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University Microfilms 30 0 N o rth Z e e b R o a d Ann A rb o r, M ic h ig a n 48106 A X ero x E d u c a tio n C o m p a n y 73-12,724 HAIN, Fred Paul, 1944THE BIOLOGY AND CONTROL OF THE PINE CANDLE MOTH, Exoteleia nepheos, ON SCOTCH PINE IN m i c h i g SFT: Michigan State University, Ph.D., 1972 Entomology U n iv ersity M ic ro film s , A XEROXCompany , A n n A rb o r, M ic h ig a n THE BIOLOGY AND CONTROL OF THE PINE CANDLE MOTH, E x o t e l e i a nepheos, ON SCOTCH PINE IN MICHIGAN By Fred Paul Hain A THESIS Submitted to Michigan S t a t e U n i v e r s i t y in p a r t i a l f u l f i l l m e n t o f the requirements f o r the degree o f Docto r o f Philosophy Department o f Entomology 1972 PLEASE Some NOTE: pages indistinct Filmed University as Microfilms, may have print. received. A Xerox Education Compa ny ABSTRACT THE BIOLOGY AND CONTROL OF THE PINE CANDLE MOTH, E x o t e l e i a nepheos, 1 / ON SCOTCH PINE IN MICHIGAN By Fred Paul Hain The pine candle moth.iL/ E x o t e l e i a nepheos, poses a serious t h r e a t t o p l a n t a t i o n and ornamental pines. I t mines in Scotch pine needles during the e a r l y stages o f l a r v a l development and en te rs the bud du rin g the f i r s t f l u s h o f s p r in g growth, thereby destroying the t r e e ' s a e s t h e t i c appearance and va lue as a C h r i s t ­ mas t r e e . u a l l y d ie . H e a v i ly atta c ke d tree s are s e v e re ly stunted and e v e n t ­ Adults are a c t i v e from mid-June to mid-August, o v i ­ p o s i t i n g in needle sheaths o r along tw ig s. Eggs hatch in about 10 days w ith l a r v a e o v e r w in t e r in g in the needles where high w i n t ­ e r m o r t a l i t y may occur. P a r a s itis m by Copidosoma deceptor M i l l e r ( E n c y r t i d a e ) is an important b i o l o g i c a l nepheos. f a c t o r in c o n t r o l l i n g E_. Approximately 37.63% o f the po pu la tio n in one Michigan stand was p a r a s i t i z e d and about 80% o f t h i s was by C^. deceptor. Peak emergence o f the pine candle moth w i l l days above 52° F. th r e sh o ld temp erature. occur 1060 ± 32 degree In se ctic id es , parasites, pheromones and host re s is t a n c e were i n v e s t i g a t e d f o r t h e i r p o t e n t ­ ial use in an i n t e g r a t e d control program. G u t h i o i ^ , diazinon and Fred Paul Hain methoxychlor gave e x c e l l e n t con trol but M e t a - S y s t o x - ^ had the l e a s t e f f e c t on the p a r a s i t e s o f the pine candle moth w h ile s i g n i f i c a n t l y reducing the pest p o p u l a tio n . Granular systemics a p p li e d to the s o i l were i n e f f e c t i v e in c o n t r o l l i n g the pest. A pheromone, produc­ ed by the females and a t t r a c t i n g the males, was discovered and found most e f f e c t i v e a t distances o f 3 m. or le s s . t e s t o f various chemical m a t e r i a l s , be i d e n t i f i e d . But in a screening the actual compound could not Attempts to i n v e s t i g a t e host res istanc e w i t h i n v a r - i t i e s o f Scotch pine were not successful as the a r t i f i c i a l mental experi­ techniques were not conducive to e s t a b l i s h i n g any a t t a c k pattern. —^ L e p id o p t e ra : G e le c h iid a e —^ Common name accepted by the Entomological f o r E x o t e l e i a nepheos, 7 February 1972. Society o f America ACKNOWLEDGEMENTS I am indebted to Drs. W i l l i a m E. W alln e r and Dean L. Haynes, and Mr, John H. Newman o f the Department o f Entomology, and to Dr. J. W, Wright o f the Department o f F o r e s t r y , Michigan S t a t e U n i v e r s i t y , and to Dr. Louis F. Wilson o f the USDA Forest Ser vic e f o r t h e i r advice and encouragement, and als o to the numerous Michigan Christmas t r e e growers f o r t h e i r cooperation. I would a l s o l i k e to thank Drs. James W. Butcher, Gordon E. Guyer, James W. Hanover and S. N. Stephenson f o r reviewing the manuscript. TABLE OF CONTENTS LIST OF TABLES.............................................................................. Page iv LIST OF FIGURES............................................................................. v INTRODUCTION.................................................................................. 1 LITERATURE REVIEW......................................................................... 3 B i o lo g y .................................................................................. 3 I n s e c t i c i d e s ............................... 4 A t t r a c t a n t s ......................................................................... 5 Host Re s is ta n c e ................................................................ 6 BIONOMICS OF THE PINE CANDLE MOTH..................................... 8 M a t e r i a l s and Methods...................„ ............................... 8 L i f e H i s t o r y and Damage.................................................. 10 Parasitoids. .............................................. 18 A d u lt Eclosion P e r i o d ........................................ * . . . 20 MANAGEMENT OF PINE CANDLE MOTH POPULATIONS.................. 25 . . . . . . M a t e r i a l s and Methods. .................................................. 25 Chemical Control ................................................... 28 A t t r a c t a n t s .............................................................................. 34 Host R e sis ta n ce ..................................................................... 40 DISCUSSION........................................................................................... 42 CONCLUSIONS AND RECOMMENDATIONS......................................... 45 LITERATURE CITED.............................................................................. 47 APPENDICES........................................................................................... 51 iii LIST OF TABLES Table 1. Page Head capsule width measurements o f Exoteleia nepheos....................................................................................... 2. R e l a t i v e r a t e s o f p a r a s i t i s m on E x o t e l e i a nepheos during 1970........................................................... 3. 13 In f lu e n c e o f g r a n u l a r s o i l 20 system!cs on the pine candle moth..........................................................................31 4. In f lu e n c e o f g r a n u l a r s o i l systemics on the p a r a s it e s o f the pine candle moth................................... 31 5. Control o f the pine candle moth w ith chemical s p r a y s -1 9 6 9 .................................................................................. 6. 32 Control o f the pine candle moth w ith chemical s p r a y s -1 9 7 1 ..................................................................................... 33 7. E f f e c t o f chemical sprays on p a r a s i t e s o f the pine candle moth............................................. 8. Screening o f m a t e r i a l s as a t t r a c t a n t s f o r E x o t e l e i a nepheos................................................................ 9. 33 37 S u s c e p t i b i l i t y o f 15 v a r i e t i e s o f Pinus s y l v e s t r i s to a t t a c k by E x o t e l e i a nepheos when caged on i n ­ dividual branches........................................................................41 iv LIST OF FIGURES Figure 1. Page Michigan counties w ith known i n f e s t a t i o n s o f the pine candle moth ( 1 9 7 1 ) ................................ * ...................... 2. Seasonal development o f the pine candle moth in M ic h iga n........................................................................................... 3. . ..................................................................... 15 o f E x o t e l e i a nepheos........................ 17 6. Sex d i f f e r e n t i a l 7. Threshold temperature o f l a r v a l 8. Ad ult emergence o f E x o t e l e i a nepheos from 1969, 1970, 10. 14 Head capsule leng ths o f 203 l a r v a e o f the pine candle moth. 9. 12 Head capsule widths o f 208 l a r v a e of the pine candle moth.................................................................................. 5. 