The insect fauna of Canada thistle, Cirsium arvense (L.) Scop. in southern Montana by Hilde De Smet-Moens A thesis submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE in Biological Sciences Montana State University © Copyright by Hilde De Smet-Moens (1982) Abstract: Insects associated with Cirsium arvense (L.) Scop, in southern Montana are reported. Fifty-six phytophagous species and 47 visiting insects were collected, identified and tabulated. Four insect species, Corythucha distincta Osborn and Drake (Hemiptera: Tingidae), Bans sp., poss. cirsii Gilbert (Coleoptera: Curculionidae), Vanessa cardui L. (Lepidoptera: Nymphalidae), and Orellia ruficauda (Fabricius)(Diptera: Tephritidae), were considered conspicuous, because of their damage inflicted to the thistle plant. More insects were found associated with the developing seed heads than with foliage, stems or roots. The information gathered on this local survey can be valuable for future introductions of insect biological control agents. It forms the foundation for follow-up studies with indigenous insect species. Augmentation and redistribution of established monophagous insects, such as Ceuthorynchus litura and Baris sp. should be considered. Transmission experiments are necessary to evaluate the potential of these monophagous insects as thistle pathogen vectors. The combination of two stress-factors will increase the impact on the thistle plant in the field. STATEMENT OF PERMISSION TO COPY In p r e s e n t i n g t h i s t h e s i s in p a r t i a l f u l f i l l m e n t o f t h e r e q u i r e ­ ments f o r an advanced de gre e a t Montana S t a t e U n i v e r s i t y , I agree t h a t t h e L i b r a r y s h a l l make i t f r e e l y a v a i l a b l e f o r i n s p e c t i o n . I further agree t h a t pe rm is si o n f o r e x t e n s i v e copying of t h i s t h e s i s f o r s c h o l a r l y purposes may be g r a n t e d by my major p r o f e s s o r , o r , in h i s abse nc e, by t h e D i r e c t o r of L i b r a r i e s . I t is understood t h a t any copying or p u b l i ­ c a t i o n o f t h i s t h e s i s f o r f i n a n c i a l ga in s h a l l not be allowed witho ut my w r i t t e n p e r m is s i o n . S i g n a t u Ie Date THE INSECT FAUNA OF CANADA THISTLE, CIRSIUM ARVENSE (L .) SCOP IN SOUTHERN MONTANA by . . HILDE DE SMET-MOENS A t h e s i s sub mit te d in p a r t i a l f u l f i l l m e n t of t h e re q u ir e m e n ts f o r t h e degree of ' MASTER OF SCIENCE in B i o l o g i c a l Scie nce s Approved: MONTANA STATE UNIVERSITY Bozeman, Montana August, 1982 / / iii . ACKNOWLEDGMENTS I wish t o acknowledge and e x p r e s s my a p p r e c i a t i o n f o r t h e c o n t r i ­ b u t i o n s of t h e f o l l o w i n g people: Dr. W. L. M o r r i l l , my major p r o f e s s o r , f o r h i s p r o f e s s i o n a l a s s i s t ­ ance th ro u g h o u t t h i s r e s e a r c h p r o j e c t ; The members of my t h e s i s committee. Dr. P. K. F a y , Dr. S. R. Eversman, and Mr. J . M. S t o r y , f o r t h e i r tim e and i n v a l u a b l e a dvice ; Mrs. S. D. Rose, C u r a t o r of t h e M.S.U. Entomological C o l l e c t i o n , f o r h e r taxonomic a d v ic e and he lp ; All t h e s y s t e m a t i c i s t s who i d e n t i f i e d t h e i n s e c t s mentioned in t h i s r e p o r t : t h e t a x o n o m i s t s . o f t h e Sy s te m a ti c Entomology L a b o ra to ry , USDA, B e l t s v i l . l e , MD; and t h e N a ti o n al Museum of Na tural H i s t o r y , Smithsonian I n s t i t u t i o n , Washington, DC; J . L a t t i n , P. Oman, K. A. P h i l l i p s , and G. M. S t o n e d a h l , Oregon S t a t e U n i v e r s i t y , C o r v a l l i s ; L. A. K e l t o n , Bios y s t e m a t i c s Research I n s t i t u t e , Ottawa; M. W. N ie ls o n , Forage I n s e c t s Research L a b o r a to r y , Tucson, AZ; G. J . M ic he ls , J r . , Texas A&M Univer­ s i t y , A m a ril lo ; R. J . B e s h e a r , The U n i v e r s i t y of Ge orgia, Experiment; J . A. Onsager and E. A. Oma, U.S.D.A. Rangeland I n s e c t La b o ra to ry , Bozeman; R. M. B o h a r t , U n i v e r s i t y