and mates. I n t h e Cedar Creek V a l l e y , t h i s was u s u a l l y u p - v a l l e y , as a r e s u l t o f t h e c h a r a c t e r i s t i c d i u r n a l u p - v a l l e y w i n d p a t t e r n s i n v a l l e y systems ( B a r r y 1981). I t i s i m p o r t a n t t o n o t e however t h a t d u r i n g t h e s e i d e a l weather c o n d i t i o n s , t h e wind sometimes d e v i a t e d from an u p - v a l l e y d i r e c t i o n . T h i s was n o t i c e d t w i c e a t s i t e B, and may be a r e s u l t o f w i n d e d d i e s f o r m i n g a t t h e edge o f t h e f o r e s t s e t t i n g . As e x p l a i n e d by Kimmins (1987), a l a m i n a r a i r f l o w o v e r a c o o l f o r e s t w i l l c u r l i n t o t h e f o r e s t a t t h e upwind edge o f an open s i t e . T h i s phenomenon may have been more pronounced on t h e two o c c a s i o n s t h a t s i t e B 0 W d i f f e r e d from t h e J0W a t t h e o t h e r two measurement s i t e s . An i m p o r t a n t outcome o f t h i s s t u d y i s t h a t T. l i n e a t u m c l e a r l y e i t h e r f l i e s b e t t e r i n t h e f o r e s t , o r i e n t s t o h o s t s T a b l e 17. Comparison between an open and f o r e s t e d r o u t e f o r l o n g d i s t a n c e f l y i n g Trypodendron 1ineatum, d u r i n g 4 r e l e a s e s i n Exp e r i m e n t 2, w i t h i n t h e C o q u i t l a m Lake Watershed i n B r i t i s h Columbia d u r i n g 1988. X Number o f Marked B e e t l e s Caught/Release x S e t t i n g D own-valley U p - v a l l e y T r a p s S.D. Traps S.D. Open ( 3 ) 2 0.0 b 3 0.0 (10) 4.0 b 3.1 F o r e s t e d (4) 3.5 a 2.4 (9) 11.3 a 3.6 B e e t l e s from p o p u l a t i o n s A and B were combined, u s i n g r e c a p t u r e d a t a from one r e l e a s e p e r i o d t o t h e n e x t . Number i n p a r e n t h e s e s r e p r e s e n t s t r a p group number from F i g u r e 23B. D i f f e r e n t l e t t e r s i n a column i n d i c a t e s s i g n i f i c a n t d i f f e r e n c e s between s e t t i n g s (Mann-Whitney U t e s t ; p < 0.05). 157 b e t t e r i n t h e f o r e s t , o r b o t h , r a t h e r t h a n i n an open s e t t i n g . I n ex p e r i m e n t 1, more b e e t l e s were r e c a p t u r e d i n t r a p groups i n f o r e s t s t h a n i n open s e t t i n g s , o f w h i c h t h e l a t t e r were a c t u a l l y c l o s e r t o t h e r e l e a s e s i t e s . I n e x p e r i m e n t 2, t r a p groups i n f o r e s t s c o n s i s t e n t l y r e c a p t u r e d more b e e t l e s t h a n i n open s i t e s . D u r i n g t h e f i r s t v a l l e y s t u d y i n 1987, more b e e t l e s were r e c a p t u r e d a t t h e edge o f an opening t h a n i n t h e c e n t e r , when t h e edge was f u r t h e r away from t h e r e l e a s e s i t e s ( C h apter 6 ) . Thus, we have extended t h i s continuum i n t o t h e f o r e s t . The r e l a t i v e l y l i g h t wind speed c h a r a c t e r i s t i c under a f o r e s t canopy i s most l i k e l y r e s p o n s i b l e f o r t h e i n c r e a s e d number o f b e e t l e s r e c a p t u r e d . O t h e r s have s u g g e s t e d t h a t v e r t i c a l s i l h o u e t t e s w i t h i n t h e f o r e s t s e r v e t o s t i m u l a t e b e e t l e f l i g h t (S. Bombosch p e r s o n a l communication)! 6-. However, f o r a b e e t l e t h a t i s i n i t i a l l y p h o t o p o s i t i v e upon s p r i n g emergence, and has e v o l v e d t o a t t a c k b o t h windthrown ( h o r i z o n t a l ) and b a r k b e e t l e k i l l e d t i m b e r ( v e r t i c a l ) , t y p i c a l l y o r i e n t e d h o r i z o n t a l l y , t h i s h y p o t h e s i s i s l e s s p l a u s i b l e . A d d i t i o n a l l y , i n a wind t u n n e l s t u d y d e s c r i b e d i n A p pendix 1, no p r e f e r e n c e was shown f o r s e m i o c h e m i c a l -b a i t e d s u b s t r a t e s o r i e n t e d v e r t i c a l l y o r h o r i z o n t a l l y . The r e c a p t u r e d a t a a l s o s u g g e s t t h a t f l i g h t d i s t a n c e s o f 1.5 km f o r s p r i n g d i s p e r s i n g T. l i n e a t u m a r e p o s s i b l e . I n t h e f i r s t v a l l e y s t u d y , T. l i n e a t u m e x h i b i t e d t h e a b i l i t y t o f l y 1.9 km, however, t h e f r e q u e n c y o f r e c a p t u r e was n e v e r 16 Dr. S. Bombosch, I n s t i t u t f u r F o r s t z o o l o g i e , U n i v e r s i t a t G o t t i n g e n , Busgenweg 3, D-3400 G o t t i n g e n , W. Germany 158 h i g h e r t h a n one, f o r a p a r t i c u l a r r e l e a s e . The f r e q u e n c i e s o f r e c a p t u r e from t h i s s t u d y a t d i s t a n c e s o f s i m i l a r magnitude were much h i g h e r . The r e a s o n f o r t h e d i f f e r e n c e s between t h e two s t u d i e s i s t h a t d u r i n g t h e most r e c e n t s t u d y , t h e l o n g d i s t a n c e t r a p s were p l a c e d i n t h e f o r e s t . A l s o t h e use o f t r a p groups, i n s t e a d o f i n d i v i d u a l t r a p s i n a l i n e , may have been a more e f f e c t i v e s t r a t e g y i n a t t r a c t i n g and s u b s e q u e n t l y c a p t u r i n g b e e t l e s , by i n c r e a s i n g t h e s i z e and c o n c e n t r a t i o n o f t h e s e m i o c h e m i c a l plumes produced. The d a t a from t h i s s t u d y p r o v i d e a b a s i s f o r c h a r a c t e r i z i n g t h e movement o f T. l i n e a t u m d u r i n g t h e mass f l i g h t p e r i o d i n t h e s p r i n g . I n most woodlands o p e r a t i o n s a l o n g t h e west c o a s t o f Canada, t h e p r e v i o u s y e a r ' s h a r v e s t i n v e n t o r y , n o t y e t removed from t h e f o r e s t b e f o r e t h i s p e r i o d , i s s u s c e p t i b l e t o a t t a c k by T. l i n e a t u m . The d a t a from t h i s and t h e p r e v i o u s c h a p t e r s u g g e s t t h a t placement o f i n v e n t o r y can p l a y an i m p o r t a n t r o l e i n i t s u l t i m a t e a t t a c k . L o g s ^ p r o b a b l y s a f e s t when s t o r e d as f a r away from f o r e s t m a r gins as p o s s i b l e , i n t h e c e n t e r o f an open s e t t i n g . E v e n t u a l l y t h e b e e t l e s w i l l f i n d t h i s m a t e r i a l , b u t c o n d i t i o n s i n an open s e t t i n g i n mountainous t e r r a i n a r e l e s s l i k e l y t o be c o n d u c i v e f o r b e e t l e a t t a c k . A l t e r n a t i v e l y , l o g s n e a r f o r e s t margins s h o u l d be removed more s w i f t l y t h a n t h e l o g s i n t h e c e n t e r o f t h e open s e t t i n g s . Knowledge o f wind p a t t e r n s , e s p e c i a l l y p r e v a i l i n g winds i f t h e y o c c u r , on warm, sunny days i n an a r e a 159 c o n t a i n i n g i n v e n t o r y , can be used as a g u i d e t o d e t e r m i n e w h i c h p o r t i o n o f t h e i n v e n t o r y would be most s u s c e p t i b l e t o a t t a c k based on i t s l o c a t i o n r e l a t i v e t o t h e s u r r o u n d i n g f o r e s t . The i m p l e m e n t a t i o n o f m a s s - t r a p p i n g t e c h n i q u e s used i n l o g s o r t s and s t o r a g e a r e a s has been shown t o s u c c e s s f u l l y r educe p o p u l a t i o n s o f European T. l i n e a t u m w i t h i n a f o r e s t s t a n d d u r i n g a t h r e e y e a r p e r i o d , d e s p i t e a y e a r l y r e p l a c e m e n t o f s u s c e p t i b l e l o g s on t h e s i t e ( K o n i g 1988). McLean and Salom ( u n p u b l i s h e d data) a t t e m p t e d t o r e duce t h e a t t a c k o f s u s c e p t i b l e l o g s on a r i g h t - o f - w a y i n a c o m m e r c i a l f o r e s t on Vancouver I s l a n d , u s i n g m a s s - t r a p p i n g t e c h n i q u e s , however t h e y were u n s u c c e s s f u l . Based on t h e d a t a from C h a p t e r 6 and t h i s c h a p t e r , i t i s l i k e l y t h a t t h e l a c k o f s u c c e s s i n u s i n g t h i s s t r a t e g y r e s u l t e d from t h e placement o f t r a p s n e a r s u s c e p t i b l e l o g s , r a t h e r t h a n a t t h e s i t e s where b e e t l e p o p u l a t i o n s were l i k e l y t o o v e r w i n t e r . I f e e l m a s s - t r a p p i n g does have p o t e n t i a l as a p e s t management t o o l i n p r o t e c t i n g l o g s b e f o r e t h e y a r e t r a n s p o r t e d from t h e f o r e s t . I n a r e a s where h a r v e s t i n g i s c o n t i n u o u s , i n v e n t o r y s u s c e p t i b l e t o s p r i n g a t t a c k and removed from t h e s i t e a f t e r June, s h o u l d be sampled f o r T. l i n e a t u m a t t a c k . I f a t t a c k i s abundant, a l a r g e brood p o p u l a t i o n most l i k e l y has f l o w n t o t h e nearby f o r e s t s t o o v e r w i n t e r (Dyer and K i n g h o r n 1961). I t i s i n t h i s f o r e s t t h a t s e m i o c h e m i c a l - b a i t e d t r a p groups s h o u l d be p u t i n p l a c e t h e f o l l o w i n g s p r i n g . The t r a p s s h o u l d n o t o n l y be p l a c e d a l o n g t h e f o r e s t m a r g i n , b u t 160 as f a r as 100 m i n t o t h e f o r e s t . I n t h e s p r i n g , many o f t h e emerging b e e t l e s s h o u l d be c a p t u r e d i n t h e t r a p s , p r e v e n t i n g them from a t t a c k i n g s u s c e p t i b l e h o s t m a t e r i a l a t new h a r v e s t s i t e s . W h i l e n o t a l l b e e t l e s w i l l be c a p t u r e d t h i s way, i t i s l i k e l y t h a t t h i s t r a p p i n g can reduce t h e p o p u l a t i o n s i n t h e f o r e s t . Combined w i t h s h o r t t i m e i n t e r v a l s between h a r v e s t and t r a n s p o r t o f i n v e n t o r y , a l o n g w i t h t h e c o n t i n u e d m a s s - t r a p p i n g a t t h e l o g s o r t s , I b e l i e v e t h a t p o p u l a t i o n s o f T. l i n e a t u m can be reduced s i g n i f i c a n t l y a t comm e r c i a l woodlands d i v i s i o n s , r e s u l t i n g i n r e d u c t i o n s i n degrade l o s s due t o t h i s s p e c i e s . R e s e a r c h s h o u l d be c o n t i n u e d i n t h i s a r e a t o see i f t h e proposed approach i s p l a u s i b l e and a l s o t o t e s t m o d i f i c a t i o n s t h a t can make t h i s s t r a t e g y work most s u c c e s s f u l l y . 