11 Stages, damage and p a r a s i t i s m o f the pine candle moth.................................................................................................... 4. 9 development. . . 1971 ....................................................................................... 22 23 Pheromone traps and f i e l d cages................................................29 Distance a d u l t females, E x o t e l e i a nepheos, could a t t r a c t males from a Scotch pine s t an d ...................... v . 35 INTRODUCTION The pine candle moth, E x o t e l e i a nepheos Freeman, is a r e c e n t l y discovered pest o f Michigan p l a n t a t i o n pines. main host is Scotch p i n e , Pinus s y l v e s t r i s L . , but i t on red p in e , P_. resinosa A i t o n , A u s tr ia n p in e , and Mugho p in e , F\ mugo Tu rra (Freeman, 1966). range o f the pest i s not known but i t Its als o occurs n i g r a A r n o ld , The complete is also pre sent in Ohi^ (Neiswander, 1 9 6 6 ) , and Toronto and Ottawa, Canada ( L i n d q u i s t and T r i n n e l l , 19 67 ). There is very l i t t l e a t u r e in regards to i t s life in f o r m a t io n in the l i t e r ­ h is to r y , biology, or c o n tro l. In Michigan i t appears to be spreading throughout the southern h a l f o f the lower p e n i n s u l a , a t t a c k i n g mainly Christmas t r e e p l a n t a t ­ ions o f Scotch p in e . 10 to 25 f e e t t a l l However, since i t is also found on t r e e s ( L i n d q u i s t and T r i n n e l l , 1967), i t must be considered a po s s ib le t h r e a t to f o r e s t t r e e s as w e l l . The o v e r a l l h a b its and l i f e o b j e c t i v e of the study was to l e a r n the p e s t ' s cy cle and to use t h i s in f o r m a t io n in de veloping a management program t h a t would no t be i n j u r i o u s t o the e n v i r o n ­ ment. With the pre sent concern over environmental by p e s t i c i d e s a rhemical contamination t h a t would r e q u i r e repeated a p p l i c a t i o n s t o suppress the pest population is not d e s i r a b l e . Therefore a f u r t h e r o b j e c t i v e was t o determine what in f lu e n c e c e r t a i n chemicals 1 2 had on the e n t i r e environmental complex and how we could best op e ra te w i t h i n these c o n d itio n s to c o n t r o l the pest problem w ith a minimum o f p e s t i c i d e usage. The research on the bio lo g y and l i f e candle moth i n d i c a t e d the p o t e n t i a l ed c o n t r o l program. h i s t o r y o f the pine and n e c e s s it y o f an i n t e g r a t ­ The f o l l o w i n g f a c t s about the pest and ho s t, i n f l u e n c e d the approach o f the present i n v e s t i g a t i o n s : ( 1 ) More than 20 v a r i e t i e s o f Scotch pine have demonstrated d i f f e r e n c e s in s u s c e p t i b i l i t y to a t t a c k by several in s e c t s ( W r ig h t , e t a l . , 1966; W r ig h t , e t a l . 1967; Wright and Wils on , 1972); ( 2 ) A high percentage o f the bud i n f e s t i n g la rv ae are p a r a s i t i z e d which may serve as an impor tant b i o l o g i c a l f a c t o r in c o n t r o l l i n g the pine candle moth (Campbell and B a lders to n, 1971; L i n d q u is t and T r i n n e l l , 1967; Neiswander, 1 96 6); ( 3 ) Ad ult females produce a pheromone t h a t a t t r a c t s males, and pheromones have been used in the past as a s u r v e i l l a n c e technique to d e t e c t the presence o f a pest ( S t e i n e r , et a l . , 1 9 6 1 ). Besides the l i f e h i s t o r y s t u d i e s , then, the f o l l o w ­ ing s t u d i e s were made to le a r n t h e i r p o t e n t i a l c o n tr o l program: in an i n t e g r a t e d ( 1 ) p e s t i c i d e s and t h e i r e f f e c t on p a r a s i t i s m , ( 2 ) a t t r a c t i o n by n a t u r a l and s y n t h e t i c pheromones, and ( 3 ) host r e s i s t a n c e between v a r i e t i e s o f Scotch p in e . LITERATURE REVIEW Biology E x o t e l e i a nepheos was f i r s t descr ibed by Freeman (1967) and a companion paper by L i n d q u i s t and T r i n n e l l describes the b io lo g y o f the pest in Canada. from e a r l y J u ly to e a r l y August. (1967) b r i e f l y Ad ult f l i g h t occurs Females l a y t h e i r eggs s i n g l y or in small c l u s t e r s on needle sheaths o f the previous y e a r ' s f o l i a g e or under bark scales o f tw ig s. The la r v a e mine along the edges o f the needles and o v e r w in t e r in the needle mines. Mining term inates in mid to l a t e May, when the l a r v a e m ig ra te and feed in staminate flowe rs or e l o n g a t i n g buds. Pupation occurs about mid-June in the flowers o r shoots and the moths emerge in about 2 weeks. 5 larval Thera are instars. P a r a s i tis m plays a major r o l e in the bio lo g y o f £ . nepheos. L in d q u is t and T r i n n e l l (1967) c o l l e c t e d the f o l l o w i n g p a r a s i t e s on E^ nepheos: Achrysocharis sp. ( E u l o p h i d a e ) , Copidosoma geniculatum (Dalman) ( E n c y r t i d a e ) and Eurytoma sp. ( E u r y t o m id a e ) . a lso found a number o f p a r a s i t e s on a r e l a t e d h o s t, M a r t i n (1959) dodece1la L. Neiswander (1966) found t h a t 43% o f the la r v a e in 2 Ohio counties were p a r a s i t i z e d by C. d e c e p t o r ; and M a rt in (1959) found 50% o f the l a r v a l po pu la tio n o f £ . dodecel1 a ( p i n e bud moth) in two Ontario plantations p a ra s itiz e d . 3 4 Insecticid es Campbell (1 9 6 8 ) and Campbell and B alde rsto n ( 1 9 7 1 ) are the only res ea rc he rs who have t r i e d chemical c o n t r o l In a p r e l i m i n a r y ex pe riment Campbell on E. nepheos. ( 1 96 8) found t h a t M e ta - S y s to x - f ^ a p p a r e n t l y reduced th e pin e candle moth p o p u l a t i o n by le s s than 50% w h i l e having l i t t l e e f f e c t on the p a r a s i t e s . In an i n t e g r a t e d c o n t r o l experiment on Scotch pine in Ohio (Campbell 1971) to c o n tr o l the pine candle moth, the European pine s a w f l y , Ne o dip rio n s e r t i f e r ( G e o f f r o y ) , nus a g i l i s and B a ld e r s t o n , ( Kal t e n b a c h ) . and the sp o tt e d pin e a p h id , Eulach- M e t a - S y s t o x - ^ was a p p l i e d in e a r l y May w h i l e the l a r v a e o f E_. nepheos were boring i n the buds. This did not reduce the number o f l a r v a e i n f e s t i n g the candles but d i d reduce the number o f needle m ining l a r v a e o f the ensuing g e n e r a t i o n . Fu rther m or e, p a r a s i t i s m o f the l a r v a e was no t lessened by t h i s t r e a t ­ ment. M e t a - S y s t o x - I ^ h a s a l s o been observed to be an e f f e c t i v e control o f cone i n s e c t s on Douglas f i r (Hedlin, Zin g g , 1 9 6 7 ) , the w a ln u t aphid (H eff erm an; on ash and elm, and the elm l e a f b e e t l e 1 9 6 7 ) , v a ri o u s aphids (K o e h le r and Campbell, O ther systemics t h a t have shown a c t i v i t y (Kearby, e t a l . , 1970), 1966; Johnson and 19 68 ). in pine i n c l u d e Temi i@> Di-Systoi$^, F u r a d a ^ ^ a n d Cygoii^ Temlk has been found e f f e c t i v e a g a i n s t the Saratoga s p i t t l e b u g on red pine (Wilson and Kennedy, 1971) and i s more e f f e c t i v e than D i - S y s t o r ® i n c o n t r o l l i n g c e r t a i n aphids on Scotch p in e (Kearby and B l i s s , 19 69 ). F u r a d a i ^ has als o been used s u c c e s s f u l l y a g a i n s t the Sar atoga s p i t t l e ­ bug (Wilson and Kennedy, 1971) as w e l l as the pinyon s t u n t needle 5 midge (Brewer, 19 7 2). i s a successful con tr ol ins ec ts on Douglas f i r agent o f cone ( H e d l i n , 1966; Johnson and Zingy, 1967) and the western pine t i p moth on ponderosa pine (Haverbeke, 1971). Reynolds and M e t c a l f (1962) t h e o r i z e t h a t w a te r s o l u b i l i t y and s o il mositure play an important r o l e in t h e e f f e c t i v e n e s s o f granu­ l a t e d systemics a p p l i e d to the s o i l . Compounds o f extremely low w ate r s o l u b i l i t y may move too slowly i n t o the r oot zone to o b tain optimum i n s e c t i c i d a l v a l u e ; conversely , r e a d i l y s o lu b le compounds may lose t h e i r e f f e c t i v e n e s s more q u i c k l y by widespread movement in the s o i l and thus become