o f C a l i f o r n i a , Davis; D. K. Young, Michigan S t a t e U n i v e r s i t y , Ea st Lansing; . The Western A g r i c u l t u r a l Research C en t er and t h e Montana weed d i s t r i c t s f o r t h e fun din g of t h i s r e s e a r c h p r o j e c t . TABLE OF CONTENTS Page VITA .................................................... ACKNOWLEDGMENTS i.i ...................... iii LIST OF TABLES..................................................................... LIST OF FIGURES . . . . .............................................................................. vi ABSTRACT ................................................................. vii INTRODUCTION.................................................... LITERATURE REVIEW .................... I . . . . . . . . 3 CO (T i The Host P l a n t ............................................ ..................... B i o l o g i c a l Co ntrol of Canada T h i s t l e .................. B i o l o g i c a l Co ntrol of Canada T h i s t l e in Montana MATERIALS AND METHODS v . , ...................... - . ....................... .... 12 Study S i t e s ..................................................................................................... C o l l e c t i n g M e t h o d s ........................................... Experimental R e a r i n g s ................................................ .... . ' .................. F i e l d S t u d i e s ..................... I n s e c t I d e n t i f i c a t i o n .................. RESULTS AND DISCUSSION.................................. . Phytophagous I n s e c t s .. ............................................ . V i s i t o r s , P r e d a t o r s and P a r a s i t o i d s . . . . . . . . . . . . Summ ary...................... .....................................................■ LITERATURE CITED .......................................................................... 12 12 15 16 16 18 18 38 42 46 V LIST OF TABLES Table 1. 2. 3. 4. 5. 6. 7. Page In fo rm a tio n on t h e i n s e c t s r e l e a s e d in Montana f o r t h e b i o l o g i c a l c o n t r o l of Canada t h i s t l e ............................... 10 S e l e c t e d c o l l e c t i o n s i t e s of t h e 1981 i n s e c t survey on Canada t h i s t l e ...................... ...................................... .... 14 Phytophagous i n s e c t s c o l l e c t e d from Canada t h i s t l e , Cirsium a ry e ns e (L .) S c o p . , in s o u th e r n Montana, 1981 . . . . ............................... .............................. 19 Average p l a n t h e i g h t s o f e i g h t B a r is i n f e s t e d and e i g h t uni n f e s t e d t h i s t l e p l a n t s in s i t e s 5 and 6, June 19, 1981 ............................... . . ................................... 30 I n f e s t a t i o n o f Canada t h i s t l e heads by O r e l l i a r u f i c a u d a ( F a b r i c i u s ) , September 3, 1981 . . . ........................... 37 V i s i t o r s , p r e d a t o r s , and p a r a s i t o i d s c o l l e c t e d from Canada t h i s t l e , Cirsium a rv e nse (L .) Scop. in so u th er n Montana, 1981 T ...................... ..................... ..................... 39 P a r a s i t i s m of O r e l l i a r u f icauda ( F a b r i c i u s ) pupae in Canada t h i s t l e seed head s, S e p t e m b e r ^ , 1981 43 . . . vi LIST OF FIGURES F ig u r e 1. Page C o l l e c t i o n s i t e s o f t h e i n s e c t survey on Canada t h i s t l e , 1 9 8 1 ............................................................................... 13 2. Emergence t r a p in s i t e 5 . . . ................................. ........................ 17 3. Corythucha d i s t i n c t a a d u l t s f e e d i n g on Canada t h i s t l e le av e s . . ............................................ ..................... 23 Feeding damage o f Corythucha d i s t i n c t a on Canada t h i s t l e . . ............................................... 23 4. 5 . • Aggregation of Corythucha d i s t i n c t a nymphs on t h e u n d e r - s i d e o f Canada t h i s t l e le ave s . . . . . . . . 