161 CHAPTER 8: CONCLUSIONS D i s p e r s a l The r e a l i z a t i o n t h a t d i s p e r s a l i s a c r i t i c a l a s p e c t o f t h e l i f e h i s t o r y and p o p u l a t i o n dynamics o f many i n s e c t s p e c i e s r e s u l t e d from i m p o r t a n t r e v i e w s on t h e s u b j e c t (Southwood 1962; Johnson 1969) and t h e i m p l i c a t i o n s o f t h e s e v i e w s on p e s t management have been t h e s u b j e c t o f s e v e r a l symposia s i n c e t h e n (Berryman and S a f r a n y i k 1980; Danthanarayana 1986; D e l u c c h i and B a l t e n s w e i l e r 1979; Rabb and Kennedy 1979). The d e f i n i t i o n s used t o d e s c r i b e and c o n t r a s t d i s p e r s a l f r e q u e n t l y d i f f e r . D i n g l e (1985) c o n s i d e r s d i s p e r s a l t o be random o r h aphazard movements whi c h o c c u r p u r e l y as a r e s u l t o f chance c i r c u m s t a n c e s , whereas m i g r a t i o n i s d e f i n e d as s p e c i a l i z e d b e h a v i o r e v o l v e d f o r t h e d i s p l a c e m e n t o f t h e i n d i v i d u a l i n space. Kennedy (1975) s t a t e d t h a t d i s p e r s a l can i n v o l v e s p e c i a l i z e d b e h a v i o r , b u t movements o c c u r w i t h i n t h e h a b i t a t . T h i s c o n t r a s t s w i t h m i g r a t i o n w h i c h r e p r e s e n t s movement between d i f f e r e n t h a b i t a t l o c a l i t i e s . W e l l i n g t o n (1980) d e f i n e d d i s p e r s a l as an a d a p t i v e r e s p o n s e t o h a b i t a t , o f t e n i n v o l v i n g a p e r i o d o f o b l i g a t o r y t r a v e l i ndependent o f p r e v i o u s o r c u r r e n t p o p u l a t i o n d e n s i t i e s . The d e f i n i t i o n p r o v i d e d by W e l l i n g t o n f i t s i n w e l l w i t h t h e l i f e h i s t o r y p a t t e r n s o f s c o l y t i d b e e t l e s w h i c h b e g i n t h e i r d i s p e r s a l s t a g e when t h e y emerge from t h e b r o o d t r e e o r l o g and end when t h e y respond t o h o s t s t i m u l i and/or 162 a t t r a c t i v e pheromones (D. Wood 1982). The f a c t o r s i n d u c i n g d i s p e r s a l and i t s c h a r a c t e r i s t i c s , such as b e h a v i o r a l r e v e r s a l s i n f l i g h t , h o s t s e l e c t i o n , p e r c e p t i o n o f pheromones, and mass response t o a g g r e g a t i o n pheromones a r e c l e a r l y s p e c i e s - s p e c i f i c (Borden 1984) and a r e v e r y much r e l a t e d t o t h e i r h a b i t a t ( A t k i n s 1966b). S p e c i e s - s p e c i f i c v a r i a t i o n o f d i s p e r s a l i s s u b s t a n t i a l and can be i l l u s t r a t e d by c o n s i d e r i n g f l i g h t d i s t a n c e t o h o s t m a t e r i a l . Wollerman (1979) found t h a t t h e maximum d i s t a n c e f l o w n by S c o l y t u s  m u l t i s t r i a t u s (Marsham) was 150 m i n a m a r k - r e c a p t u r e s t u d y . I n c o n t r a s t , B o t t e r w e r g (1982) showed t h a t I p s t y p o q r a p h u s L. a r e c a p a b l e o f f l y i n g 8 km. An even more d r a m a t i c example comes from N i l s s e n (1984) who found t h r e e s c o l y t i d s p e c i e s t r a v e l l e d ( l i k e l y wind blown) 171 km i n n o r t h e r n F i n l a n d . H ost s e l e c t i o n , t h e n e x t s t a g e i n h o s t c o l o n i z a t i o n s t r a t e g i e s f o r s c o l y t i d s i s e i t h e r a random p r o c e s s , i n w h i c h b e e t l e s l a n d i n d i s c r i m i n a n t l y on h o s t s , o r a d i r e c t e d p r o c e s s , i n w h i c h t h e y l a n d p r e f e r e n t i a l l y on h o s t s v s . nonhosts (D. Wood 1982). S e l e c t i o n o f h o s t s by p i o n e e r b e e t l e s a r e f a c i l i t a t e d by t h e i r p e r c e p t i o n o f h o s t v o l a t i l e s (1° a t t r a c t a n t ) , f o l l o w e d by p r o d u c t i o n o f a g g r e g a t i o n pheromones (2° a t t r a c t a n t ) , and t h e i r p e r c e p t i o n and mass re s p o n s e t o them (Borden 1984). A t k i n s (1966b) s u g g e s t e d t h a t 2° a t t r a c t i o n f o r s c o l y t i d s e v o l v e d i n i t i a l l y i n c o n j u n c t i o n w i t h t h e i r u t i l i z a t i o n o f temporary h a b i t a t s , a s s u r i n g maximum u t i l i z a t i o n o f s c a t t e r e d h o s t m a t e r i a l . 163 The a b i l i t y o f a g g r e s s i v e b a r k b e e t l e s t o a t t a c k l i v i n g , h e a l t h y t r e e s t h r o u g h m a s s - a t t a c k s p r o b a b l y a r o s e l a t e r . I n terms o f p e s t management, we must c o n s i d e r d i s p e r s a l and h o s t s e l e c t i o n i n t h e c o n t e x t o f p o p u l a t i o n s t r a t e g i e s . F o r T. l i n e a t u m t h i s r e q u i r e d a c l o s e e x a m i n a t i o n o f 1) t h e e f f e c t i v e d i s t a n c e a p o p u l a t i o n i s c a p a b l e o f f l y i n g t o cause economic d e v a l u a t i o n o f s u s c e p t i b l e l o g s ; 2) t h e i n f l u e n c e o f w i n d , f o r e s t c o v e r , and t o pography on o v e r a l l d i s t r i b u t i o n o f d i s p e r s i n g b e e t l e s ; 3) o p t i m a l e n v i r o n m e n t a l c o n d i t i o n s r e q u i r e d f o r T. l i n e a t u m r e s p o n s e and o r i e n t a t i o n t o o l f a c t o r y s t i m u l i ; and 4) t h e i m p o r t a n c e and r o l e s o f t h e s e m i o c h e m i c a l s used by T. l i n e a t u m . and how such i n f o r m a t i o n r e l a t e s t o b e e t l e t r a p p i n g s t r a t e g i e s . A summary t h a t a d d r e s s e s t h e s e c o n s i d e r a t i o n s based on an i n t e g r a t i o n o f p r e v i o u s r e s e a r c h w i t h t h e r e s e a r c h from t h i s r e p o r t i s p r e s e n t e d below. S p r i n g D i s p e r s a l o f Trypodendron l i n e a t u m O v e r w i n t e r i n g b e e t l e s t h a t emerge and f l y i n t h e s p r i n g when t e m p e r a t u r e s warm up above 15°C (Chapman and K i n g h o r n 1958; R u d i n s k y and Daterman 1964), w i l l u s u a l l y t a k e - o f f t o w a r d t h e open sky (Graham 1959). T h i s has been i n t e r p r e t e d as a p h o t o p o s i t i v e r e s p o n s e . I f no a t t r a c t a n t s a r e p r e s e n t , b e e t l e s w i l l g e n e r a l l y t a k e - o f f i n a l l d i r e c t i o n s under v e r y low w ind speed c o n d i t i o n s (< 1 km/h), w i t h i n c r e a s i n g f r e q u e n c y o f downwind t a k e - o f f as w i n d speed i n c r e a s e s ( F i g . 4 ) . I f a t t r a c t i v e s u s c e p t i b l e h o s t m a t e r i a l 164' i s n e a r t h e emergence s i t e s (+ 25 m), some b e e t l e s w i l l r e s p o n d i m m e d i a t e l y and f l y upwind t o t h e s o u r c e o f a t t r a c t i o n , w h i l e o t h e r n o n - r e s p o n d i n g b e e t l e s w i l l c o n t i n u e on t h e i r way ( F i g s . 17 and 2 0 ) . I n a homogeneous f o r e s t s e t t i n g , b e e t l e s w i l l f l y i n a l l d i r e c t i o n s , w i t h t h e g r e a t e s t d i s p e r s a l d i s t a n c e s r e c o r d e d downwind from t h e emergence s i t e ( F i g . 18). Under h i g h e r w ind speed c o n d i t i o n s , where t h e f o r e s t i s b r o k e n up and may be more i n f l u e n c e d by p r e v a i l i n g winds, b e e t l e f l i g h t w i l l t e n d t o be downwind ( T a b l e 1 5 ) , u n t i l p e r c e p t i o n and r e s p o n s e t o s e m i o c h e m i c a l s s e t s them on an upwind t r a c k t o t h e a t t r a c t i v e s o u r c e . A l o n g t h e windward s i d e o f a f o r e s t m a r g i n , b e e t l e s w i l l f l y p r e d o m i n a n t l y downwind i n t o t h e f o r e s t ( F i g s . 26 and 2 7 ) . However, from t h e l e e w a r d s i d e o f a p r e v a i l i n g w ind, w h i l e many b e e t l e s w i l l f l y downwind i n t o o p e nings a t h i g h wind speeds, some b e e t l e s w i l l a l s o f l y upwind t h r o u g h t h e f o r e s t ( F i g s . 26 and 2 7 ) , where t h e wind speed i s m a r k e d l y l o w e r ( T a b l e 1 3 ) . I n f a c t , b e e t l e c a p t u r e o r l o g a t t a c k i s most s u c c e s s f u l i n t h e f o r e s t ( T a b l e 1 7 ) , where wind speeds a r e low, l e s s s u c c e s s f u l a t t h e edge o f an open s e t t i n g where wind speeds a r e h i g h e r , and l e a s t s u c c e s s f u l i n t h e c e n t e r o f t h e opening where wind speeds and t u r b u l e n c e a r e g r e a t e s t ( F i g . 2 1 ) . Trypodendron l i n e a t u m a r e adapted t o f i n d r a r e h o s t m a t e r i a l i n n a t u r a l f o r e s t s i t u a t i o n s . T h e i r s m a l l s i z e r e s t r i c t s them from f l y i n g a g a i n s t s t r o n g winds (Chapman 19 6 2 ) , and t h e r e f o r e t h e y must o p t i m i z e t h e i r s e a r c h f o r 165 h o s t m a t e r i a l by u s i n g wind t o a i d i n t h e i r d i s p e r s a l . T h i s was i n d i c a t e d by t h e i n c r e a s e d p r o p o r t i o n o f downwind ( F i g . 5 ) , d i r e c t ( F i g . 6) f l i g h t o b s e r v e d , as wind speed i n c r e a s e d i n a wind t u n n e l , and g e n e r a l downwind movement o b s e r v e d i n t h e f i e l d ( T a b l e 15). Our s t u d i e s i n d i c a t e t h a t t h e b e e t l e s can e a s i l y f l y 1.5 km ( F i g s . 24 - 2 7 ) , and can l i k e l y f l y f u r t h e r t h a n 2 km as l o n g as h o s t m a t e r i a l o c c u r s n e a r o r w i t h i n a f o r e s t s e t t i n g . They can f l y t h e s e d i s t a n c e s i n one day, y e t t h e y can a l s o l o c a t e a s o u r c e o f a t t r a c t a n t a f t e r a week ( F i g . 2 2 ) . F l i g h t from one s i d e o f a v a l l e y t o a n o t h e r i s l i t t l e p roblem f o r t h e s e b e e t l e s ( F i g s . 24 and 25) . P r i m a r y a t t r a c t a n t s , e s p e c i a l l y e t h a n o l , a r e i m p o r t a n t f o r f e males s e a r c h i n g o u t , r e s p o n d i n g t o , and a c c e p t i n g h o s t m a t e r i a l (Moeck 1970; F i g s . 