l o s t to the r o o t zone. For example, more weeks l o n g e r to produce cabbage aphid m o r t a l i t i e s ob tained in a few days w it h A ttractants Chemical a t t r a c t a n t s in d e t e c t i o n and co n tr o l o f in s e c t pests o f f e r a promising means o f reducing p e s t i c i d e contamination o f the environment (Bero za, 1 9 7 1 ) . This approach takes advantage o f spec­ i a l i z e d organs which guide insects t o f o o d , a mate, or to a place of oviposition. Many o f these a t t r a c t a n t s are h i g h l y s p e c i f i c and traps can be b a it e d w i t h them f o r i n s e c t d e t e c t i o n . These traps t e l l where and when an i n s e c t is present and help reduce the need­ less spread o f i n s e c t i c i d e s i n t o areas where the pest is absent. Roelofs and Comeau (1970) discovered new sex a t t r a c t a n t s f o r 37 lepidop terous species in f i e l d screening t e s t s w ith 26 monoun­ s a tu r a t e d long-chain a c e t a t e s and a l c o h o l s . t e s t s were made p r a c t i c a l Systematic screening by the development o f l o n g - l a s t i n g p o l y - 6 e th y le n e wicks (Glass e t a l . , 1 9 7 0 ) , which r e le a s e chemicals f o r several months a t a low r a t e , and by the a v a i l a b i l i t y o f economi­ cal f o l d - u p , s t i c k y - c o a t e d t ra p s (3M C o .) . Some o f these a t t r a c t i o n s can be very stro ng. Daterman and McComb (1970) found the a t t r a c t a n t o f the European pine shoot moth, Rhyacionia bu oliana ( S c h i f f e r m a l l e r ) , to be e f f e c t i v e over a dis tanc e o f 86m where 11.5% o f the r ele as e d males where captured in tra p s b a i t e d w i t h females. However, the inc id enc e o f male sexual response depended upon the le n g th o f time males were con dit io n ed a t high i l l u m i n a t i o n (Daterman, 1972). exposed to high i l l u m i n a t i o n The lon ge r the males were (up to 48 hours) the g r e a t e r was the response t o the sex pheromone. Host Resistance Host r e s i s t a n c e to a t t a c k by various i n s e c t pests has been e x h i b i t e d in a number o f Pinus s p . . W r ig h t , e t a l . (1966 and 1967) observed f o u r p l a n t a t i o n s t h a t had 108 s e e d lo ts each o f P_. s y l v e s t r i s i n f e s t e d w ith the European pine s a w f l y , N_. s e r t i f e r . They found t h a t the percentage o f t r e e s a t ta c k e d v a r i e d from 0 to 6 f o r north Eurasian v a r i e t i e s , 6 to 11 f o r south European v a r i e t i e s , and 12 to 26 f o r c e n t r a l European v a r i e t i e s . G e n e r a l l y , the t a l l e s t v a r ­ i e t i e s were a t ta c k e d most h e a v i l y , w ith the e xc eption o f va r. u ra l- ensis which was attac ke d much less than expected f o r i t s h e i g h t . Compared w ith o t h e r v a r i e t i e s , it has w id e r needles, a lower f o l i a g e co n c e n tra tio n o f N, P, Na, Mg, Pe, and B, and the l a r v a e fe ed ing on va r. u r a l e n s i s were 1/2 to 1 i n s t a r behind in development. in an a r t i f i c i a l However, feeding experiment w ith l a t e - i n s t a r l a r v a e in the 7 l a b o r a t o r y , no p r efere n ce was shown between v a r i e t i e s . In a f i e l d a n a ly s is o f pin e root c o l l a r w e e v i l , Hylobies r a d i c i s Buch., r e s i s t a n c e between v a r i e t i e s was s t a t i s t i c a l l y s i g n i f i c a n t (W rig ht and Wilson, 1972). Bennett (1954) r ep or ted a host r e s i s t a n c e f a c t o r w ith the pine needle miner, E x o t e l e i a p i n i f o l i e l l a ies o f host t r e e s . (Cham.), a t t a c k i n g various spec­ He found a c o r r e l a t i o n between the i n t e r n a l s t r u c t u r e o f a pine needle and i t s s u s c e p t i b i l i t y t o i n j u r y . Of the many species t h i s pest a t t a c k s , lodgepole pine and ja c k pine were the most favored. The needles o f these 2 species have only 2 modera te-s ize d r e s i n canals locate d in each corner o f the needle. Scotch p i n e , on the o t h e r hand, was d e f i n i t e l y u n d e r s ir a b le as a host t r e e , with conspicuous resin canals varyin g from 6 to 16; the h ig h e r the number, the g r e a t e r the r e s i s t a n c e to the i n s e c t a p p a re n tly due to the copious resin flo w . BIONOMICS OF THE PINE CANDLE MOTH The p o t e n t i a l economic impact o f the pine candle moth is severe; reducin g the a e s t h e t i c appearance and va lue of h a r v e s t able Christmas t r e e s , and slowing the growth r a t e o f younger stock. Since the in s e c t presents a major t h r e a t to p l a n t a t i o n and ornamental pines in Michigan ( F i g . o f i t s h a b i t s and n a t u r a l c o n t r o l s , This s e c ti o n r e p o r t s on the l i f e 1 . ) , and l i t t l e is known i n v e s t i g a t i o n s were e s s e n t i a l . h i s t o r y and b io lo g y o f £ . nepheos, i n c lu d in g damage c h a r a c t e r i s t i c s and degree o f p a r a s i t i s m . M a t e r i a l s and Methods Three Scotch pine p l a n t a t i o n s i n f e s t e d w i t h the pine candle moth in lower Michigan were s e l e c t e d f o r study; one each in the western (West O l i v e ) , c e n t r a l regions o f the S t a t e . (East La nsing), and eas te rn ( M a y v i l l e ) Each s i t e con sisted o f a pp roxim ate ly 5 acres; the tre e s were g e n e r a l l y 5 - 7 f e e t t a l l w i t h a basal diameter o f 2 - 4 inches. A d u l t emergence data were c o l l e c t e d from these s i t e s in 1969 (West O l i v e and East L a n s i n g ) , 1970 (West O l i v e ) and 1971 (West O l i v e , East Lansing and M a y v i l l e ) . Adults in f l i g h t were c o l l e c t e d by use o f a r o t a r y b l a c k - l i g h t t r a p (West O liv e and East L a n s in g ), and by pheromone traps ( M a y v i l l e ) which consisted o f adu lts caged in p i n t s i z e d i c e cream carton s w i t h screening on both ends. were sprayed w ith T a n g l e - f o o ^ and placed in the f i e l d . Cartons Various numbers o f males and females were placed w i t h i n the cartons but 8 9 *AHE S UPtRIOR IMM m ■ t I Fig , 1. Michigan counties w ith known I n f e s t a t i o n s of the pine candle moth (1971). 10 a t t r a c t i o n was best where t h e r e was a preponderance o f females. The black l i g h t traps ran from 8 p.m. to 8 a.m. Climatological da ta were gathered a t each s i t e w i t h a hydro-thermograph. P a r a s it e s were c o l l e c t e d a t West O l i v e , reared in an i n s e c t a r y a t Michigan S t a t e U n i v e r s i t y and sent t o the USDA Entomology Research D i v i s i o n for id e n tific a tio n . Rearing was accomplished by c o l l e c t i n g i n f e s t e d buds in l a t e May o r e a r l y June and p la c in g the m a t e r i a l cages. The data c o l l e c t e d f o r l a r v a l in screen development came from i n f e s t e d needles and buds in the West O l i v e p l a n t a t i o n . L i f e H i s t o r y and Damage The l i f e h i s t o r y of the pine candle moth in Michigan is s i m i l a r to t h a t in O n t a r i o , Canada as r ep orted by L i n d q u i s t and T r i n n e l l (1967). There is only one ge n era tio n per y e a r ( F i g . 2). In Michigan, a d u l t a c t i v i t y occurs somewhat e a r l i e r than in O n t a r i o ; a d u lts are found from mid-June t o mid-August w i t h i n the v i c i n i t y o f the host tree. They are weak f l y e r s and d is p e rs a l ed nursery stock. The a d u lt s ( F i g . is probably through i n f e s t ­ 3A) a re a golden brown w i t h 3 g r a y is h and w h it e tra n sve rse f a s c i a e on t h e i r fore-w ing s (Freeman, 1 967). They have a mean (± S . D . ) wingspan o f 9 . 2 3 ± .47mm and a l eng th o f 5 . 5 ± .35mm (based on measurements o f 50 i n s e c t s ) . After mating the females o v i p o s i t in needle sheaths or along the twigs (Fig. 3B and 3C); the eggs are l a i d s i n g l y o r in small groups o f 2 - 4 and are a l i g h t brown, 0 . 2 6 ± .02mm wide and 0 . 3 7 ± .01mm long. The l a r v a e hatch in about 10 days and begin to excavate needle mines FEB MARCH APRIL lorvo* in N m e p*ne c a r d i e meth: A, ^ e e d l e 5 ~ : . t n ; C, egg on b a r k o f s n o o t ; shoot; r>: , . ,:nd:e ooened t o show l a r v a ; -e . $ -.