25 6. B a r is sp. a d u l t fe e d i n g on t h i s t l e r o s e t t e ............................... 28 7. Feeding damage o f B a r is sp. a d u l t s on p o tt e d t h i s t l e p l a n t s in t h e i n s e c t a r y ...................... .... 31 8. B ar is s p. l a r v a in Canada t h i s t l e r o o t ...................... 31 9. Wilted t h i s t l e p l a n t in s i t e 5, i n f e s t e d with B ar is s p . l a r v a e .......................................................................... 31 L i n e a r r e g r e s s i o n of p l a n t h e i g h t v e rs us l a r v a l f r e q u e n c y , August 19 8 1 ........................................................ 33 Feeding damage o f O r e l l i a r u f i c a u d a . l a r v a e on Canada t h i s t l e seeds. . -................................... .. ........................... 36 P a r a s i t i z e d O r e l l i a r u f i c a u d a pupa . . . ................................... 44 10. 11. .12. . ABSTRACT I n s e c t s a s s o c i a t e d with Cirsium a rv e nse ( I . ) Scop, in south ern Montana a r e r e p o r t e d . F i f t y - s i x phytophagous species, and 4 7 . v i s i t i n g i n s e c t s were c o l l e c t e d , i d e n t i f i e d and t a b u l a t e d . Four i n s e c t s p e c i e s , Corythucha d i s t i n c t a Osborn and Drake (Hemiptera: T i n g i d a e ) , B a n s s p . , p o s s . c i r s i i G i l b e r t ( C ole opte ra : C u r c u l i o n i d a e ) , Vanessa^cardui L. ( L e p i d o p t e r a: Nyrnphalidae),. and O r e l l i a r u f i c a u d a ( F a b r i c i u s ) ( D i p t e r a : T e p h r it i d a e ) , were c o n s i d e r e d , c o n s p ic u o u s , because of t h e i r damage i n f l i c t e d t o t h e t h i s t l e p l a n t . More i n s e c t s were found a s s o c i a t e d with t h e - d e v e l o p i n g seed heads th a n with f o l i a g e , stems or r o o t s . The i n fo r m a t io n g a th e r e d o h . t h i s lo c a l survey can be v a l u a b l e f o r f u t u r e i n t r o d u c t i o n s of in s ec t, b i o l o g i c a l c o n t r o l a g e n t s . I t forms t h e fo u n d a ti o n f o r fo ll o w - u p s t u d i e s with indigenous i n s e c t s p e c i e s . Augmentation and r e d i s t r i b u t i o n of e s t a b l i s h e d monophagous i n s e c t s such as Ceuthorynchus l i t u r a and B a r is sp. should, be c o n s i d e r e d . Trans mi ssi on ex periments a r e n e c e s s a r y t o e v a l u a t e t h e p o t e n t i a l of these ' monophagous ins ects, as t h i s t l e pathogen v e c t o r s . The combination of . two s t r e s s - f a c t o r s w i l l i n c r e a s e t h e impact on t h e t h i s t l e p l a n t in . the f i e l d . - INTRODUCTION C u l t u r a l and chemical c o n t r o l p r a c t i c e s have h i s t o r i c a l l y been t h e main approaches t o weed c o n t r o l . Both methods a r e aimed a t removing unwanted p l a n t s as q u i c k l y as p o s s i b l e , a s h o r t term approach r e q u i r i n g c o n s i d e r a b l e annual e x p e n d i t u r e s of r e s o u r c e s and energy (Andres and Gpeden, 1971). B i o l o g i c a l c o n t r o l o f weeds has become a p o p u la r a l t e r n a ­ t i v e because i t i s a means o f c o n t r o l l i n g weeds w i t h o u t t h e high energy c o s t s o f c u l t u r a l p r a c t i c e s and w i t h o u t t h e r e s i d u e and p o l l u t i o n prob­ lems o f h e r b i c i d e s . B i o l o g i c a l c o n t r o l o f weeds i s t h e d e l i b e r a t e use of i n s e c t s or o t h e r p l a n t p a r a s i t e s t o reduce t h e d e n s i t y o f a weed t o an a c c e p t a b l e l e v e l ( H a r r i s , 1971b). B i o c o n t r o l , when e f f e c t i v e , i s r e l a t i v e l y in e x­ p e n s i v e , l o n g - l a s t i n g and t h e b e n e f i t s a r e c u m u la ti v e . This approach has s t r e n g t h s and weaknesses d i f f e r e n t from o t h e r methods and hence i s advantageous under c e r t a i n c o n d i t i o n s ( H a r r i s , 1971b; Maw, 1982). c ro ps of^arablle land lhave many s p e c i e s of weeds as c o m p e t i t o r s . c o n t r o l l i n g one s p e c i e s through b i o