13 - 15; T a b l e 4 ) . Once t h e female f i n d s and a c c e p t s t h e h o s t , she p r o d u c e s a p o w e r f u l a g g r e g a t i o n pheromone (Borden and S l a t e r 1969; M a c C o n n e l l e t a l . 1977), l i n e a t i n , t h a t a t t r a c t s b o t h males and f e m a l e s t o mass a t t a c k t h e h o s t . M a l e s , w h i l e l e s s i n f l u e n c e d by e t h a n o l t h a n females a r e ( T a b l e 4 ) , do respond t o i t i n a p o s i t i v e manner w i t h r e s p e c t t o f l i g h t d i r e c t i o n ( F i g . 13) and speed o f r e s p o n s e ( F i g . 1 5 ) . Yet males appear t o r e l y m o s t l y on l i n e a t i n t o f i n d and l a n d on s u i t a b l e h o s t m a t e r i a l ( F i g . 14; T a b l e 4 ) , f a c i l i t a t i n g t h e m a t i n g p r o c e s s w i t h f e m a l e s . The s i g n i f i c a n c e o f q u i c k e r r e s p o n s e s by r e s p o n d i n g males and females when e t h a n o l was p r e s e n t ( F i g . 166 15) p r o b a b l y r e s u l t s from a more complete c h e m i c a l message from h o s t and mates. S t r i k i n g d i f f e r e n c e s i n r e s p o n s e t o h o s t a t t r a c t a n t s have been o b s e r v e d between N o r t h American and European p o p u l a t i o n s o f T. l i n e a t u m . I n Europe, h o s t a t t r a c t a n t s , e t h a n o l and <9£-pinene, when combined w i t h l i n e a t i n , have a s y n e r g i s t i c e f f e c t on t h e number o f b e e t l e s c a p t u r e d (Borden e t a l . 1982; P a i v a and K i e s e l 1985; V i t e and Bakke 1979), r e g a r d l e s s o f what t y p e o f t r a p i s used (Borden e t a l . 1981; McLean e t a l . 1987). I n N o r t h A m e r i c a , h o s t a t t r a c t a n t s a r e i m p o r t a n t t o a l e s s e r degree; t h e r e i s a s t r o n g e r t r a p by a t t r a c t a n t i n t e r a c t i o n a f f e c t i n g T. l i n e a t u m c a p t u r e . B e e t l e s r e s pond t o h o s t a t t r a c t a n t s i n d r a i n p i p e t r a p s (Shore and McLean 1983; F i g . 14; T a b l e 4 ) , y e t do n o t r e s p o n d t o h o s t a t t r a c t a n t s i n a f u n n e l t r a p (Salom and McLean 1988; F i g . 8 ) . V i t e and Bakke (1979) e x p l a i n e d t h a t t h e d r a i n p i p e t r a p r e q u i r e s an e x t r a b e h a v i o r a l r e s p o n s e by t h e b e e t l e s t o e n t e r t h e s m a l l h o l e s t h a t s i m u l a t e g a l l e r y o p e n i n g s . Host v o l a t i l e s enhance t h i s r e s p o n s e . T h i s b e h a v i o r i s n o t n e c e s s a r y f o r t h e c a p t u r e o f b e e t l e s i n f u n n e l t r a p s ( L i n d g r e n 1983). Upon p e r c e p t i o n o f t h e h o s t , b e e t l e s f l y a n e m o t a c t i c a l l y t o i t i n a s t e a d y f l i g h t i n low w ind c o n d i t i o n s . Once near t h e h o s t , t h e b e e t l e s seem t o t e s t e i t h e r t h e c h e m i c a l g r a d i e n t , plume s t r u c t u r e , o r t h e wind, b e f o r e l a n d i n g on t h e l o g , and e i t h e r a t t a c k i t o r mate w i t h a n o t h e r b e e t l e . Under s t i l l w ind c o n d i t i o n s , t h e b e e t l e s 167 w i l l l a n d i n h i g h e r f r e q u e n c i e s , t h a n i f wind i s p r e s e n t ( F i g s . 2 and 7 ) . They appear t o d i v e i n t o t h e h o s t , somehow l a n d i n g u p r i g h t on t h e i r l e g s ( p e r s o n a l o b s e r v a t i o n ) . I t appears t h a t d i f f e r e n t mechanisms o f h o s t f i n d i n g a r e used by T. l i n e a t u m under v a r y i n g wind speed c o n d i t i o n s . I n s t i l l w i n d c o n d i t i o n s , b e e t l e s a r e a r r e s t e d and o r i e n t t o s e m i o c h e m i c a l s . I n t h e p r e s e n c e o f low wind speeds, b e e t l e s a t t r a c t e d t o t h e s e m i o c h e m i c a l s respond t o t h e odor plume u s i n g chemo-anemotaxis (Chapter 2 ) . The a b i l i t y o f t h e b e e t l e s t o use t h e s e d i f f e r e n t mechanisms t o s e a r c h o u t and a t t a c k h o s t s under c o n t i n u o u s l y c h a n g i n g c o n d i t i o n s i n n a t u r a l f o r e s t s e t t i n g s , a l l o w t h e b e e t l e s t o o p t i m i z e t h e i r s e a r c h f o r a r a r e h o s t . New C o n s i d e r a t i o n s i n P e s t Management The r e s u l t s from my e x p e r i m e n t s can a i d i n i m p r o v i n g management o f t h i s p e s t a t t h e l o g s o r t s and s t o r a g e a r e a s , as w e l l as i n t h e commercial f o r e s t s e t t i n g s . C u r r e n t l y , m a s s - t r a p p i n g programs d e p l o y s e m i o c h e m i c a l - b a i t e d f u n n e l t r a p s a l o n g t h e f o r e s t margin s u r r o u n d i n g open a r e a s c o n t a i n i n g s u s c e p t i b l e l o g s . The t r a p s i n most c a s e s a r e p l a c e d no f u r t h e r t h a n 10 m i n t o t h e margins (Phero Tech. T e c h n i c a l B u l l e t i n No. 84001). I s u g g e s t p l a c i n g t r a p s up t o 50 - 100 m i n t o t h e f o r e s t , i f enough f o r e s t a r e a i s p r e s e n t . S i n c e b e e t l e s a r e c a p t u r e d a t h i g h e r f r e q u e n c i e s i n s t i l l a i r c o n d i t i o n s o f a f o r e s t , t h a n i n open a r e a s , t h e r e i s an o p p o r t u n i t y t o a t t r a c t b e e t l e s away from t h e 168 l o g s s e t i n t h e open a r e a s , i n s t e a d o f j u s t t r y i n g t o s e t up a b a r r i e r w h i c h w i l l a t t r a c t b e e t l e s i n t h e d i r e c t i o n o f t h e l o g s . I n c ommercial f o r e s t s , where l o g i n v e n t o r i e s c o u l d n ot be removed from t h e s e t t i n g s b e f o r e mass f l i g h t o f T. l i n e a t u m i n t h e s p r i n g , l o g s s h o u l d be moved from t h e edges o f t h e open s e t t i n g and p i l e d o r s t a c k e d n e a r i t s c e n t e r . T h i s c o u l d reduce t h e amount o f l o g s a t t a c k e d because o f t h e d i f f i c u l t i e s t h e b e e t l e s seem t o have i n o r i e n t i n g t o o l f a c t o r y s t i m u l i i n open a r e a s . F i e l d s t u d i e s need t o be c o n d u c t e d t o see i f m a s s - t r a p p i n g can work. K o n i g (1988) has shown t h a t b e e t l e numbers can be reduced i n t h e f o r e s t u s i n g t h i s t e c h n i q u e . I suggest placement o f s e m i o c h e m i c a l -b a i t e d f u n n e l t r a p s p l a c e d i n t h e f o r e s t , 0 - 100 m deep i n t o t h e f o r e s t , around a l o g g e d s e t t i n g t h a t was h e a v i l y a t t a c k e d by T. l i n e a t u m t h e p r e v i o u s season. P o p u l a t i o n movement out o f t h e f o r e s t t o a new s i t e can be impeded by c a p t u r i n g t h e s e b e e t l e s t h e f o l l o w i n g s p r i n g . G i v e n a c h o i c e , emerging b e e t l e s w i l l f l y i n t h e f o r e s t r a t h e r t h a n an open s e t t i n g , t h e r e f o r e t h e chance o f c a p t u r i n g them b e f o r e t h e y d i s p e r s e l o n g d i s t a n c e s t o new s i t e s i s good. Placement o f c u t b l o c k s i n a commercial f o r e s t can be m o d i f i e d t o m i n i m i z e b e e t l e a t t a c k o f h i g h q u a l i t y l o g s . The g e n e r a l t r e n d i n l o g g i n g o l d - g r o w t h f o r e s t s a l o n g t h e B.C. c o a s t i s t o c u t s t a n d s i n t h e l o w e r v a l l e y e l e v a t i o n s f i r s t , f o l l o w e d by h a r v e s t s a t h i g h e r e l e v a t i o n s . These l a t t e r c u t b l o c k s a r e v e r y s u s c e p t i b l e t o T. l i n e a t u m 169 a t t a c k , because u p - v a l l e y winds w i l l a i d i n d i s p e r s i n g b e e t l e s from o l d c u t b l o c k s t o new ones. I f new c u t b l o c k s can be p l a c e d upwind ( p r o v i d i n g t h e p r e s e n c e o f a p r e v a i l i n g wind) t o o l d c u t b l o c k s , t h e a b i l i t y o f t h e p o p u l a t i o n from t h e o l d c u t b l o c k s u c c e s s f u l l y f i n d i n g t h e new s i t e would be, i n my o p i n i o n , s e v e r e l y r e s t r i c t e d . I n B.C., as second-growth f o r e s t s come i n t o use o r where r o a d networks a r e a l r e a d y e s t a b l i s h e d , placement o f c u t b l o c k s upwind t o p r e v i o u s s e t t i n g s may be become more p l a u s i b l e . I n t h e U n i t e d S t a t e s , where second growth f o r e s t r y now dom i n a t e s , t h i s approach can p r e s e n t l y be c o n s i d e r e d on a l a r g e r s c a l e . W h i l e improvements have and can be made to w a r d managing T. l i n e a t u m p o p u l a t i o n s , more i n f o r m a t i o n on d i s p e r s a l , e s p e c i a l l y d u r i n g t h e f a l l s eason, can p r o v i d e a means t o d e v e l o p a comprehensive p e s t management s t r a t e g y e x t e n d i n g from t h e f o r e s t s e t t i n g s i n t o t h e l o g s o r t and s t o r a g e a r e a s . Such a program c o u l d save t h e B r i t i s h Columbia F o r e s t I n d u s t r y m i l l i o n s o f d o l l a r s each y e a r . 170 REFERENCES A n n i l a , E., A. Bakke, B. B e j e r - P e t e r s o n , B. Lekander. 1972. F l i g h t p e r i o d and b r o o d emergence i n Trypodendron  l i n e a t u m ( O l i v . ) ( C o l . , S c o l y t i d a e ) i n t h e N o r d i c c o u n t r i e s . Comm. I n s t . F o r . Fenn. 76:7-28. A n g e r i l l i , N. and J.A. McLean. 1984. W i n d t u n n e l and f i e l d o b s e r v a t i o n s o f w e s t e r n s p r u c e budworm r e s p o n s e s t o pheromone-baited t r a p s . J . Ent. Soc. B.C. 81:10-16. A t k i n s , M.D. 19 66a. L a b o r a t o r y s t u d i e s on t h e b e h a v i o u r o f t h e D o u g l a s - f i r b e e t l e , Dendroctonus pseudotsugae Hopk. Can. E n t . 98:953-991. A t k i n s , M.D. 1966b. B e h a v i o r a l v a r i a t i o n among s c o l y t i d s i n r e l a t i o n t o t h e i r h a b i t a t . Can. E n t . 