-r • - j l)ry - i ;j !?, l a r v a e a iu it; e n t r a n c e h o l e or c , c a n d l e opened t o s^o1 p.: do i t l a r v a pa^as 1‘ o'- zed cy Cop-idosofna d e ce p : c ' ~ . 13 in the c u r r e n t y e a r ' s f o l i a g e ; w i n t e r is passed in the needles and e xcavation continues in the e a r l y s p r in g . About the middle o f May the l a r v a e t r a n s f e r to newly developing buds where they resume boring ( F i g . 3D and 3E). Pupation occurs w i t h i n the bud ( F i g . from e a r l y June through J u l y . 3F) The pupae are about 4 . 5 0 ± .06mm long. The pine candle moth appears to have 5 l a r v a l instars. Measure­ ments o f over 200 head capsule widths and lengths ( F i g s . 4 and 5) produced sizes s i m i l a r to those of L in d q u i s t and T r i n n e l l (1967). Table 1. shows a comparison o f the observed head capsule widths w ith the c a l c u l a t e d widths according t o the theory o f geometric progression (D y a r, 1 8 9 0 ) , and a ls o shows the f i g u r e s c a l c u l a t e d by L i n d q u i s t and T r i n n e l l ( 1 9 6 7 ) , which are s i m i l a r , except f o r some variation instar. in the f i f t h Table 1. Head capsule width measurements o f E x o t e l e i a nepheos 1arvae. Head Capsule Widths (mm. ) Instar C a lc u la t e d Observed L&T* 1 0.143 0 .1 4 3 0 .1 3 6 11 0 .1 8 7 0.187 0 .1 9 0 0 .2 44 0.246 0 .2 60 IV 0.318 0 .3 1 5 0 .355 V 0.415 0. 515 0 .4 85 m * L i n d q u i s t and T r i n n e l l (1967). NUMBER of MEASUREMENTS M Ul MEASUREMENT (m 3 M CM u CM 4 Number ■ Intiar NUMBER of MEASUREMENTS 40- MEASUREMENT (mm.) Hg. 5. Head capsule lengths of 203 larvae o f the pine candle moth. 16 The la r v a e which are a pale y e ll o w brown, o v e r w in t e r as a 3rd o r 4 t h i n s t a r and are in the 5th i n s t a r by mid-May. As development progresses the la r v a e increase in le ng th from 0 . 5 - 1 . Omm as 1st i n s t a r , to as much as 9.0mm f o r 5th i n s t a r s . The le ng th o f the needle mine also increases from a pp roxim ate ly 3.0mm to as much as 48.0mm. L in d q u is t and T r i n n e l l (1967) s t a t e t h a t in the spring the l a r v a e vacate the mine and e n t e r the same needle a t a lower p o i n t , o r an a d ja c e n t needle. T h e r e f o r e , the leng th o f the mine (Appendices la and l b ) depends upon whether the l a r v a remains in the needle o r t r a n s f e r s to another. Sex r a t i o o f the emerging a d u lts v a r i e s throughout the season but the average r a t i o during 1971 was 1 .9 2 ^ : 1.0 0 o ? e a r l y stages of emergence males predominate ( F i g . During the 6 ) , but as the season progresses the females become more abundant, e s p e c i a l l y when the number o f emerging moths is high. The browning needles caused by the mining a c t i v i t i e s o f E_. nepheos is inc onsequential damage. compared to the serio us e f f e c t s o f bud I f the i n f e s t a t i o n is se ve re , the dead candles ( e lo n g a t i n g shoots) which are f r e q u e n t l y crooked, and the r e s u l t i n g t u f t e d appearance of the f o l i a g e along w ith the d i s t o r t e d t r e e shape may make the t r e e unmarketable. may occur to the candles; Fig . 3G shows the e x t e n t o f damage t h a t the la r v a e f r e q u e n t l y bore 2 o r 3 inches i n t o the shoot, but o t h e r times only the shoot t i p is brown. Even mature Christmas t r e e stands w ith l i g h t i n f e s t a t i o n s pr e se nt a hazard as a brood source. For example, in one l i g h t l y a t ta c k e d stand, t h e r e was no dramatic a l t e r a t i o n in the s a le value o f the i n d i v i d u a l D IF F E M N T IA L SEX | :PEAK in EMERGENCE DAYS o ftff tU START of EMERGENCE (1971) Fig. 6 Sex differential of Exotelela nepheos. The horizontal line narked 0 indicates no difference in the number o f emerging adults (a 1:1 sex ra tio ,). 18 trees. However, a d ja c e n t t o t h i s stand was a 3 y e ar old p l a n t i n g which became i n f e s t e d . In a sample o f 25 young t r e e s , 25.5% o f the candles were attacked w ith a range o f 0 t o 80%. T h e r e f o r e , due t o an obvious reduction in growth r a t e and t r e e form, a s l i g h t nuisance in a mature stand becomes a serious economic t h r e a t in a younger one. Parasi toids The f o l l o w i n g p a r a s i t e s , reared a t Michigan State U n i v e r s i t y from m a t e r i a l c o l l e c t e d a t West O l i v e , M ichigan, were i d e n t i f i e d by the USDA Entomology Research D i v i s i o n , E x e r is te s comstockii ( C r . ) ( Ichneumonidae) Scambus hispae ( H a r r . ) ( Ichneumonidae) I t o p l e c t i s c o n q u i s i t o r (Say) Campolex sp. B e l t s v i l l e , M ary la nd .—^ (Ichneumonidae) (Ichneumonidae) Chelonus r e c u r v a r i a e McComb (Braconidae) Bracon g e le c h ia e Ashm. Phanerotoma sp. (Braconidae) (Braconidae) Apanteles p a r a le c h ia e Mues. (Braconidae) Copidosoma deceptor M i l l e r ( E n c y r t i d a e ) Elasmus s e t o s o s c u t e l l a t u s Crawford (Eulophidae) Sympiesis s t ig m a tip e n n is G i r a u l t (Eulophidae) Eupelmella v e s i c u l a r i s (R etz.) (Eupelmidae) 3/ Ichneumonidae i d e n t i f i e d by R. W. C a r ls o n , Braconidae by P. M. Marsh, Syrphidae by L. V. Knutson, and E n c r y r t i d a e , Eulophidae, Eupelmidae, Pteromalidae and Eurytomidae by B. D. Burks. 19 Habrocytus phyci di s Ashm. Eurytoma p i n i (Pterom alidae) Bugbee ( E u ryto m id a e) A l l o g r a p t a o b l i q u a (Say) ( S y r p h id a e ) L i n d q u i s t and T r i n n e l l ( 1 9 6 7 ) a ls o c o l l e c t e d a species o f A c hry soc haris ( E u l o p h i d a e ) p a r a s i t i z i n g ( 1959 ) E_. nepheos and M a r t i n found Pim plopterus parvus ( C r e s s . ) (Ichneumonidae) on E_. d o d e c e l1 a L. C_. de c ep to r is by f a r the most e f f i c i e n t o f the above parasites. F ig s . 3H ..nd 31 show l a r v a e o f E_. nepheos p a r a s i t i z e d by C_. de c ep to r (pu pa e ). T a b le 2 shows t h e r e l a t i v e abundance o f t h i s p a r a s i t e as compared t o the o t h e r s . appears to be i n c r e a s i n g i n Michigan. p o p u l a t i o n s was p a r a s i t i z e d 1971 t h i s The amount o f p a r a s i t i s m In 1969, 19.18% o f the l a r v a l (1152 i n f e s t e d buds examined) and by in cre as ed t o 37.63% (782 i n f e s t e d buds examined). In Ohio and O n t a r i o , where the pine candle moth i s o f c o n s i d e r a b l y les s concern, degree o f p a r a s i t i s m has been h i g h e r . Neiswander ( 1 9 6 6 ) r e p o r t s t h a t a p p r o x i m a t e l y 43% o f the l a r v a e in two Ohio cou ntie s were p a r a s i t i z e d and M a r t i n of ed. (1959) found 50% o f the l a r v a l population do dec e l1 a ( p i n e bud moth) in two O n t a r i o p l a n t a t i o n s p a r a s i t i z ­ Degree o f p a r a s i t i s m reaching the se f i g u r e s have o n ly been o b t ai n e d r e c e n t l y in Michigan i n d i c a t i n g t h a t an in t r o d u c e d nepheos p o p u l a t i o n was ab le to spread t h r o u g h to u t lower Mich igan w h i l e the p o p u la tio n s o f b i o l o g i c a l control agents lagged behind. 20 Table 2. R e l a t i v e r a t e s o f p a r a s i t i s m on E x o t e l e i a nepheos during 1970*. Parasite Family P a r a s i t i z e d Larvae Number Percent o f Total 1083 7 9.5 0 152 1 1 .3 8 Ichneumoni dae 62 4.64 Eurytomidae 12 0.90 Eulophidae 9 0.67 Pteromalidae 8 0.60 Syrphi dae 5 0 .3 7 Eupelmidae 4 0.30 E n cyrtidae (Copidosoma deceptor) Braconi dae * Over 3000 i n f e s t e d Pinus s y l v e s t r i s buds were c o l l e c t e d a t West O l i v e , Michigan. O verw in tering m o r t a l i t y may also be a strong l i m i t i n g factor. In 3 po pu la tio n samples ( r a n g i n g from 13 to 69 l a r v a e ) taken in January and March, from West O l i v e , Michigan (Appendix I I ) , percentage o f dead l a r v a e ranged from 46 to 100%. the This may e x p l a in why the pine candle moth has not been re p o r t e d in the n o r t h e r n p a r t o f the lower peninsula o f Michigan; being unable to s u r v i v e the harsh w i n t e r s o f these no rth e r n reg ions. A d u lt Eclosion Period The peak o f a d u l t e c l o s i o n is im p orta n t f o r tim ing chemical 21 c o n tr o l programs. Temperature (Appendix I I I ) alone compared to a d u l t t r a p c o l l e c t i o n s was too v a r i a b l e - ranging between 54° and 8 0 ° F - to be u s e f u l . I n s t e a d , p r e d i c t i n g peak o f ec lo s io n was attempted by the degree-day method. To do t h i s i t is necessary to determine the most r e l i a b l e thresh old temperature above which degree days are accumulated. This is t y p i c a l l y done by p l o t t i n g perc ent development over d i f f e r e n t temperatures and e x t r a p o l a t i n g to the p o i n t where the regression l i n e crosses the a b s c is s a , but Casagrande (1971) p o in ts out t h a t t h i s method is o b j e c t i o n a b l e because i t is hard to j u s t i f y extending a regression l i n e beyond data p o i n t s . The method used in stead is s i m i l a r to t h a t developed by Casagrande ( 1 9 7 1 ) . Degree days were determined f o r various th r e s h o ld temperatures ranging from 42° to 76° F (Appendices IVa IV f). Accumulated degree days were c a l c u l a t e d from the d a i l y maximum and minimum temperatures by using the sine curve method d e sc ribe d by B a s k e r v i l l e and Emin ( 1 9 6 9 ) . N ex t, the degree days a t which peak emergence occurs was recorded f o r a l l standard e r r o r determined (Appendix V). s i t e s and the This was transformed i n t o the c o e f f i c i e n t o f v a r i a b i l i t y and p l o t t e d over th r e s h o ld tempera­ tures . Development appears to s t a r t around the thr esh old temperature of 52°F ( F i g . 7). T h e r e f o r e by t h i s method peak emergence can be expected to occur a t about 1 0 6 0 ± 3 2 ( ± S . E . ) degree days above 52°F. t h r e s h o l d temperature. rough e s t i m a t e . 1971 ( F i g . In p r a c t i c e , however, t h i s gives only a The emergence p a t t e r n a t East Lansing f o r 1969, 1970, 8) shows a good p r e d i c t i o n f o r 1969 but t h e r e was a delayed 17 16 15 14 13 » 12 K i n i* 0 9 bm I «• 2Ik 7 3 * 5 41 3 2 1- ■i 'T f ■! ~r i- 42 44 46 48 50 52 i I i 1 I i I l 5 6 5 8 6 0 6 2 6 4 6 6 6 0 7 0 7 2 THRESHOLD TEMPERATURE <°F) Fig. 7. Threshold tenperature of larval development. appears to begin between 50 and 54°F. Development i ■ i 74 i ' 76 i 78 u i 321969 1970 1971 2624- o NUMBER EMERGIN 22 - 18- 12 10 - DECREE DAYS ( * 100) Fig, 8. Adult emergence of Exoteleia nepheos for 1969, 1970, 1971, 24 p r e d i c t i o n f o r 1970 when 930 degree days would have given the peak. There were t h r e e unusually d i s t i n c t peaks in 1971 which cannot be e xplain ed on the basis o f degree days alone. I f degree days was the only f a c t o r c o n t r o l l i n g emergence then one would expect a s i n g l e peak curve as e x h i b i t e d during the previous two y e a rs . MANAGEMENT OF PINE CANDLE MOTH POPULATIONS Research to develop i n t e g r a t e d c on tro l n e c e s s it a t e s a broad approach to i n s e c t pest problems. given to a l l Consideration must be p o t e n t i a l methods o f po pu la tio n red uction i n c l u d in g b i o l o g i c a l , chemical and c u l t u r a l and p l a n t r e s i s t a n c e . c o n t r o l s , pheromones, a t t r a c t a n t s This in v o lv e s comprehensive in f o r m a t io n on the bio lo g y and ecology o f the pest species on the host p l a n t . U ltim ately a ll o f these techniques must be o r c h e s t r a t e d to produce economical con tr ol w it h o u t damage to the environment. M a t e r ia ls and Methods CHEMICAL CONTROL.-All chemical con trol experiments were done in i n f e s t e d p l a n t a t i o n s near West O l i v e , Michigan; t r e e s ranged in h e i g h t from 1 .5 to 2 m. and had a basal dia m ete r between 5 and 10 cm. The g r a n u l a r systemics inc lu de d Temi , and Dasani Furada During e a r l y May, p r i o r to movement o f the l a r v a e from the needles to the buds, granules were a p p l ie d to the s o i l around the p e r ip h e r y of the t r e e crown. The randomized block design had 5 r e p l i c a t e d trees f o r each dosage r a t e o f 2 2 . 8 , 4 5 . 6 , 9 1 .2 and 1 14.0 grams o f systemic per 1 . 0 cm. of t r e e dia m ete r ( 2 , 4 , 8 , and 10 o z / i n . to a llow f o r s o i l of d ia .). penetration. S u f f i c i e n t r a i n f ollow ed a p p l i c a t i o n A n alysis o f the treatments was done by c o l l e c t i n g the i n f e s t e d buds and examining them f o r both pest and p a r a s i t e s . 25 26 In 1969 the f o l l o w i n g sprays were analyzed f o r c o n tr o l of nepheos l a r v a e w it h o u t regard to p a r a s i t i s m : 1) DDT - 1 gal o f 25% emulsion/100 gal H20; 2) dia zin on - 1 pt o f 48% emulsion/100 gal H2 O; 3) S e v i $ ^ - 4 lbs o f 50% w e t t a b l e powder/100 gal methoxychlor - 1 gal o f a 2 l b / g a l 3 pts o f a 30.5% emulsion/100 gal a 22% e m u ls io n /1 00 gal H20. replications; ^0; 4 ) emu lsion/100 gal H20; 5) C y g o i ® H2 O; and 6) Guthio The randomized block design had 6 each block consisted o f 400 t r e e s so t h a t each t r ea tm en t covered app roximately 2400 t r e e s . A p p l i c a t i o n was done w ith a h y d r a u l i c spr a ye r (300 lbs o f pr essur e) u n t i l o f f and i t 1 qt of ru n­ took 80-100 ga ls o f i n s e c t i c i d e to cover the 2400 t r e e s . The program was i n i t i a t e d app roxim ate ly 10 days a f t e r peak a d u l t emergence, about the end o f J u l y , as determined by counts from a b la c k lig h t trap. Results were analyzed by randomly s e l e c t i n g a s i n g l e t r e e from each block (6 tree s f o r each t r e a t m e n t ) and cou nt­ ing the In number o f dead buds and crooked candles. 1971 t h i s experiment was repeated w i th a con s id e ra ti o n o f e f f e c t s on p a r a s i t i s m as w e l l . e xperim en t, and M e ta -S ys to xwere as f o l l o w s : DDT and C y g o i^ w e r e dropped from the and L a n n a t ^ ^ w e r e added. 1) M e t a - S y s t o x - 1 ^ - The r a t e s 1.5 pts o r a 25% emulsion/100 gal H^O; 2) G u t h i o i ^ - 2 qts of a 22% emulsion/100 gal H2G; 3) d i a z i n o n same as in 1969; 4) S e v i r ^ - 1.5 lbs o f 80% w e t t a b l e powder/100 gal H20; 5) L a n n a t ^ ^ - 0 . 5 lbs o f 90% w e t t a b l e powder/100 gal H20. sprays were a p p l ie d by a h y d r a u l i c spr ay e r u n t i l The r u n o f f ; a pp roxim ate ly 8 ga ls o f spray were used to cover 5 t r e e s in a randomized block 27 design. A p p l i c a t i o n again was done a pp roxim ate ly 10 days a f t e r peak emergence (end o f J u l y ) to co n tr o l the ha tching la r v a e and measurements o f the te r m in a l growth o f the 35 sprayed and c o n tr o l t r e e s found the average t o be app roximately 14 .0 0 ± 3 .5 4 inches (+ S.D. ) a t t h i s t im e. The experiment was analyzed by c o l l e c t i n g i n f e s t e d buds and examining f o r h e a lt h y and p a r a s i t i z e d 1arvae. ATTRACTANTS.— Experimental a t t r a c t i o n work was done in an i n f e s t e d Scotch pine p l a n t a t i o n near M a y v i l l e , Michigan. Five to ten l i v e a d u l t moths, r eare d in an i n s e c t a r y a t Michigan S t a t e U n i v e r s i t y , were placed in an ic e cream ca rt o n t r a p ( F i g . placed various dis ta nc es from the i n f e s t e d stand to determine d i s t a n c e over which the a t t r a c t i o n is e f f e c t i v e . placed w i t h i n the traps was not determined u n t i l 9A) and Sex o f the ad u lts a f t e r the f i e l d experiment was complete. A f i e l d screening t e s t o f 51 d i f f e r e n t chemical m a t e r i a l s —^ (T a b le 8) f o r a t t r a c t i o n p r o p e r t i e s was also performed by pla cin g 10 u l i t e r s o f each t e s t chemical e t a l . 1970) i n t o a p o ly e th y l e n e cap (Glass ( S c i e n t i f i c Products, 05-6 n a t u r a l c lo s u re f o r 6 0 9 7 5 - L , v i a l s , 2-dram ). t ra p s (Fig. 9B) and hung randomly (Roe lofs and Comeau, 1 9 7 0 ) . p o ly e th yle ne The caps were placed in on Scotch pine in the in f e s t e d area They remained in the f i e l d through the month o f J u ly and trapped i n s e c t s were p e r i o d i c a l l y c o l l e c t e d . 