98: 285-288. A t k i n s , M.D. 1969. L i p i d l o s s w i t h f l i g h t i n t h e D o u g l a s -f i r b e e t l e . Can. E n t . 101:164-165. Bak e r , T.C. and L.P.S. Kuenen. 1982. Pheromone s o u r c e l o c a t i o n i n f l y i n g moths: A supplementary non-a n e m o t a c t i c mechanism. S c i e n c e . 216:424-427. Baker, T.C. and C.E. L i n n , J r . 1984. Wind t u n n e l s i n pheromone r e s e a r c h , pp. 75-110. I n Hummel, H.E. and T.A. M i l l e r ( E d s . ) . Techniques i n Pheromone R e s e a r c h . S p r i n g e r - V e r l a g , New York. 464 pp. Banks, H.T. P.M. K a r e i v a , and L. Z i a . 1988. A n a l y z i n g f i e l d s t u d i e s o f i n s e c t d i s p e r s a l u s i n g t w o - d i m e n s i o n a l t r a n s p o r t e q u a t i o n s . E n v i r o n . Entomol. 17:815-82 0. B a r r y , R.G. 1981. Mountain Weather and C l i m a t e . Metheun, London. 313 pp. B a t s c h e l e t , E. 1981. C i r c u l a r S t a t i s t i c s i n B i o l o g y . Academic P r e s s . 371 pp. Bauer, J . and J.P. V i t e . 1975. Host s e l e c t i o n by Trypodendron l i n e a t u m . N a t u r w i s s . 62: 539. B e l l , W.J. and R.T. Carde ( E d s . ) . 1984. C h e m i c a l E c o l o g y o f I n s e c t s . Chapman & H a l l , London. 524 pp. B e n n e t t , R.B. and J.H. Borden. 1971. F l i g h t a r r e s t m e n t o f t e t h e r e d Dendroctonus pseudotsugae and Trypodendron  l i n e a t u m ( C o l e o p t e r a : S c o l y t i d a e ) i n r e s p o n s e t o o l f a c t o r y s t i m u l i . Ann. Entomol. Soc. Am. 64:1273-1286. 171 Berryman, A.A. and L. S a f r a n y i k ( E d s . ) * 1980. P r o c . IUFRO Conf., D i s p e r s a l o f F o r e s t I n s e c t s : E v a l u a t i o n , Theory and Management I m p l i c a t i o n s . Idaho. 278 pp. B i r c h , M.C. and P.R. White. 1988. Responses o f f l y i n g male Anobium punctatum ( C o l e o p t e r a : A n o b i i d a e ) t o female sex pheromone i n a w ind t u n n e l . J . I n s . Behav. 1:111-115. Borden, J.H. 1977. B e h a v i o r a l r e s p o n s e s o f C o l e o p t e r a t o pheromones, a l l o m o n e s , and kairomones. pp. 169-198. I n Shorey, H.H. and J . J . McKelevey, J r . ( E d s . ) . C h e m i c a l C o n t r o l o f I n s e c t B e h a v i o r : Theory and A p p l i c a t i o n . John W i l e y and Sons, I n c . Borden, J.H. 1984. S e m i o c h e m i c a l - m e d i a t e d a g g r e g a t i o n and d i s p e r s i o n i n t h e C o l e o p t e r a . pp. 123-149. I n L e w i s , T. (Ed.) I n s e c t Communication. Academic P r e s s , London. Borden, J.H. 1985. A g g r e g a t i o n pheromones. pp. 257-285. I n B e h a v i o r . K e r k u t , G.A. ( E d . ) . V o l 9. I n Comprehensive I n s e c t P h y s i o l o g y , B i o c h e m i s t r y , and Pharmacology. K e r k u t , G.A. and L . I . G i l b e r t ( E d s . ) . Pergamon P r e s s , O x f o r d . Borden, J.H. 1988a. The s t r i p e d a m b r o s i a b e e t l e , Trypodendron l i n e a t u m ( O l i v i e r ) pp. 579-596. I n Berryman, A.A. ( E d . ) . Dynamics o f F o r e s t I n s e c t P o p u l a t i o n s . Plenum P u b l i s h i n g Co. 603 pp. Borden, J.H. 1988b. Use o f s e m i o c h e m i c a l s t o manage c o n i f e r o u s t r e e p e s t s i n w e s t e r n Canada. I n Ridgeway, R., R.M. S i l v e r s t e i n , and M. I n s c o e ( E d s . ) . P r a c t i c a l A p p l i c a t i o n s o f I n s e c t Pheromones and O t h e r A t t r a c t a n t s . ( i n p r e s s ) . Borden, J.H. and J.A. McLean. 1981. Pheromone-based s u p p r e s s i o n o f a mbrosia b e e t l e s i n i n d u s t r i a l t i m b e r p r o c e s s i n g a r e a s , pp. 133-154. I n M i t c h e l l , E.R. (ED.). Management o f I n s e c t P e s t s W i t h S e m i o c h e m i c a l s : Concepts and P r a c t i c e . Plenum P r e s s , New York. 514 pp. Borden, J.H. and C.E. S l a t e r . 1969. Sex pheromone o f Trypodendron l i n e a t u m : p r o d u c t i o n i n t h e f e m a l e h i n d g u t - M a l p i g h i a n t u b u l e r e g i o n . Ann. Entomol. Soc. Am. 62: 454-455. Borden, J.H., L. Chong, J.A. McLean, K.N. S l e s s o r , and K. M o r i . 1976. G n a t h o t r i c h u s s u l c a t u s : s y n e r g i s t i c e n a n t i o m e r s o f t h e a g g r e g a t i o n pheromone s u l c a t o l . S c i e n c e 192:894-896. 172 Borden, J.H., L. Chong, K.N. S l e s s o e , A.C. O e h l s c h l a g e r , H.D. P i e r c e , J r . and B.S. L i n d g r e n . 1981. A l l e l o c h e m i c a c t i v i t y o f a g g r e g a t i o n pheromones between t h r e e s y m p a t r i c s p e c i e s o f ambrosia b e e t l e s ( C o l e o p t e r a : S c o l y t i d a e ) . Can. E n t . 113:557-563. Borden, J.H., J.R. Handley, B.D. J o h n s t o n , J.G. MacConnel, R.M. S i l v e r s t e i n , K.N. S l e s s o r , A.A. Swi g a r , and D.T.W. Wong. 1979. S y n t h e s i s and f i e l d t e s t i n g o f 4,6,6-l i n e a t i n , t h e a g g r e g a t i o n pheromone o f Trypodendron  l i n e a t u m ( C o l e o p t e r a : S c o l y t i d a e ) . J . Chern. E c o l . 5:681-689. Borden, J.H., J.R. Handley, J.A. McLean, R.M. S i l v e r s t e i n , L. Chong, K.N. S l e s s o r , B.D. J o h n s t o n , H.R. S c h u l e r . 1980. Enantiomer-based s p e c i f i c i t y i n pheromone communication by two s y m p a t r i c G n a t h o t r i c h u s s p e c i e s ( C o l e o p t e r a : S c o l y t i d a e ) . J . Chern. E c o l . 6:445-456. Borden, J.H., D.W.A. Hunt, D.R. M i l l e r , and K.N. S l e s s o r . 1986. O r i e n t a t i o n i n f o r e s t C o l e o p t e r a : an u n c e r t a i n outcome o f r e s p o n s e s by i n d i v i d u a l b e e t l e s t o v a r i a b l e s t i m u l i . pp. 97-110. I n Payne, T.L., M.C. B i r c h , and C.E.J. Kennedy ( E d s . ) . Mechanisms i n I n s e c t O l f a c t i o n . C l a r e n d o n P r e s s , O x f o r d . 3 64 pp. Borden, J.H., C.J. K i n g , S. L i n d g r e n , L. Chong, D.R. Gray, A.C. O e h l s c h l a g e r , K.N. S l e s s o r , and H.D. P i e r c e , J r . 1982. V a r i a t i o n i n r e sponse o f Trypodendron l i n e a t u m from two c o n t i n e n t s t o s e m i o c h e m i c a l s and t r a p form. E n v i r o n . Entomol. 11:403-408. B o t t e r w e g , P.F. 1982. D i s p e r s a l and f l i g h t b e h a v i o u r o f t h e s p r u c e b a r k b e e t l e I p s t y p o g r a p h u s i n r e l a t i o n t o sex, s i z e , and f a t c o n t e n t . Z. ang. E n t . 94:466-489. B y e r s , J.A. 1988. Upwind f l i g h t o r i e n t a t i o n t o pheromone i n w e s t e r n p i n e b e e t l e t e s t e d w i t h r o t a t i n g windvane t r a p s . J . Chern. E c o l . 14:189-198. Carde, R.T. 1984. C h e m o - o r i e n t a t i o n i n f l y i n g i n s e c t s , pp. 111-124. I n B e l l , W.J. and R.T. Carde ( E d s . ) . C h e m i c a l E c o l o g y o f I n s e c t s . S i n a u e r A s s o c . , I n c . M a s s a c h u s e t t s . Chapman, J.A. 1956. F l i g h t muscle changes d u r i n g a d u l t l i f e i n a s c o l y t i d b e e t l e . N a t u r e 177:1183. Chapman, J.A. 1962. F i e l d s t u d i e s on a t t a c k f l i g h t and l o g s e l e c t i o n by t h e ambrosia b e e t l e Trypodendron l i n e a t u m ( O l i v . ) ( C o l e o p t e r a : S c o l y t i d a e ) . Can. E n t . 94:74-92. Chapman, J.A. and J.M. K i n g h o r n . 1958. S t u d i e s o f f l i g h t and a t t a c k a c t i v i t y o f t h e ambrosia b e e t l e , 173 Trypodendron l i n e a t u m ( O l i v . ) , and o t h e r s c o l y t i d s . Can. E n t . 90:362-372. Chapman, J.A. and W.W. N i j h o l t . 1980. Time and a t t a c k f l i g h t o f Ambrosia b e e t l e Trypodendron l i n e a t u m ( C o l e o p t e r a : S c o l y t i d a e ) i n r e l a t i o n t o weather i n c o a s t a l B r i t i s h Columbia. Can. F o r . S e r v . I n f . Rept. BC-R-5. 2 3 pp. Choudhury, J.H. and J.S. Kennedy. 1980. L i g h t v e r s u s pheromone-bearing wind i n c o n t r o l o f f l i g h t d i r e c t i o n by b e e t l e s , S c o l y t u s m u l t i s t r i a t u s . P h y s i o l . Entomol. 5:207-214. Danthanarayana, W. ( E d . ) . 1986. I n s e c t F l i g h t : D i s p e r s a l and M i g r a t i o n . S p r i n g e r - V e r l a g , B e r l i n . 289 pp. Daust, D.K. 1985. The i n f l u e n c e o f s i t e c h a r a c t e r i s t i c s on l o g g i n g t e c h n i q u e s and ambrosia b e e t l e damage i n t h e Lake Cowichan A r e a . B.S.F. T h e s i s . U n i v . o f B r i t i s h C o lumbia. 69 pp. D e l u c c h i , V. and W. B a l t e n s w e i l e r ( E d s . ) . 1979. P r o c . IUFRO Conf., D i s p e r s a l o f F o r e s t I n s e c t s : E v a l u a t i o n , Theory and Management I m p l i c a t i o n s . S w i t z e r l a n d . 342 pp. D i c k e n s , J.C. 1981. B e h a v i o u r a l and e l c t r o p h y s i o l o g i c a l r e s p o n s e s o f t h e b a r k b e e t l e , I p s ty p o g r a p h u s t o p o t e n t i a l pheromone components. P h y s i o l . Entomol. 6:251-261. D i n g l e , H. 1985. M i g r a t i o n , pp. 375-414. I n B e h a v i o u r . K e r k u t , G.A. ( E d . ) . V o l . 9. I n Comprehensive I n s e c t P h y s i o l o g y , B i o c h e m i s t r y , and Pharmacology. K e r k u t , G.A. and L . I . G i l b e r t ( E d s . ) . Pergamon P r e s s , O x f o r d . Dyer, E.D.A. 1961. F l i g h t c a p a b i l i t y o f a m b r o s i a b e e t l e (Trypodendron). Can. Dep. A g r i c . F o r . B i o l . D i v . B i -Mon. P r o g . Rep. 17 ( 1 ) : 4 . Dyer, E.D.A. and J.A. Chapman. 1965. F l i g h t and a t t a c k o f t h e a m b r o s i a b e e t l e , Trypodendron l i n e a t u m ( O l i v . ) i n r e l a t i o n t o f e l l i n g d a t e s o f l o g s . Can. E n t . 97:42-57. Dyer, E.D.A. and J.M. K i n g h o r n . 1961. F a c t o r s i n f l u e n c i n g t h e d i s t r i b u t i o n o f o v e r w i n t e r i n g a m b r o s i a b e e t l e s , Trypodendron l i n e a t u m ( O l i v . ) . Can. E n t . 93:746-759. Edson, L . J . 