4/ — M a t e r i a l s obtain ed from M a r t i n Jacobson, P e s t i c i d e Chemicals Research Branch, Entomology Research D i v i s i o n , USDA, B e l t s v i l l e , Maryl and. 28 HOST RESISTANCE.-Two methods were attempted in i n v e s t i g a t i n g po s s ib le host r e s i s t a n c e in the many v a r i e t i e s of Scotch pine: cage experiments on mature t r e e s ( F i g . experiments ( F i g . 9D). Host m a t e r i a l sleeve 9C) and caged se ed lin g f o r the sle eve cage experiments came from a Scotch pine geographic o r i g i n study near East Lansing, Michigan ( W r i g h t , e t a l . 1966). F i f t e e n d i f f e r e n t v a r i e t i e s o f 10 y ea r o l d Scotch pine were t e s t e d w i t h 4 r e p l i c a t i o n s , by caging one branch w ith a 20 x 20 mesh screen cage and r e l e a s i n g 10 a d u l t moths i n t o the cage. The f o l l o w i n g f i e l d season the r e s u l t s were analyzed by removing the caged branches and counting the number o f mined needles and buds. Red pine seedlin gs and eleven seed sources o f Scotch p in e , (2 years o l d ) were ob tained from Van's Pines in West O l i v e , Michigan and placed in s i x (12' x 1 9 .5 ' x 6 ') cages. Each cage had 5 seed­ l in g s o f each v a r i e t y randomly dispersed and 70 a d u l t moths were r e ­ leased in each. The r e s u l t s were analyzed the f o l l o w i n g f i e l d season by i n s p e c t in g the needles and buds. Chemical Control M a t e r i a l s were t e s t e d not only f o r t h e i r a b i l i t y to control the pine candle moth but als o f o r t h e i r e f f e c t on the l a r g e number o f p a r a s i t e s t h a t a t t a c k £. nepheos. In one Michigan p l a n t a t i o n 37.63% o f 782 i n f e s t e d buds examined had l a r v a e t h a t were p a r a s i t i z e d ; most o f t h i s p a r a s i t i s m was by Copidosoma deceptor ( E n c y r t i d a e ; Lepidoptera). T h e r e f o r e , the id e a l i n s e c t i c i d e is one t h a t reduces ro ID 9. A.’1 pr sro^ore tran baibed '■"! tn ?jAe cardie piotris -PC' s l e r e cage in w h i r ,'iults -.ere released p., pneroTOre C^ap ir re^mater'aU l " ' x l q.5'x6' rage ccr : ‘-; " '- ‘ p O'1 ■■.•£!■, r ’ s r c 30 the pine candle moth p o p u l a ti o n to where the n a t u r a l enemies o f the pest can keep i t below an economic t h r e s h o ld . In a p r e l i m i n a r y experiment w ith the g r a n u la r systemics i t was de sired to determine the proper r a t e of a p p l i c a t i o n . The granules were a p p li e d in a band between t r e e rows a t a r a t e o f 1 8 .7 grams and 4 6 .5 gram/m ( 0 . 2 and 0 . 5 o z / f t . ) . as c o n t r o l s . Rows between treatments were used None o f the systemics showed any a c t i v i t y lin g the larvae. in c o n t r o l ­ There was no s i g n i f i c a n t d i f f e r e n c e in the number o f dead buds between t r e a t e d and u n tre a te d rows. The f o l l o w i n g y e a r the r a t e s o f a p p l i c a t i o n were increased and the s i z e o f the t r e e s was taken i n t o c o n s i d e r a t i o n . were a p p l i e d to the s o i l The granules around the p eriph e ry o f the t r e e crown a t 2 2 . 8 , 4 5 . 6 , 91.2 and 114.0 grams/cm o f t r e e diam e te r. and i n f l u e n c e on p a r a s i t e s was i n v e s t i g a t e d . the average percent m o r t a l i t y Both con trol Although in most cases (Table 3) and the average r a t e o f p a r a s i t i s m (Table 4) was h i g h e r than the c o n t r o l s , t h i s 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 and i t appears t h a t these g r a n u l a r systemics are not a c t i v e enough in Scotch p i n e , even a t high dosages, to serve as adequate c o n tr o ls o f the pine candle moth. No atte m pt was made to enhance the uptake o f the i n s e c t i c i d e i n t o the t r e e by w a te r in g or r ak in g the granules i n t o the s o i l , but r a i n was f re q u e n t in the area. 31 Table 3. I n f lu e n c e o f g r a n u l a r s o i l systemics on the pine candle moth. Percent M o r t a l i t y Systemic/ Tree d i a . grams/cm Furadar^ 70.83 6 0.53 4 6.23 64.05 2 2 .8 4 5 .6 9 1 .2 114.0 Table 4. Dosage Sys temi c / Tree d i a . grams/cm 22.8 45.6 91.2 114.0 Di-Systo r( ® Temi 6 6.2 0 5 9.3 0 6 5 .7 7 6 5.3 3 60.50 6 7.2 7 5 6.3 8 60.33 Influ en ce o f g r a n u l a r s o i l pine candle moth. Dasani 1 ® Cygorf&) Control 60.23 57.45 6 5.4 0 6 1.0 8 5 0.0 0 6 0.6 7 55.55 59.39 5 8.3 8 54.92 51.36 46.08 systemics on the p a r a s i t e s o f the Percent M o r t a l i t y Furadai$^ Temilf^ 50.59 4 5.7 3 37.68 44.44 5 3.79 40.69 36.91 34.66 D i-S ysto r^ gave adequate control 36.30 42.71 31,11 3 4.5 8 4 3 ,69 4 6 .36 4 2.5 5 34.71 Of the f o l i a r sprays t e s t e d Guthio Das an i Cygo^ Control 48.53 4 2.5 4 4 3.2 4 3 7.29 4 4 .7 2 3 3 .6 7 39.09 33.04 diazinon and methoxychlor in 1969 (T a ble 5) w h il e DDT and Cygor$^ showed no s i g n i f i c a n t d i f f e r e n c e from the un tre a te d c o n t r o l s . dia zin on and methoxychlor gave complete control In 1971 G u t h i o $ ^ (Table 6) w h i l e Sevi and M eta-Systox- £ > s i g n i f i c a n t l y reduced the number o f i n f e s t e d buds. 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 the u n t re a t e d t r e e s . and Sevi Lannati^ Meta-Systox- had the l e a s t e f f e c t on the p a r a s it e s o f the pine candle moth 32 (Table 7) although these e f f e c t s were 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 according to Tukey’ s m u l t i p l e comparison t e s t . Table 5. Control o f the pine candle moth with chemical sprays - 1969 Dead bu d s /t re e T reatment Average Number Range Guthiorf^ 2.83c* 0 -7 Di azinon 2 .8 3 c 0-8 Methoxychl or 2.92c 0 -9 Sevif(E) 3.67bc 0 -9 Cygofr^ 7 . 75abc 0-36 DDT 1 1 . 1 7ab 0-28 Control 1 1.67a 0-34 * Treatments which do not have the same l e t t e r a re s i g n i f i c a n t l y d i f f e r e n t according to Tukey's m u l t i p l e comparison t e s t a t the 5 % l e v e l . 33 Table 6. Control o f the pine candle moth w ith chemical sprays - 1971, I n f e s t e d b u d s /t r e e T reatment Average Number Range G u th io ^ 0 b* 0 Di azinon 0 b 0 Methoxychlor 0 b 0 Meta-Systox-^^ 1 .5 0 b 1-3 Sevi 4 .0 0 b 3-6 Lannat^^ 6.75 a 0-17 1 6.3 0 a 9 -29 Control * Treatments which do not have the same l e t t e r are s i g n i f i c a n t l y d i f f e r e n t a c c o r d in g ly to Tukey's m u l t i p l e comparison t e s t a t the 5% l e v e l . Table 7. E f f e c t o f chemical sprays on parasi tes o f the pine candle Treatment P a r a s i t i zed Larvae Percentage Range Guthi o ! ^ 0 0 Di azinon 0 0 Methoxychlor 0 0 Lannat^^ 4.18 0-16.7 S e v ir^ 9.83 0-25.0 M eta-Systox-f® 20.83 0-50.0 Control 2 2 .7 0 15. 7 9 - 2 8 . 0 34 A tt ractants During the 1970 f i e l d season a p r e l i m i n a r y t e s t was performed by pla cin g 10 a d u l t moths w i th Scotch pine f o l i a g e in each of 2 traps s i m i l a r to those in f i g . were s e t ; empty. 9A. Three o t h e r c o n tr o l traps 2 w i t h Scotch pine f o l i a g e and the o t h e r completely A l l were sprayed w i t h T a n g le f o o ^ ^ a n d placed around the edge o f an i n f e s t e d p l a n t a t i o n . F iv e days l a t e r they were examined and the b a i t e d traps had a t t r a c t e d 26 a d d i t i o n a l moths from the f i e l d p o p u la tio n w hile the c o n tr o l traps had no c a p t u r e s . This was the f i r s t demonstration o f an a t t r a c t i o n a ss ocia ted w ith the pine candle moth. During the next f i e l d season a d d i t i o n a l performed in M a y v i l i e and East Lansing. o f the a t t r a c t i o n , traps ( F i g . fie ld t e s t s were To demonstrate the power 9A) were placed a t vario us distances ( 1 , 3, 7 and 10m) from an i n f e s t e d stand in M a y v i l l e . are shown in F i g . 10. The r e s u l t s A strong a t t r a c t i o n occurred a t the s h o r t e r distances and, from w ith in 3m., was s i g n i f i c a n t l y h ig h e r than a t t r a c t i o n a t g r e a t e r d is t a n c e s . Again the co n tr o l traps had no c aptures . The sex o f the adults placed w i t h i n the traps v a r i e d from predominately male to predo minately female but on ly the l a t t e r produced an a t t r a c t i o n . When t r a p s were male or had a n e a r ly even m ixture o f males and females, very few adu lts were captured. On the o t h e r hand, the sex r a t i o o f the ad u lt s a t t r a c t e d to the traps 12 11 a 10 9 8 7 w U1 6 5 4 3 2 I T" 3 -r 4 T 6 -r 7 nr 8 T T“ 9 10 Dittanc* from Pin* Stand (mcttn) ig. 10. distance adult females, Exoteleia repneos, cou’ d attract males from a Scotch pine stand, instances *hicr. do ret have the same le tte r are s:grificar.tly different according to Trey's lultiple comparison test at the SI le.el. 36 was app ro x im ately 9:1 in f a v o r o f males ( t h i s is only an a p p r o x i ­ mation s in c e the c on dition o f many captured a d u l t s made sex d e t e r m i n a t io n d i f f i c u l t ) . A r e l e a s e experiment was performed in an open f i e l d in East Lansing; 8 t r a p s c o n ta in in g a preponderance o f females, were staggered in 2 c o n c e n tric c i r c l e s o f 1 and 2m r a d i i emerged a d u l t s were re lea se d from the c e n t e r . and newly No released adu lts were recaptured a t the pheromone t r a p s i n d i c a t i n g the newly emerged moths were not immediately responsive to the a t t r a c t ­ ion o f females. In a f i e l d ( M a y v i l l e ) screening t e s t , 51 d i f f e r e n t chemicals were t e s t e d as possible pheromones o f the pin e candle moth ( F i g . 9 8 ). The m a t e r i a l s are l i s t e d in Table 8 along w ith the number o f a d u l t E, nepheos ca ptured. Results show no s i g n i f i c a n t d i f f e r ­ ences between chemicals and, e v i d e n t l y , the pheromone o f the pine candle moth was not one o f the m a t e r i a l s , o r , a t l e a s t , was not present in a co n c en tra tio n c o m p e t i tiv e w i t h the n a t u r a l In an open f i e l d experiment (E as t Lansing) 9 m a t e r i a l s attraction . ( E n t . No. 1996, 17442, 34392, 3299, 21161, 21534, 2 4 7 9 5 , 522, and 33993; only these nine were used because o f l i m i t e d q u a n t i t i e s o f the m a t e r i a l ) were placed randomly in a s e m i - c i r c l e with a 2m radius and pine candle moths were r elea s ed from the ce n te r (upwind). The r e s u l t s were also ne gativ e (no adu lts c a p t u r e d ) . One i n t e r e s t i n g s i d e l i g h t o f the screening t e s t done a t M a y v i l l e 37 Table 8. Screening o f M a t e r i a l s as A t t r a c t a n t s f o r E x o t e l e i a nepheos Number 522 5859 941 1025 1955 1996 11016 11586 11595 17442 17540 18152 18258 18262 18336 207 21161 21209 21282 21527 21537 21902 21534 24138 24795 3294 33824 3429 3878 30099 30725 31164 31166 31183 31186 31227 31833 33265 32266-b 33478-h 33538 33845 33971 33993 33994 34180 Terpineol, acetate Cyclohexanol , 2 - p h e n y l - , a c e ta t e L in a lo o l, acetate Ethanol, 2 - e t h o x y - , a c e t a t e A c e t ic a c i d , benzyl e s t e r Furfuryl a lc o h o l, acetate 1- T e t r a d e c a n o l , a c e t a t e 1-Dodecanol, a c e t a t e Cyclohexaneethanol , aceta te Fenchyl a l c o h o l , a c e t a t e Benzyl a l c o h o l , a l p h a - m e t h y l - , a c e t a t e Cyclohexanol, 4 - i s o p r o p y l - , a c e t a t e Cyclohexanol, 2 - sec- b u t y l - , a c e t a t e Cyclohexanol, 2 - c y c l o h e x y l - , a c e t a t e G eraniol, acetate Benzyl a l c o h o l , j> -ethox y-, a c e t a t e F u rf u ry l a l c o h o l , t e t r a h y d r o - , a c e t a t e Benzyl alcohol, 3, 4-d im e th y l-, acetate Benzyl a l c o h o l , p - e t h y l - , a c e ta t e Benzyl a l c o h o l , O - m e t h y l - , a c e t a t e Benzyl a l c o h o l , m , e t h y l - , a c e t a t e Benzyl alcohol, F, 5-d im eth yl-, acetate Ace tic a c i d , undecyl e s t e r 2 - B u t a n o l , 3-methy1- 4 - p h e n y l - , a c e t a t e A c e t ic a c i d , cyclohexyl e s t e r t r a n s - 5 - h e x a d e c e n - l - o l , a c e ta t e Carvacrol, acetate A c e t ic a c i d , phenethyl e s t e r 3 - C y c lo h e x e n e - l- m e t h a n o l, a lph a , a l p h a - 6 t r im e t h y l- , acetate A c e tic a c i d , esoo ctyl e s t e r 1, 3 - D i o x o l a n e -4 -m e th a n o l, 2 - p h e n y l - , a c e ta t e 1 , 3 - D i o x o l a n e -4 -m e th a n o l, 2 - ( 6 - m e t h y l - 3 c y c l o h e x e n - 1 - y l ) , a c e ta t e 1 , 3 - D i o x o l a ne -4 -m e th a no l, 2 - b e n z y l - , a c e ta t e 1, 3 - D i o x o l a ne-4 -m eth a no l, 2 - p - t o l y l - , a c e ta t e 1 , 3 - D i o x o l a ne -4 -m e th ano l, 2 - e t h y l - 2 - m e t h y l - , a c e ta t e 2-Butanone, 4-(]D-hydroxyphenyl ) - , a c e t a t e 7 -D o d e c y n -l-o l, acetate c i s - 7 - D o d e c e n - l- o l , a c e ta t e c i s - 7 - H e x a d e c e n - l- o l , ace ta te 1 0 -Isopropyl-1 1 -m e th y l-c is -5 , 9-Dodecadien-lo l , acetate 1 0 - I s o p r o p y l - c i s - 5 , 9 - T r i d e c a d i e n - l - o l , a c e ta t e c i s - 9 - D o d e c e n - l - o l , ac eta te c i s - 8 - D o d e c e n - l - o l , aceta te t r a n s - 8 - D o d e c e n - l - o l , ace ta te c i s - 4 - H e x a d e c e n - l - o l , ac eta te 15 19 1 7 10 10 2 12 6 16 6 6 10 1 2 8 9 3 8 3 4 3 18 6 16 13 8 12 9 6 8 3 5 8 4 0 7 3 1 3 11 3 4 11 5 8 38 Table 8 Continued 34370 34392 34395 34412 34462 8 - N o n e n - l - o l , a c e ta t e c i s - 3 - h e x e n - l - o l , a c e ta t e 2 - N o n e n - l - o l , a c e ta t e 2 , 4 - U n d e c a d i e n - l - o l , a c e ta t e 7-B u to x y -l-h e p ta n o l, acetate 5 14 8 8 9 * Ent. No. appear in " M a t e r ia l s t e s t e d as i n s e c t a t t r a c t a n t s " , A g r i c u l ­ t u r e Handbook No. 239, Agr. Res. Ser. USDA. 39 5/ was t h a t 160 i n d i v i d u a l s o f E l a t o b i a r o a r t i n e l l a — Walker ( T i n e i n a e ) were captured in the 3 t r a p s c o n ta in in g c i s - 9 Dodecen-1-01 a c e t a t e . This a t t r a c t i o n was very stro ng as these moths were observed f l u t t e r i n g around the f i e l d lab as the m a t e r i a l s were being placed in the t r a p s . As f a r as can be determined t h i s is the f i r s t record o f t h i s i n s e c t appearing in Michigan. According to Forbes (1923) the la r v a e bore in the bark o f Pinus spp. during e a r l y sp rin g and the a d u l t s appear in June. 5/ — But, a p p a re n tly they are o f no economic importance. I d e n t i f i e d by J. H. Newman, Michigan S tate U n i v e r s i t y . 40 Host Resistance The sleeve cage experiments on mature t r e e s gave h ig h ly variable resu lts (Table 9) t h a t showed no s i g n i f i c a n t d i f f e r e n c e s between v a r i e t i e s a t the .05 l e v e l o f s i g n i f i c a n c e . The experiment in which a d u l t pine candle moths were r e le a s e d in cages c on tain ing various v a r i e t i e s o f Scotch pine was not succ e ss ful. o v i p o s i t i o n o r in s e c t damage was found. No evidence o f Under these h i g h ly a r t i f i c i a l c on dit io n s any number o f f a c t o r s could have i n f lu e n c e d the a b i l i t y o f the a d u l t s to mate and o v i p o s i t e and the r e s u l t s do not n e c e s s a r i l y i n d i c a t e t h a t t here is no host r e s is t a n c e . 41 Table 9. S u s c e p t i b i l i t y o f 15 v a r i e t i e s o f Pinus s y l v e s t r i s to a t t a c k by E x o t e le ia nepheos when caged on i n d i v i d u a l branches. Variety Seed Source Average Dead Needles Number Dead Suds a l t a i ca Southern S i b e r i a 21.50+33,75* 0 . 5 0 ± 1 .00 s e pe n tr ion al i s So uth-ce ntra l Sweden and Norway, Southern Fin la nd 2 0 . 25 + 23 .4 7 3.00+4.08 rigens is La tv ia n SSR, S. Sweden 1 9 . 5 0 ± 2 4 .24 1 . 50±1.91 u r a le n s is Ural Mountains 2 5 . 7 5 ± 3 0 . 73 2 . 0 0 ± 2 . 83 polonica Poland 2 0 . 7 5 ± 2 7 . 89 2.00+2.83 borussica Lowlands o f Northeastern Germany 5 0 .3 3+ 4 2.8 3 0 . 6 7 ± 0 . 58 hercynica Germany, Czechoslovakia 25 .60+37.29 2.00+4.47 haguenensis Belgium, Vosges Mountains 6 5 . 2 5 ± 9 5 .8 2 o f France, a d ja c e n t Germany 2.00+2.45 ' East A n g l i a ' Engl and 3 6 . 2 5 ± 7 0 . 51 0 . 75 ±1.50 'N. Northern I t a l y 3 4 . 5 0 ± 3 5 . 91 0 . 75±0.96 i 1l y r i c a Yugoslav!a 40.00±65,00 0.33+0.58 i beri ca Spain 3 0 . 