1978. Host c o l o n i z a t i o n and t h e a r r i v a l sequence o f t h e mountain p i n e b e e t l e and i t s i n s e c t a n a s s o c i a t e s . Ph.D. D i s s e r t a t i o n , U n i v e r s i t y o f C a l i f o r n i a , B e r k e l e y . 196 pp. 174 Edwards, D.K. 1960. E f f e c t s o f e x p e r i m e n t a l l y a l t e r e d u n i p o l a r a i r - i o n d e n s i t y upon t h e amount o f a c t i v i t y o f t h e b l o w f l y , C a l l i p h o r a v i c i n a R.D. Can. J . Z o o l . 38:1079-1091. F a r k a s , S.R. and H.H. Shorey. 1972. Che m i c a l t r a i l -f o l l o w i n g by f l y i n g i n s e c t s : A mechanism f o r o r i e n t a t i o n t o a d i s t a n t odor s o u r c e . S c i e n c e . 178:67-68. F a r k a s , S.R. and H.H. Shorey. 1974. Mechanisms o f o r i e n t a t i o n t o a d i s t a n t pheromone s o u r c e . I n B i r c h , M.C. ( E d . ) . Pheromones. N o r t h - H o l l a n d P u b l . Co. 495 pp. F a r e s , Y., P.J.H. Sharpe, C.E. Magnuson. 1980. Pheromone d i s p e r s i o n i n f o r e s t s . J . Theor. B i o l . 84:335-359. F i s h e r , R.C., G.H. Thompson, and W.E. Webb. 1953. Ambrosia b e e t l e s i n f o r e s t and s a w m i l l : t h e i r b i o l o g y , economic i m p o r t a n c e and c o n t r o l . P a r t 1. F o r . A b s t r . 14:381-389. F o c k l e r , C.E. and J.H. Borden. 1972. S e x u a l b e h a v i o r and s e a s o n a l m a t i n g a c t i v i t y o f Trypodendron l i n e a t u m ( C o l e o p t e r a : S c o l y t i d a e ) . Can. E n t . 104:1841-1853. Fox, D.J. and K.E. G u i r e . 1976. Documentation f o r M i d a s . S t a t . Res. Lab., U n i v e r s i t y o f M i c h i g a n . 2 03 pp. F r a n c i a , F.C. and K. Graham. 1967. A s p e c t s o f o r i e n t a t i o n b e h a v i o r i n t h e ambrosia b e e t l e Trypodendron l i n e a t u m ( O l i v i e r ) . Can. J . Z o o l . 45:985-1002. Francke-Grossman, H. 1963. Some new a s p e c t s i n f o r e s t entomology. Ann. Rev. Entomol. 8:415-438. Funk, A. 1965. The s y m b i o t i c fung o f c e r t a i n a m b r o s i a b e e t l e s i n B r i t i s h Columbia. Can J . Bot. 43:929-932. F u r n i s s , M.M. and R.L. F u r n i s s . 1972. S c o l y t i d s ( C o l e o p t e r a ) on s n o w - f i e l d s above t i m b e r l i n e i n Oregon and Washington. Can. E n t . 104:1471-1478. Gara, R . I . 1963. S t u d i e s on t h e f l i g h t b e h a v i o r o f I p s c o n f u s u s (Lec.) ( C o l e o p t e r a : S c o l y t i d a e ) i n r e s p o n s e t o a t t r a c t i v e m a t e r i a l . C o n t r i b . Boyce Thompson I n s t . 21:51-66. Graham, K. 1959. R e l e a s e by f l i g h t e x e r c i s e o f t h e c h e m o t r o p i c response from p h o t o p o s i t i v e d o m i n a t i o n i n a s c o l y t i d b e e t l e . N a t u r e . 184:284-85. 175 Gray, D.R. and J.H. Borden. 1985. Ambrosia b e e t l e a t t a c k on l o g s b e f o r e and a f t e r p r o c e s s i n g t h r o u g h a d r y l a n d s o r t i n g a r e a . F o r . Chron. 61:299-302. Haufe, W.O. 1954. The e f f e c t s o f a t m o s p h e r i c p r e s s u r e on t h e f l i g h t r e s p o n s e s o f Aedes a e g y p t i (L.) B u l l . E ntomol. Res. 45:507-526. H e l l a n d , I.S., J.M. H o f f , and O. A n d e r b r a n t . 1984. A t t r a c t i o n o f b a r k b e e t l e s ( C o l e o p t e r a : S c o l y t i d a e ) t o a pheromone t r a p : e x p e r i m e n t and m a t h e m a t i c a l models. J . Chem. E c o l . 10:723-752. Henson, W.R. 1962. L a b o r a t o r y s t u d i e s on t h e a d u l t b e h a v i o r o f Conopthorus c o n i p e r d a ( C o l . : S c o l y t . ) . I I I . F l i g h t . Ann. Entomol. Soc. Am. 55:524-530. Johnson, C.G. 1969. M i g r a t i o n and D i s p e r s a l o f I n s e c t s by F l i g h t . Metheun, London. 763 pp. Kennedy, J.S. 1975. I n s e c t d i s p e r s a l . pp. 103-119. I n P i m e n t a l , D. ( E d . ) . I n s e c t s , S c i e n c e and S o c i e t y . Academic P r e s s , New York. Kennedy, J.S. 1978. The c o n c e p t s o f o l f a c t o r y a r r e s t m e n t and a t t r a c t i o n . P h y s i o l . Entomol. 3:91-98. Kennedy, J.S. 1986. Some c u r r e n t i s s u e s i n o r i e n t a t i o n t o odour s o u r c e s . pp. 11-25. I n Payne, T.L., M.C. B i r c h , and C.E.J. Kennedy ( E d s . ) . Mechanisms i n I n s e c t O l f a c t i o n . C l a r e n d o n P r e s s , O x f o r d . Kimmins, J.P. 1987. F o r e s t E c o l o g y . M a c M i l l a n P u b l i s h i n g Co. New York. 531 pp. K i n g h o r n , J.M. and J.A. Chapman. 1959. The o v e r w i n t e r i n g o f t h e ambrosia b e e t l e Trypodendron l i n e a t u m ( O l i v . ) F o r . S c i . 5:81-92. Kleinbaum, D.G. and L.L. Kupper. 1978. A p p l i e d R e g r e s s i o n A n a l y s i s and Other M u l t i v a r i a b l e Methods. Duxbury P r e s s , C a l i f o r n i a . 486 pp. K o n i g , E. 1988. M a s s - t r a p p i n g o f Trypodendron l i n e a t u m 01. ( C o l . , S c o l y t i d a e ) : e f f e c t o f i n f e s t a t i o n o f c u t t i m b e r t e m p o r a r i l y l e f t i n t h e f o r e s t . Z. ang. E n t . 106:262-265. K y d o n i e u s , A.F. and M. Be r o z a . 1981. The hereo n d i s p e n s e r f o r m u l a t i o n and r e c e n t t e s t r e s u l t s . pp. 445-454. I n M i t c h e l l , E.R. ( E d . ) . Management o f I n s e c t P e s t s W i t h S e m i o c h e m i c a l s : Concepts and P r a c t i c e . Plenum P r e s s , New York. 514 pp. 176 L i n d g r e n , B.S. 1983. A m u l t i p l e f u n n e l t r a p f o r s c o l y t i d b e e t l e s ( C o l e o p t e r a ) . Can. E n t . 115:299-302. L i n d g r e n , B.S. and J.H. Borden. 1983. Survey and mass t r a p p i n g o f ambrosia b e e t l e s ( C o l e o p t e r a : S c o l y t i d a e ) i n t i m b e r p r o c e s s i n g a r e a s on Vancouver I s l a n d . Can. J . F o r . Res. 13:481-493. L i n d g r e n , B.S., J.H. Borden, L.Chong, L.M. F r i s k i e , D.B. O r r . 1983. F a c t o r s i n f l u e n c i n g t h e e f f i c i e n c y o f pheromone-baited t r a p s f o r t h r e e s p e c i e s o f a m b r o s i a b e e t l e s ( C o l e o p t e r a : S c o l y t i d a e ) . Can. E n t . 115:303-313. L i n t o n , D.A., L. S a f r a n y i k , L.H. McMullen, and R. B e t t s . 1987. F i e l d t e c h n i q u e s f o r r e a r i n g and m a r k i n g mountain p i n e b e e t l e f o r use i n d i s p e r s a l s t u d i e s . J . E n t . Soc. B.C. 84:53-57. M a c C o n n e l l , J.G., J.H. Borden, R.M. S i l v e r s t e i n , and E. S t o k k i n k . 1977. I s o l a t i o n and t e n t a t i v e i d e n t i f i c a t i o n o f l i n e a t i n , a pheromone from t h e f r a s s o f Trypodendron l i n e a t u m ( C o l e o p t e r a : S c o l y t i d a e ) . J . Chern. E c o l . 5:549-561. McLean, J.A. 1985. Ambrosia b e e t l e s : a m u l t i m i l l i o n d o l l a r degrade problem o f sawlogs i n c o a s t a l B r i t i s h C olumbia. F o r . Chron. 61:295-298. McLean, J.H. and J.H. Borden. 1977. A t t a c k by G n a t h o t h r i c h u s s u l c a t u s ( C o l e o p t e r a : S c o l y t i d a e ) on stumps and f e l l e d t r e e s b a i t e d w i t h s u l c a t o l and e t h a n o l . Can. E n t . 109:675-686. McLean, J.A., A. Bakke, H. Niemeyer. 1987. An e v a l u a t i o n o f t h r e e t r a p s and two l u r e s f o r t h e a m b r o s i a b e e t l e Trypodendron l i n e a t u m ( O l i v . ) ( C o l e o p t e r a : S c o l y t i d a e ) i n Canada, Norway, and West Germany. Can. E n t . 119:273-280. Mc M u l l e n , L.H. and M.D. A t k i n s . 1962. On t h e f l i g h t and h o s t s e l e c t i o n o f t h e D o u g l a s - f i r b e e t l e , Dendroctonus  pseudotsucfae Hopk. ( C o l e o p t e r a : S c o l y t i d a e ) . Can. E n t . 94: 1309-1325. M a t h e r s , W.G. 1935. Time o f f e l l i n g i n r e l a t i o n t o i n j u r y from a m b r o s i a b e e t l e s , o r pinworms. B.C. Lumberman. 19:14. M i z e l l , R.F. I l l , J . L . F r a z i e r , and T.E. Nebeker. 1984. Response o f t h e c l e r i d p r e d a t o r Thanasimus d u b i u s ( F . ) , t o b a r k b e e t l e pheromones and t r e e v o l a t i l e s i n a wind t u n n e l . J . Chem. E c o l . 10:177-187. 1.77 Moeck, H.A. 1970. Eth a n o l as the primary a t t r a c t a n t f o r the ambrosia b e e t l e Trypodendron li n e a t u m ( C o l e o p t e r a : S c o l y t i d a e ) . Can. Ent. 102:985-995. Moeck, H.A. 1971. F i e l d t e s t o f e t h a n o l as a s c o l y t i d a t t r a c t a n t . Can. Dept. F i s h , and For. Bi-mon. Res. Notes. 27 (2):11-12 N i j h o l t , W.W. 1967. Mo i s t u r e and f a t content d u r i n g the a d u l t l i f e o f the ambrosia b e e t l e , Trypodendron  lin e a t u m ( O l i v . ) J . Ento Soc. B.C. 64:51-55. N i j h o l t , W.W. 1979. The s t r i p e d ambrosia b e e t l e , Trypodendron lineatum ( O l i v i e r ) : an annotated b i b l i o g r a p h y . Can. For. Serv. I n f . Rept. BC-X-121. N i j h o l t , W.W. and J . Schonherr. 1976. Chemical response b e h a v i o r of s c o l y t i d s i n West Germany and western Canada. Can. For. Serv. Bi-mon. Res. Notes. 32:31-32. N i l s s e n , A.C. 1984. Long-range a e r i a l d i s p e r s a l o f bark b e e t l e s and bark w e e v i l s (Coleoptera, S c o l y t i d a e and C u r c u l i o n i d a e ) i n no r t h e r n F i n l a n d . Ann. Ent. Fenn. 50:37-42. Pa i v a , M.R. and K. K i e s e l . 1985. F i e l d responses of Trypodendron spp. ( C o l . , S c o l y t i d a e ) t o d i f f e r e n t c o n c e n t r a t i o n s o f l i n e a t i n and -pinene. Z. ang. Ent. 99:442-448. Payne, T.L. M.C. B i r c h , and C.E.J. Kennedy. (Eds.). 1986. Mechanisms i n I n s e c t O l f a c t i o n . Clarendon Press, Oxford. 3 64 pp. Pedgley, D. 1982. Windborne Pest and Diseases: Meteorology of a i r b o r n e organisms. E. Horwood L t d . England. 250 pp. Phelan, P.L. and J.R. M i l l e r . 1982. P o s t - l a n d i n g b e h a v i o r of a l a t e Myzus p e r s i c a e as a l t e r e d by ( E ) - B — f a r n e s e n e and t h r e e c a r b o x y l i c a c i d s . Ent. Exp. Ap p l . 32:46-53. Pitman, G.B. and J.P. V i t e . 