50±38.90 3 . 0 0 + 6 .0 0 aqui tana Centr al Massif o f France 2 4 . 0 0 + 2 8 .1 4 0 .0 0 rhodopaea Greece 3 4.7 5±4 3.89 3.00+1.58 armenea Turkey, Caucasus Mtns. 3 2.7 5+25.2 4 2.0 0+2 .71 Ita ly ' * Standard d e v i a t i o n . DISCUSSION CHEMICAL CONTROL.-The biology o f the pine candle moth is such t h a t the p o s s i b i l i t y o f developing an e f f i c i e n t i n t e g r a t e d co n tr o l program t h a t takes advantage o f the high r a t e o f n a t u r a l p a r a s i t i s m appears e x tr em e ly promising. Although Guthi o r ® , di a zin o n and methoxychlor gave e x c e l l e n t c o n tr o l o f the pine candle moth, the evidence i n d i c a t e s t h a t M e t a - S y s t o x - m a y be a more e f f e c t i v e m a t e r ia l M eta -Sys to x - in a co n tr o l program. In the pres ent study i g n i f i c a n t l y reduced the number o f i n f e s t e d buds w h i l e not completely e l i m i n a t i n g them; thus l e a v i n g breeding m aterial f o r the numerous p a r a s ite s t h a t a t t a c k £ . nepheos and a l lo w in g them to keep the pine candle moth po p u lati o n under an economic thresh old and po ssibly avoiding the n e c e s s i t y o f f u t u r e spray programs. In a p r e l i m i n a r y experiment in Ohio, Campbell (1968) found t h a t M e t a - S y s t o x - f ^ a p p a r e n t l y reduced the pine candle moth po pu la tio n by about 50% w h i le having l i t t l e parasites. e f f e c t on the In an i n t e g r a t e d control experiment on Scotch pine (Campbell and B a ld e r s to n , 1971) to c o n tr o l the pin e candle moth, the European pine s a w f l y , Neodiprion s e r t i f e r ( G e o f f r o y , and the s potte d pine aphid, Eulachnus a g i l i s (K a lte n b a c h ) Meta-Sy stox- was a p p l ie d in e a r l y May w h i l e the l a r v a e o f the pine candle moth were boring in the buds. This did not reduce the number o f l a rv a e i n f e s t i n g the candles but did reduce the number o f needle mining l a r v a e o f the ensuing g e n e r a t i o n . Furth erm ore, p a r a s i t i s m o f the l a r v a e was not lessened by t h i s t r e a t m e n t . 42 43 ATTRACTANTS.-The pheromone t e s t s i n d i c a t e t h a t the chemical, once i d e n t i f i e d , could be used as a survey too l to determine i n f e s t a t i o n s w i t h i n a stan d; thus avoiding unnecessary spraying o f u n i n f e s te d or l i g h t l y i n f e s t e d stands. The n e g a tiv e r e s u l t s o f the rele a se experiments i n d i c a t e s t h a t some p r e c o n d i t i o n i n g o f the pine candle moth may be necessary before the a d u l t male i s r e c e p t i v e to the pheromone. Daterman (1972) found t h a t the incid ence o f male sexual response o f Rhyacionia buo liana ( S c h i f f e r m u l e r ) depended upon the length o f tim e males were c o n d itio n e d a t high i l l u m i n a t i o n . HOST RESISTANCE. -The host r e s i s t a n c e s tudie s were in c o n c l u s iv e . I t appears t h a t o n l y under n a t u r a l c o n d it io n s could host r e s is t a n c e to the pine candle moth be v e r i f i e d . For example, W r ig h t , e t a l . (1966 and 1967) observed f o u r p l a n t a t i o n s t h a t had 108 s e e d lo ts each o f Pinus s y l v e s t r i s L. Neodiprion s e r t i f e r . i n f e s t e d w i th the European pine s a w f l y , They found t h a t the percentage o f t r e e s a t ta c k e d va rie d from 0 to 6 f o r north Eurasian v a r i e t i e s , 6 to 11 f o r south European v a r i e t i e s , and 12 to 26 f o r c e n t r a l European v a r i e t i e s . Something s i m i l a r may occur w ith the pine candle moth. Bennett (1954) r e p o r t e d a host r e s i s t a n c e f a c t o r w ith the pine needle m in e r, E x o t e l e i a p i n i f o l i e l l a species o f host t r e e s . (Chamb.), a t t a c k i n g various He found a c o r r e l a t i o n between the i n t e r n a l s t r u c t u r e o f a pine needle and i t s s u s c e p t i b i l i t y to i n j u r y . Of the many species t h i s pest a t t a c k s , lodgepole pine and j a c k pine were the most favored. The needles o f these 2 species have only 2 moderate­ s ized r e s i n canals l o c a t e d in each corner o f the ne ed le . Scotch p i n e , 44 on the o t h e r hand, was d e f i n i t e l y u n d e s ir a b le as a host t r e e with conspicuous r e s i n canals varying from 6 to 16; the h ig h e r the number, the g r e a t e r the res is t a n c e to the i n s e c t ap p a re n tl y due to the copious r e s i n f lo w . CONCLUSIONS AND RECOMMENDATIONS Although an estim ate o f peak a d u l t emergence o f the pine candle moth was made a t 1060+32 degree days above 52°F. thre sh old t e m p e r a tu re , the v a r i a b i l i t y involved in t h i s e stim ate appears too g r e a t t o be r e l i a b l e from y e a r to y e a r . T h e re f o r e , black l i g h t traps or p o s s ib ly pheromone t r a p s should be employed to determine peak e c lo s io n . An exact measure is important f o r determining the best time f o r con tr ol o p e ra t io n s . P e s t i c id e a p p l i c a t i o n s ten days a f t e r peak a d u l t emergence, o r a t the time o f egg hatch, gives good c o n t r o l . ot h er time in the l i f e c y c l e , the pest is w ell At any p r ote cted w i t h i n the needle o r candle mines and c o n t r o l s would be less e f f e c t i v e . Choice o f i n s e c t i c i d e is very important. Since the re are a la rg e number o f p a r a s ite s which a t t a c k t h i s species and since they appear to be o f some importance in keeping the population l e v e l s o f £. nepheos down, c o n s id e r a tio n should be given to the e f f e c t o f the p e s t i c i d e on the p a r a s i t e po pu la ti o ns. diazin on and methoxychlor a l l In the present study Guthioi@, gave e x c e l l e n t c o n t r o l . However, these m a t e r i a l s a l s o appeared t o have an adverse e f f e c t on the p a r a s i t e s . Meta-Systox- I ® on the o t h e r hand, gave less co n tr o l but also appeared to have less e f f e c t on the p a r a s i t e s . P o ssib ly t h i s m a t e r i a l can b r in g the pest po p u lati o n down to acceptable l e v e l s where i t can be maintained by the p a r a s i t e po pu lati o n. Future research should attempt to v e r i f y these statements by a long-t erm study on the e f f e c t s o f these m a t e r i a l s on both the pest and 45 46 p a r a s i t e p o p u la tio n s . This would in v o l v e an i n t e n s i v e population dynamics study t h a t would c l o s e l y examine the r e l a t i o n s h i p o f p a r a s i t e to p e s t , i n c l u d i n g when the p a r a s i t e s are p r e s e n t , which species are most im p o r t a n t , and what d e n s i t i e s would r e s u l t in a s i g n i f i c a n t reduction in the pest p o p u la tio n . Furthermore, m o r t a l i t y due to o v e r w i n t e r in g appears to be an im portant p a r t o f the l i f e h i s t o r y o f the pine candle moth and should be examined more c l o s e l y . Other t o o l s t h a t may be useful a t t r a c t a n t s and host r e s i s t a n c e . in a c o n tr o l program include This study has demonstrated t h a t a sex pheromone does e x i s t , but f u r t h e r research is needed to determine the exact chemical. The a t t r a c t a n t we used is not very powerful a t t r a c t i n g a d u l t males ov er s h o rt d is t a n c e s . in Perhaps the pheromone would have been s t r o n g e r when in a s s o c ia t io n w i th host v o l a t i l e s . If a strong a t t r a c t a n t can be i d e n t i f i e d and s y n t h e s iz e d , i t may prove o f immense va lue in surv eying f o r i n f e s t e d a re as . I f the pine candle moth should become an even more serious pest and become a t h r e a t to areas not now i n f e s t e d , surveying with pheromone t ra p s could p i n - p o i n t these areas r a p i d l y so t h a t prompt con tr ol measures can be taken. 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APPENDICES 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 MEASUREMENT ( mm.) pend1x la. Lengths of 191 needle mines excavated fcy lanae of the pine cardie ^cth. M IN E NEEDLE ol LENGTH AVERAGE Juno Appendix