1969. Aggregation b e h a v i o r of Dendroctonus ponderosae (C o l e o p t e r a : S c o l y t i d a e ) i n response t o chemical messengers. Can. Ent. 101:143-149. P o j a r , J . 1983. F o r e s t Ecology. pp. 221-318. In Watts, S.B. (Ed.). F o r e s t r y Handbook f o r B r i t i s h Columbia. For. Undergrad. S o c , Vancouver, B.C. 611 pp. Prebble, M.L. and K. Graham. 1957. S t u d i e s o f a t t a c k by ambrosia b e e t l e s i n softwood l o g s on Vancouver I s l a n d , B r i t i s h Columbia. For. S c i . 3:9.0-112. 178 P r e i s s , R. and E. Kramer. 1986. Pheromone i n d u c e d anemotaxis i n s i m u l a t e d f r e e f l i g h t . I n Payne, T.L., M.C. B i r c h , and C.E.J. Kennedy ( E d s . ) . Mechanisms i n I n s e c t O l f a c t i o n . C l a r e n d o n P r e s s , O x f o r d . 364 pp. Rabb, R.L. and G.G. Kennedy ( E d s . ) . 1979. Movement o f H i g h l y M o b i l e I n s e c t s : Concepts and Methodology i n R e s e a r c h . N o r t h C a r o l i n a S t . U n i v . P r e e . 456 pp. R u d i n s k y , J.A. and G.E. Daterman. 1964. F i e l d s t u d i e s on f l i g h t p a t t e r n s and o l f a c t o r y r e s p o n s e s o f a m b r o s i a b e e t l e s on D o u g l a s - f i r f o r e s t s o f w e s t e r n Oregon. Can. E n t . 96:1339-1352. Salom, S.M. and J.A. McLean. 1988. S e m i o c h e m i c a l s f o r c a p t u r i n g t h e ambrosia b e e t l e , Trypodendron l i n e a t u m , i n m u l t i p l e f u n n e l t r a p s i n B r i t i s h Columbia. J . E n t . Soc. B.C. 85:34-39. Sanders, C.J. 1985. F l i g h t speed o f male s p r u c e budworm moths i n a w ind t u n n e l a t d i f f e r e n t wind speeds and a t d i f f e r e n t d i s t a n c e s from a pheromone s o u r c e . P h y s i o l . Entomol. 10:83-88. SAS I n s t i t u t e . 1985. SAS u s e r ' s g u i d e : S t a t i s t i c s . SAS I n s t i t u t e , Cary N.C. 956 pp. S c h l y t e r , F., J . L o f q v i s t , and J.A. B y e r s . 1987a. The b e h a v i o r a l sequence i n a t t r a c t i o n t o pheromone s o u r c e s i n t h e b a r k b e e t l e I p s typocrraphus. P h y s i o l . Entomol. 12:185-196. S c h l y t e r , F., J.A. B y e r s , and J . L o f q v i s t . 1987b. A t t r a c t i o n t o pheromone s o u r c e s o f d i f f e r e n t q u a n t i t y , q u a l i t y , and s p a c i n g : D e n s i t y - r e g u l a t i o n mechanisms i n b a r k b e e t l e I p s t y p o q r a p h u s. J . Chern. E c o l . 13:1503-1523. S c h m i t z , R.F. 1979. D i s p e r s a l o f p i n e e n g r a v e r b e e t l e s i n second growth ponderosa p i n e f o r e s t s , pp. 41-50. I n Berryman, A.a. and L. S a f r a n y i k ( E d s . ) . P r o c . IUFRO Conf., D i s p e r s a l o f F o r e s t I n s e c t s : E v a l u a t i o n , Theory and Management I m p l i c a t i o n s . Idaho Shepherd, R.F. 1966. F a c t o r s i n f l u e n c i n g t h e o r i e n t a t i o n and r a t e s o f a c t i v i t y o f Dendroctonus ponderosae H o p k i n s ( C o l e o p t e r a : S c o l y t i d a e ) . Can. E n t . 98:507-518. Shore, T.L. and J.A. McLean. 1983. A f u r t h e r e v a l u a t i o n o f t h e i n t e r a c t i o n s between t h e pheromones and two h o s t kairomones o f t h e a mbrosia b e e t l e s Trypodendron 179 l i n e a t u m and G n a t h o t r i c h u s s u l c a t u s ( C o l e o p t e r a : S c o l y t i d a e ) . Can. E n t . 115:1-5. Shore, T.L. and J.A. McLean. 1984. The e f f e c t o f h e i g h t o f t h e pheromone-baited t r a p s on c a t c h e s o f t h e a m b r o s i a b e e t l e , Trypodendron l i n e a t u m . J . E n t . Soc. B.C. 81:17-18. Shore, T.L. and J.A. McLean. 1985. A s u r v e y f o r t h e am b r o s i a b e e t l e s Trypodendron l i n e a t u m and G n a t h o t r i c h u s r e t u s u s ( C o l e o p t e r a : S c o l y t i d a e ) i n a s a w m i l l u s i n g pheromone-baited t r a p s . Can. E n t . 117:49-55. Shore, T.L. and J . A. McLean. 1988. The use o f mark-r e c a p t u r e t o e v a l u a t e a pheromone-based mass t r a p p i n g program f o r ambrosia b e e t l e s i n a s a w m i l l . Can. J . F o r . Res. 18:1113-1117. Shore, T.L., J.A. McLean, and J.C. Zanuncio. 1986. R e p r o d u c t i o n and s u r v i v a l o f t h e a m b r o s i a b e e t l e Trypodendron l i n e a t u m ( C o l e o p t e r a : S c o l y t i d a e ) i n D o u g l a s - f i r and w e s t e r n hemlock l o g s . Can. E n t . 119:131-139. Shorey, H.H. 1973. B e h a v i o r a l r e s p o n s e s t o i n s e c t pheromones. Ann. Rev. Entomol. 18:349-380. Southwood, T.R.E. 1962. M i g r a t i o n o f t e r r e s t i a l a r t h r o p o d s i n r e l a t i o n t o h a b i t a t . B i o l . Rev. 37:171-214. Southwood, T.R.E. 1978. E c o l o g i c a l Methods. Chapman and H a l l , London. 1978. 524 pp. S t i n n e r , R.E., C.S. B a r f i e l d , J . L . S t i m a c , and L. Dohse. 1983. D i s p e r s a l and movement o f i n s e c t p e s t s . Ann. Rev. Entomol. 1983. 28:319-335. T i l d e n , P.E., W.D. Bedard, K.Q. L i n d a h l , J r . and D.L. Wood. 1983. T r a p p i n g Dendroctonus b r e v i c o m i s : changes i n a t t r a c t a n t r e l e a s e r a t e , d i s p e r s i o n o f a t t r a c t a n t , and s i l h o u e t t e . J . Chern. E c o l . 9:311-321. Thompson, G.A. 1985. V e g e t a t i o n c l a s s i f i c a t i o n o f t h e Endowment Lands. G r e a t e r Vancouver R e g i o n a l D i s t r i c t P a r k s Department, Burnaby, B.C. Tech Paper No. 4. 115 pp. V i t e , J.P. and A. Bakke. 1979. Syn e r g i s m between c h e m i c a l and p h y s i c a l s t i m u l i i n h o s t c o l o n i z a t i o n by an ambr o s i a b e e t l e . N a t u r w i s s . 66:528-529. V i t e , J.P., R . I . Ga f a , and H.D. von S c h e l l e r . 1964. F i e l d o b s e r v a t i o n s on t h e re s p o n s e t o a t t r a c t a n t s o f b a r k 180 b e e t l e s i n f e s t i n g s o u t h e r n p i n e s . C o n t r i b . Boyce Thompson I n s t . 22:461-470. W e l l i n g t o n , W.G. 1980. D i s p e r s a l and p o p u l a t i o n change. pp. 11-24. I n Berryman, A.A. and L. S a f r a n y i k ( E d s . ) . P r o c . IUFRO Conf., D i s p e r s a l o f F o r e s t I n s e c t s : E v a l u a t i o n , Theory and Management I m p l i c a t i o n s . Idaho. W e l l i n g t o n , W.G. 1983. B i o m e t e o r o l o g y o f d i s p e r s a l . B u l l . E ntomol. Soc. Amer. 29:24-29. Wollerman, E.M. 1979. D i s p e r s i o n and i n v a s i o n by S c o l y t u s  m u l t i s t r i a t u s i n r e s p o n s e t o pheromone. E n v i r o n . Entomol. 8:1-5. Wood, D.L. 1982. The r o l e o f pheromones, kairomones, and a l l o m o n e s i n t h e h o s t s e l e c t i o n and c o l o n i z a t i o n b e h a v i o r o f b a r k b e e t l e s . Ann. Rev. Entomol. 27:411-446. Wood, S.L. 1982. The Bark and Ambrosia B e e t l e s o f N o r t h and C e n t r a l A m e r i c a : A Taxonomic Monograph. G r e a t B a s i n N a t u r a l i s t Memoirs, Number 6. Brigham Young U n i v e r s i t y . 1359 pp. Z a r , J.H. 1984. B i o s t a t i s t i c a l A n a l y s i s . P r e n t i c e - H a l l I n c . , New J e r s e y . 718 pp. 181 APPENDIX 1: RESPONSE OP TRYPODENDRON LINEATUM (OLIVIER) TO DIFFERENT SEMIOCHEMICAL-BAITED SUBSTRATES IN A WIND TUNNEL. A s t u d y was d e s i g n e d t o d e t e r m i n e t h e o p t i m a l s u b s t r a t e , and i t s o r i e n t a t i o n , f o r use i n c o n d u c t i n g d e t a i l e d b e h a v i o r a l o b s e r v a t i o n s o f Trypodendron l i n e a t u m ( O l i v i e r ) i n a wind t u n n e l . The s u b s t r a t e s t e s t e d i n c l u d e d a f u n n e l t r a p , a w e s t e r n hemlock l o g , and a s i m u l a t e d l o g . The d i m e n s i o n s o f t h e l a t t e r two s u b s t r a t e s a r e g i v e n i n Cha p t e r 2. Both t h e t r u e and s i m u l a t e d l o g were p l a c e d i n h o r i z o n t a l and v e r t i c a l p o s i t i o n s t o see i f e i t h e r p o s i t i o n was p r e f e r r e d by t h e b e e t l e s . E x p e r i m e n t a l p r o c e d u r e s such as b e e t l e p r e p a r a t i o n , r e l e a s e t r a y , d i s t a n c e t o s u b s t r a t e s , and s e m i o c h e m i c a l s a r e s i m i l a r t o t h o s e d e s c r i b e d i n C h a p t e r 2, s t u d y 1. One d i f f e r e n c e i s t h a t a wind speed o f 0.3 m/s was used h e r e . The e x p e r i m e n t was randomized complete b l o c k d e s i g n , w i t h t h e s i x e x p e r i m e n t a l days s e r v i n g as t h e b l o c k s . S e p a r a t e r e l e a s e s f o r males and fem a l e s o f 50 b e e t l e s each were c a r r i e d o u t f o r each t r e a t m e n t p e r b l o c k . F o l l o w i n g r e l e a s e , b e e t l e s were g i v e n 20 min t o respond. The d a t a o f i n t e r e s t f o r t h e f u n n e l t r a p t r e a t m e n t was t h e number o f b e e t l e s c a p t u r e d , and from t h e l o g t r e a t m e n t s , t h e number o f b e e t l e s l a n d i n g on them. The d a t a a r e p r e s e n t e d as p e r c e n t a g e s o f t h e number r e l e a s e d , and were s u b j e c t e d t o an a r c s i n e t r a n s f o r m a t i o n p r i o r t o a n a l y s i s (SAS 1985). 182 No s i g n i f i c a n t r e s p o n s e d i f f e r e n c e s were found between t h e s u b s t r a t e t r e a t m e n t s f o r b o t h males ( F 4 2 n = i * 0 1 ' P > 0.05) and f e males ( F 4 2Q = 3.34; p > 0.05) ( T a b l e 1 8 ) . No d i f f e r e n c e s i n r e s p o n s e s were found between sex (F^ 5 3 = 1.06; p > 0.05) e i t h e r . Based on t h e s e r e s u l t s , t h e e a s i e s t s u b s t r a t e t o use, t h e s i m u l a t e d l o g , was employed i n t h e d e t a i l e d b e h a v i o r w i n d t u n n e l s t u d i e s i n C h a p t e r 2. T h i s s u b s t r a t e was l i g h t w e i g h t and i t s l i g h t c o l o r a t i o n a l l o w e d f o r easy o b s e r v a t i o n o f b e e t l e s as t h e y l a n d e d on o r approached t h e s u b s t r a t e . The h o r i z o n t a l o r i e n t a t i o n o f t h e s u b s t r a t e was chosen, because i t s i m u l a t e d windthrown t i m b e r and l o g s n o r m a l l y a t t a c k e d i n t h e n a t u r a l s e t t i n g by a m b r o s i a b e e t l e s . 183 T a b l e 18. Mean r e c a p t u r e p e r c e n t a g e o f Trypodendron l i n e a t u m t o d i f f e r e n t s u b s t r a t e s b a i t e d w i t h s e m i o c h e m i c a l s i n a wind t u n n e l . S i x r e p l i c a t e s o f 50 b e e t l e s , f o r a t o t a l o f 300 b e e t l e s were r e l e a s e d f o r each t r e a t m e n t by sex c a t e g o r y . Treatment Mean % o f B e e t l e s R e c a p t u r e d + S.E. Males Females F u n n e l Trap 11.7 + 2.5 12.7 + 2.5 S i m u l a t e d Log ( H o r i z o n t a l ) 12.7 + 2.5 14.0 + 1.8 Hemlock Log ( H o r i z o n t a l ) 14.0 + 2.0 15. 0 + 1.8 S i m u l a t e d Log ( V e r t i c a l ) 14.7 + 2.8 17.0 + 3.0 Hemlock Log ( V e r t i c a l ) 13.3 + 1.6 15.0 + 2.7 No s i g n i f i c a n t d i f f e r e n c e s were found between t r e a t m e n t s and sex ( A n a l y s i s o f V a r i a n c e ; p > 0.05). 184 Appendix 2 : RESPONSE OF FLOWN AND UNFLOWN TRYPODENDRON LINEATUM TO SEMIOCHEMICAL-BAITED FUNNEL TRAPS IN A WIND TUNNEL. A.2 . 1 Introduction P h y s i o l o g i c a l r e a d i n e s s i s an e s s e n t i a l p r e r e q u i s i t e f o r d i s p e r s i n g s c o l y t i d b e e t l e s r e s p o n d i n g t o a t t r a c t i v e h o s t m a t e r i a l (Borden 1977). F o r t h e s t r i p e d a m b r o s i a b e e t l e , Trypodendron l i n e a t u m ( O l i v i e r ) , Graham (1959) found t h a t f l i g h t e x e r c i s e f o r s p r i n g emerging b e e t l e s was a n e c e s s a r y p r e r e q u i s i t e f o r s w i t c h i n g from an i n i t i a l p h o t o p o s i t i v e dominated b e h a v i o r t o t h e c h e m o t r o p i c b e h a v i o r used i n f i n d i n g h o s t m a t e r i a l . F l i g h t e x e r c i s e has s i n c e been documented as p a r t o f t h e d i s p e r s a l b e h a v i o r o f o t h e r s c o l y t i d b e e t l e s , such as Conopthorus c o n i p e r d a (Schw.) (Henson 1962), Dendroctonus ponderosae Hopkins (Shepherd 1966) , Dendroctonus pseudotsucrae Hopkins ( A t k i n s 1969) and S c o l y t u s m u l t i s t r i a t u s (Marsham) (Choudhury and Kennedy 1980). The amount o f f l i g h t e x e r c i s e needed v a r i e s between s p e c i e s as Be n n e t t and Borden (1971) found t h a t i t l a s t s an average o f 30 minutes f o r T. l i n e a t u m and 90 min u t e s f o r D. pseudotsucrae f o r t e t h e r e d b e e t l e s on a f l i g h t m i l l . F o r t h e s t u d i e s p r e s e n t e d i n t h i s t h e s i s , p r e v i o u s l y f l o w n b e e t l e s ( c a p t u r e d i n pheromone-baited f u n n e l t r a p s ) have been used, due t o t h e d i f f i c u l t y o f r e a r i n g enough b e e t l e s t o c a r r y o u t t h e e x p e r i m e n t s s a t i s f a c t o r i l y . T h i s , however, has posed a problem w i t h r e s p e c t t o i n t e r p r e t a t i o n o f e x p e r i m e n t a l r e s u l t s . P r e - f l o w n b e e t l e s may n o t 185 a c c u r a t e l y r e f l e c t b e h a v i o r o f a n a t u r a l l y emerging p o p u l a t i o n i n t h e f o r e s t . As a r e s u l t , a s t u d y was c a r r i e d o u t t o i n v e s t i g a t e d i f f e r e n c e s i n f l i g h t r e s p o n s e o f f l o w n and u n f l o w n T. l i n e a t u m t o s e m i o c h e m i c a l - b a i t e d f u n n e l t r a p s i n a wind t u n n e l . I n a d d i t i o n , t h e e f f e c t o f s t o r i n g f l o w n b e e t l e s was i n v e s t i g a t e d t o see i f t h e s e b e e t l e s needed a d d i t i o n a l f l i g h t e x e r c i s e b e f o r e r e s p o n d i n g t o s e m i o c h e m i c a l - b a i t e d t r a p s . A. 2 . 2 M a t e r i a l s a n d M e t h o d s Unflown b e e t l e s Western hemlock, Tsuqa h e t e r o p h y l l a (Raf.) S a r g . l o g s , f r e s h l y a t t a c k e d by T. l i n e a t u m a t t h e U n i v e r s i t y o f B r i t i s h C olumbia R e s e a r c h F o r e s t i n Maple R i d g e , B.C., were c u t and t h e n p l a c e d on t o p o f d u f f m a t e r i a l i n a 5.0 x 4.3 x 1.2 m canvas t e n t i n June o f 1986. The l o g s remained i n t h e t e n t u n t i l t h e f o l l o w i n g w i n t e r . B e e t l e s t h a t emerged from t h e l o g s i n t h e f a l l c o u l d n o t escape t h e t e n t and were f o r c e d t o e n t e r t h e d u f f m a t e r i a l c o v e r i n g t h e ground. The d u f f was c o l l e c t e d i n t h e w i n t e r and s t o r e d o u t s i d e i n p l a s t i c bags. As t e m p e r a t u r e s began t o warm up i n March, 1987, t h e bags were moved i n t o a w a l k - i n c o o l e r and h e l d a t 4 °C. To e x t r a c t t h e b e e t l e s , t h e s t o r e d d u f f m a t e r i a l was p u l l e d o u t o f t h e h o l d i n g bags and p l a c e d a t t h e bottom o f s e a l e d 60 x 38 x 27 cm c a r d b o a r d s e e d l i n g boxes. The l a y e r o f d u f f p l a c e d i n t h e boxes was l i m i t e d t o c a . 8 cm i n 186 t h i c k n e s s . Each box was f i t t e d a t one end w i t h a 3 00 ml mason j a r and a s m a l l s e c t i o n o f a c l o t h t o w e l i n s i d e o f i t . T h i s a l l o w e d l i g h t i n t o t h e box, i n d u c i n g warmed, p h o t o p o s i t i v e b e e t l e s t o make t h e i r way i n t o t h e j a r . Any f l i g h t t h a t would o c c u r i n t h e box was c o n s i d e r e d m i n i m a l . Opening o f some boxes showed t h a t b e e t l e s w a l k e d on t h e s i d e s o f t h e boxes and p r o b a b l y many wa l k e d i n t o t h e j a r s . However, s i n c e I d i d n o t o b s e r v e b e e t l e a c t i v i t y i n d e t a i l a t t h i s p o i n t , i t i s i m p o s s i b l e t o say how much a c t i v i t y d i d o c c u r . A t o t a l o f 21 - 3 0 boxes were used a t a t i m e . They were p l a c e d i n a b o i l e r room on t h e U.B.C. campus where t e m p e r a t u r e r e a c h e d c a . 2 5 °C. Emergence t o o k 2 4 days t o complete as 5 r e - f i l l e d ( w i t h d u f f ) s e t s o f boxes were used. J a r s were checked t w i c e d a i l y and emerging b e e t l e s were p l a c e d i n t h e w a l k - i n c o o l e r s t o r e d a t 4°C under a p h o t o p e r i o d o f 14:10 h (L:D). D e t a i l s o f t h e s t o r a g e p r o c e d u r e s used a r e d e s c r i b e d i n C h a p t e r 2. A t o t a l o f 1100 b e e t l e s were c o l l e c t e d . Flown b e e t l e s C o l l e c t i o n s o f f l o w n T. l i n e a t u m were made a t t h e b e g i n n i n g o f t h e i r mass f l i g h t p e r i o d i n A p r i l , 1987, u s i n g l i n e a t i n - b a i t e d m u l t i p l e f u n n e l t r a p s , p l a c e d a d j a c e n t t o a l a r g e l o g boom s t o r a g e a r e a a t F o r e s h o r e P a r k , U n i v e r s i t y o f B r i t i s h Columbia Endowment Lands, Vancouver, B.C. B e e t l e s 187 were c o l l e c t e d d a i l y and p l a c e d i n a w a l k - i n c o o l e r under t h e same c o n d i t i o n s as mentioned above. Flown t w i c e b e e t l e s Flown b e e t l e s were t a k e n from t h e c o o l e r and p l a c e d i n a 4 L p i c k l e j a r . The j a r was c o m p l e t e l y c o v e r e d w i t h b l a c k t a r p a per e x c e p t f o r t h e t o p o p e n i n g w h i c h was c o v e r e d t i g h t l y w i t h a f i n e mesh Lumite^- s c r e e n . T h i s p r o v i d e d t h e o n l y l i g h t s o u r c e f o r t h e b e e t l e s c a u s i n g them t o spend most o f t h e i r t i m e i n t h e j a r t r y i n g t o f l y t o and t h r o u g h t h e s c r e e n . F l i g h t e x e r c i s e p e r i o d s o f 3 0 m i n u t e s were t r i e d i n i t i a l l y , however c l o s e o b s e r v a t i o n showed t h a t t h e b e e t l e s were i n poor p h y s i c a l c o n d i t i o n a f t e r w a r d s . Thus, a 15 m inute f l i g h t e x e r c i s e p e r i o d was used i n t h e e x p e r i m e n t . A l t h o u g h B e n n e t t and Borden (1971) d e t e r m i n e d 30 m i n u t e s as t h e average f l i g h t e x e r c i s e t i m e needed f o r T. l i n e a t u m t o r e s p o n d t o o l f a c t o r y s t i m u l i , no d e s c r i p t i o n o f d i f f e r e n t t i m e i n t e r v a l s t e s t e d i s g i v e n . Thus, one must assume t h i s was an a r b i t r a r y p e r i o d o f t i m e used f o r t e s t i n g . They d i d r e p o r t , however, t h a t an i n s e c t w i t h 15 m i n u t e s o f f l i g h t e x e r c i s e e x h i b i t e d f l i g h t a r r e s t m e n t f o l l o w i n g r e s p o n s e t o a second exposure t o o l f a c t o r y s t i m u l i . T e s t p r o c e d u r e s The s t u d y was c o nducted i n t h e w ind t u n n e l d e s c r i b e d i n C h a p t e r 2. Two L i n d g r e n 8 - f u n n e l t r a p s were p l a c e d i n t h e |7 C hicopee M a n u f a c t u r i n g Co., C o r n e l i a , GA.. 30531. 188 upwind s e c t i o n o f t h e t u n n e l , 0.5 m from t h e upwind s c r e e n . B o t h t r a p s were b a i t e d w i t h s low r e l e a s e l i n e a t i n ( a g g r e g a t i o n pheromone) l u r e s . The r e l e a s e r a t e s o f t h e l u r e s a r e based on l u r e s i z e ( c a . 7.0 ug/day/cm 2 j l 2 - . Two s t a n d a r d l u r e s o f (25.8 cm 2 each) were s p l i t i n t o two, w i t h one h a l f o f a l u r e p l a c e d i n t h e t o p and t h e o t h e r h a l f i n t h e bottom o f each t r a p . T h e r e f o r e , t h e t o t a l r e l e a s e r a t e o f l i n e a t i n i n t h e wind t u n n e l was c a . 360 ug/24 h. E t h a n o l ( h o s t a t t r a c t a n t ) was a l s o used i n t h i s s t u d y and was r e l e a s e d a t a r a t e o f 75 mg/24 h from a s m a l l p l a s t i c v i a l hung 0.5 m from t h e c e i l i n g o f t h e t u n n e l and p l a c e d i n between b o t h f u n n e l t r a p s . F o r a l l groups o f b e e t l e s , a w a l k i n g t e s t was g i v e n p r i o r t o t h e ex p e r i m e n t t o make s u r e t h e b e e t l e s used were i n good p h y s i c a l c o n d i t i o n . B e e t l e s were r e l e a s e d from a t r a y 1.5 m downwind from t h e t r a p s . The t r a y has been p r e v i o u s l y d e s c r i b e d i n Chapter 2. A 3 x 2 x 2 f a c t o r i a l e x p e r i m e n t was s e t up as a c o m p l e t e l y randomized d e s i g n . Unflown, f l o w n , and f l o w n t w i c e t r e a t m e n t s were d i v i d e d by sex and by r e s p o n s e t o u n b a i t e d ( c o n t r o l ) and b a i t e d t r a p s . Thus, a t o t a l o f 12 t r e a t m e n t s were t e s t e d . The number o f b e e t l e s r e l e a s e d f o r each t r e a t m e n t ranged from 10 - 24. Lower numbers as w e l l as l e s s r e p l i c a t i o n s were used f o r t h e c o n t r o l t r e a t m e n t s because o f t h e s m a l l number o f u n f l o w n b e e t l e s a v a i l a b l e f o r t h e e x p e r i m e n t . F o r a l l t r e a t m e n t s , b e e t l e s were g i v e n a p e r i o d o f 15 minutes p r i o r t o r e l e a s e t o warm up from t h e i r 189 c o l d s t o r a g e p e r i o d . The a i r speed i n t h e t u n n e l was h e l d a t 0.15 m/s (Chapter 2 ) , and t h e t e m p e r a t u r e i n t h e t u n n e l a veraged c a . 23 + 2°C. Each e x p e r i m e n t a l r u n was s e t f o r 10 m i n u t e s w i t h t h e number o f b e e t l e s caught i n t h e t r a p s e r v i n g as t h e measurement o f i n t e r e s t . The a r c s i n e t r a n s f o r m e d d a t a were a n a l y z e d as a G e n e r a l L i n e a r Model (SAS 1985). A.2.3 R e s u l t s and D i s c u s s i o n A three-way i n t e r a c t i o n between f l i g h t t r e a t m e n t s , sex, and b a i t was n o t s t a t i s t i c a l l y s i g n i f i c a n t ( F 2 ^.26 = ° - 3 1 ' P > 0.05). Two-way i n t e r a c t i o n s between f l i g h t and sex ( F2,126 = ° - 0 8 ' P > 0.05), b a i t and sex ( F 1 ^ 1 2 6 = 2.63; p > 0.05), and b a i t and f l i g h t ( F 2 1 2 6 = ° * 8 8 ' P > 0-05), a l s o were n o t s i g n i f i c a n t . The mean p e r c e n t o f b e e t l e s caught i n b a i t e d t r a p s was s i g n i f i c a n t l y h i g h e r , r a n g i n g from 6.4 -15.6 %, t h a n i n u n b a i t e d t r a p s , r a n g i n g from 0.8 - 3.3 % ( F l , 1 2 6 = 8 6- 5'' P < 0.05)) ( T a b l e 1 9 ) . I n t h e b a i t e d t r e a t m e n t s , r e s p o n s e was h i g h e r f o r males t h a n f e m a l e s ( F l 12 6 = 7 - 2 1 ' P ^ 0.05). F o r t h e u n b a i t e d group, t h e r e were some c a s e s where t h e o b s e r v e d number o f b e e t l e s caught o c c u r r e d i n o n l y one o f t h e r e p l i c a t i o n s , r e s u l t i n g i n mean v a l u e s and s t a n d a r d e r r o r s t h a t a r e e q u a l t o each o t h e r . F l i g h t e x e r c i s e t r e a t m e n t s d i d n o t d i f f e r s i g n i f i c a n t l y from each o t h e r ( F 2 1 2 6 = 2.88; p > 0.05). However, a h i g h e r p e r c e n t a g e o f u n f l o w n b e e t l e s were c a p t u r e d t h a n Table 19. Response of flown and unflown Trypodendron lineatum to semiochemical-baited funnel traps i n a wind tunnel. No. of Total Total Mean % Trt. Sex Groups No. Beetles No. Beetles Caught Per 1 Group + S.E. (Reps) Released Caught Unbaited (Control) Unflown M 6 60 2 3.33 + 3.33 2 F 6 102 1 0.83 + 0.83 Flown M 6 120 1 0.83 + 0.83 F 6 75 1 1.11 + 1.11 Flown 2x M 6 115 2 1.67 + 1.67 F 6 74 2 2.44 + 1.61 Baited Unflown M 17 300 46 15.64 + 1.92 F 17 315 38 12.09 + 1.77 Flown M 17 337 41 12.03 + 2.34 F 17 343 22 6.43 + 1.18 Flown 2x M 17 341 44 12.80 + 1.76 F 17 335 27 7.98 + 1.66 Data subjected to arc sine transformation. S.E. equal to mean percentages is a result of observing caught beetles i n only one of the replications. 191 f l o w n and t w i c e f l o w n b e e t l e s . The most a p p a r e n t d i f f e r e n c e s f o r t h e b a i t e d t r e a t m e n t s , was t h e r e s p o n s e o f t h e u n f l o w n males and females (15.6 and 12.1%, r e s p e c t i v e l y ) compared t o a r e s p o n s e by f l o w n males and f e m a l e s (12.0 and 6.4%, r e s p e c t i v e l y ) (Table 1 9 ) . The h y p o t h e s i s t h a t a h i g h e r p e r c e n t a g e o f f l o w n b e e t l e s compared t o u n f l o w n b e e t l e s would r e s p o n d t o s e m i o c h e m i c a l - b a i t e d f u n n e l t r a p s was r e j e c t e d f o r b o t h s e x e s . A second h y p o t h e s i s t e s t e d was t o see i f s t o r i n g s p r i n g emerged f l o w n b e e t l e s a t c o l d t e m p e r a t u r e s (4 — 2 °C) f o r a p e r i o d o f 4 weeks r e q u i r e d t h e b e e t l e s t o go t h r o u g h a n o t h e r p e r i o d o f f l i g h t e x e r c i s e b e f o r e r e s p o n d i n g t o s e m i o c h e m i c a l s . T h i s h y p o t h e s i s was a l s o r e j e c t e d . W h i l e , t h e r e s u l t s from t h i s s t u d y c o n t r a s t w i t h t h e f l i g h t e x e r c i s e h y p o t h e s i s f i r s t p r o p osed by Graham (1959) f o r T. l i n e a t u m , and s u b s e q u e n t l y s u p p o r t e d by F r a n c i a and Graham (1967) and B e n n e t t and Borden (1971), a major weakness i n t h e s t u d y r e p o r t e d i n t h i s a p p e n d i x was n o t b e i n g a b l e t o m o n i t o r b e e t l e a c t i v i t y p r i o r t o c o l l e c t i n g them i n t h e emergence j a r s . The b e e t l e s may have had enough m e t a b o l i c a c t i v i t y t o s t i m u l a t e t h e i r r e s p o n s e t o s e m i o c h e m i c a l s . I n f a c t t h e h i g h e r c a p t u r e p e r c e n t a g e o f t h e u n f l o w n b e e t l e s s u g g e s t s t h a t t h e f l o w n b e e t l e may have been i n a weaker s t a t e due t o t h e i r l o n g e r exposure t o warmer t e m p e r a t u r e s . N i j h o l t (1967) has shown t h a t f a t c o n t e n t i n a d u l t T. l i n e a t u m d e c r e a s e s w i t h a c t i v i t y o v e r t i m e . 192 These r e s u l t s i n d i c a t e t h a t t h e amount o f m e t a b o l i c a c t i v i t y needed f o r T. l i n e a t u m r e s p o n s e t o s e m i o c h e m i c a l s s h o u l d n o t a l t e r o ur i n t e r p r e t a t i o n o f e x p e r i m e n t a l r e s u l t s where p r e v i o u s l y f l o w n b e e t l e s have been used. 193 APPENDIX 3: INFLUENCE OF DUSTING T. LINEATUM WITH FLUORESCENT POWDER ON THEIR RESPONSE TO SEMIOCHEMICALS IN A WIND TUNNEL. To s t u d y d i s p e r s a l o f Trypodendron l i n e a t u m ( O l i v i e r ) , m a r k - r e c a p t u r e e x p e r i m e n t s were d e s i g n e d u s i n g f l u o r e s c e n t powder as t h e m a r k i n g agent. T h i s t e c h n i q u e was p r e v i o u s l y u sed by Shore and McLean (1988) a t a s a w m i l l , b u t no i n f o r m a t i o n was g a t h e r e d t o see what e f f e c t , i f any, t h e mark had on b e e t l e b e h a v i o r . The o b j e c t i v e o f t h i s s t u d y was t o see i f m a r k i n g b e e t l e s w i t h f l u o r e s c e n t powder, o r f l u o r e s c e n t powder and gum a r a b i c , as su g g e s t e d by Southwood (1978), would reduce t h e res p o n s e o f T. l i n e a t u m t o se m i o c h e m i c a l s . The b e e t l e s used i n t h i s s t u d y were from t h e same group o f b e e t l e s c o l l e c t e d i n 1986 f o r t h e s t u d i e s d e s c r i b e d i n Ch a p t e r s 2 and 5. B e e t l e s were marked u s i n g t h e vacuum d u s t e r t e c h n i q u e ( L i n t o n e t a l . 1987). The t r e a t m e n t s used were pure f l u o r e s c e n t powder, a m i x t u r e o f 1/2 powder and 1/2 gum a r a b i c , and unmarked b e e t l e s . One gram o f d u s t was used t o l i g h t l y mark b e e t l e s f o r each group o f 200 b e e t l e s . F o l l o w i n g m a r k i n g , b e e t l e s were r e t u r n e d t o t h e w a l k - i n c o o l e r (4°C) where t h e y had been s t o r e d s i n c e t h e i r c a p t u r e . Response o f unmarked and marked b e e t l e s t o a s e m i o c h e m i c a l - b a i t e d ( e t h a n o l and l i n e a t i n ) f u n n e l t r a p was t e s t e d i n t h e wind t u n n e l d e s c r i b e d i n C h a p t e r 2. B e e t l e s were t e s t e d 1, 3, 5, and 7 days f o l l o w i n g m a r k i n g . Between 34 and 50 b e e t l e s were r e l e a s e d from a t r a y 2 m upwind from 194 t h e t r a p s a t an a i r f l o w o f 0.3 m/s. The b e e t l e s were g i v e n 45 min t o respond, and t h e number caught i n t h e t r a p s were s u b s e q u e n t l y t a l l i e d . A randomized complete b l o c k d e s i g n was used. Days s e r v e d as b l o c k s , and a l l t r e a t m e n t s were t e s t e d w i t h i n each b l o c k . Males and females were t e s t e d s e p a r a t e l y . The d a t a were a n a l y z e d by a n a l y s i s o f v a r i a n c e (SAS 1985). Male and female r e s p o n s e s d i d not d i f f e r s i g n i f i c a n t l y ( F l , 1 9 = ° - 8 7 ' P > 0.05), t h e r e f o r e t h e i r d a t a were p o o l e d . No d i f f e r e n c e s were found i n t h e number o f b e e t l e s caught f o r any o f t h e t r e a t m e n t s ( F 2 1 8 = 0.74; p > 0.05), a l t h o u g h s l i g h t l y more marked b e e t l e s were c a p t u r e d t h a n c o n t r o l b e e t l e s ( T a b l e 2 0 ) . Thus, i t was c o n c l u d e d t h a t l i g h t l y m a r k i n g T. l i n e a t u m w i t h f l u o r e s c e n t powder does n o t a d v e r s e l y a f f e c t t h e i r f l i g h t r e s p onse t o s e m i o c h e m i c a l -b a i t e d f u n n e l t r a p s . 195 T a b l e 20. The e f f e c t o f ma r k i n g t r e a t m e n t s on T. l i n e a t u m r e s p o n s e t o a s e m i o c h e m i c a l - b a i t e d f u n n e l t r a p i n a wind t u n n e l . 1 Treatment No. o f B e e t l e s Mean % C a p t u r e d R e l e a s e d + S.E. 2 F l u o r e s c e n t 374 12.0 + 1.7 Powder F l u o r e s c e n t Powder & Gum 371 11.5 ± 1.6 A r a b i c Unmarked B e e t l e s ( C o n t r o l ) 361 9.7 + 2.3 1 Male and female d a t a were p o o l e d t o g e t h e r . 2 . Data were s u b j e c t e d t o an a r c s i n e t r a n s f o r m a t i o n . 

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