l o g i c a l Most Thus, c o n t r o l would r e q u i r e s p r a y ­ ing o r c u l t i v a t i o n f o r c o n t r o l o f t h e o t h e r s p e c i e s . On t h e o t h e r hand, dominance o f one weed s p e c i e s i s t y p i c a l on range lan d. Such a domi­ na nt weed i s a very s u i t a b l e s u b j e c t f o r b i o l o g i c a l c o n t r o l . Even a s l i g h t i n c r e a s e in p r e s s u r e can have a s i g n i f i c a n t e f f e c t ( H a r r i s , 1971b). One o f t h e i n i t i a l s t e p s in de vel opin g a b i o l o g i c a l weed c o n t r o l program i s t o de te r m in e t h e n a t u r a l enemies a t t a c k i n g t h e weed s p e c i e s , 2 in both i t s n a t i v e and i t s p r e s e n t geog raphic ranges (Maw, 1980; H a r r i s , 1971a). I n i t i a l surveys expand i n s i g h t and un de rs ta ndi ng o f t h e weed e co lo gy , h o s t r a n g e s , e t h o lo g y and i n s e c t - h o s t i n t e r r e l a t i o n s h i p s . The i n fo r m a t io n g a t h e r e d in t h e l o c a l s u r v e y s . i n d i c a t e s t h e n i c h e s occupied \ ' ■ by indigenous s p e c i e s ^ so t h a t chances of i n t r o d u c i n g a b i o c o n t r o l agent t h a t may d u p l i c a t e o r compete with an a l r e a d y p r e s e n t s p e c i e s a r e m i n i ­ mized (Maw, 1976). The o b j e c t i v e s of t h i s study were: I) t o det ermi ne t h e endemic i n s e c t fauna a s s o c i a t e d with t h e d i f f e r e n t growth s t a g e s of Canada t h i s t l e , Cirsium a rv e ns e ( I . ) S c o p . , in Southern Montana, and 2). t o e v a l u a t e t h e damage i n f l i c t e d by t h e most conspicuous s p e c i e s . LITERATURE REVIEW The Host P l a n t Cirsium a rv en se ( L , ) Scop, i s a troubl eso me p e r e n n i a l weed in Montana. Indigenous t o Europe, Western Asia and Northern A f r i c a , i t was pr ob a bl y in tr odu ce d t o North America in t h e 17th Century (Detmers, 1927; Pe sch ke n, 1971). I n f e s t a t i o n s o f Canada t h i s t l e now o c cu r th rou ghout t h e a g r i c u l t u r a l a r e a s of Canada and t h e n o r t h e r n h a l f o f t h e United S t a t e s ( Pes chk en , 1971; H itc hco ck , e t a l . , 1973). A r e c e n t survey i n d i ­ c a t e d t h a t t h i s troubl eso me weed i n f e s t s 1.5 m i l l i o n a c r e s in Montana.1 Canada t h i s t l e damages a wide v a r i e t y of crops by c o m p e t i t i v e use o f l i g h t , m o i s t u r e and n u t r i e n t s (Hodgson, 1977). Heavy i n f e s t a t i o n s in p a s t u r e s and ranges reduce f o r a g e y i e l d s c o n s i d e r a b l y . The weed a l s o h a r b o r s i i n s e c t s t h a t a t t a c k economic crops and is an a l t e r n a t e ho s t f o r some p a th oge nic organisms (Moore, 1975). Cirsium a rv e ns e ( L . ) Scop, i s a polymorphic s p e c i e s . and f l o w e r c h a r a c t e r i s t i c s vary c o n s i d e r a b l y . Stem, l e a f B o t a n i s t s u s u a l l y reco g­ n i z e t h r e e o r f o u r morphological v a r i a n t s , a l l i n t e r b r e e d i n g f r e e l y (Moore and F r a n k t o n , 1974; Hodgson, 1964; Detmers, 1927). Canada t h i s t l e i s d i o e c i o u s and r e p r o d u c t i o n occurs from seed and rhizome p i e c e s . Detmers (1927) s t a t e d t h a t honey bees were t h e c h i e f p o l l i n a t i o n a g e n t s . I Jackson,: M. J . , 1982. Perso na l communication. 4 In Montana, t h e t h i s t l e p l a n t s emerge in e a r l y May, when t h e mean weekly a i r te m p e r a t u r e reach es 5°C (Moore, 1975). R o s e t t e s a r e formed followe d by stem e l o n g a t i o n app rox imate ly t h r e e weeks a f t e r emergence (Moore, 1975). September. Flowering be gin s in mid-June and c o n t i n u e s i n t o Growth be gi ns d e c r e a s i n g in J u l y , and c e a s e s by e a r l y August The obnoxious c h a r a c t e r of t h i s weed is due mainly t o t h e rapid v e g e t a t i v e p r o p a g a t i o n o f i t s c r e e p i n g h o r i z o n t a l rhizomes, giv in g r i s e t o numerous a e r i a l sh oots (Moore, 1975). This e x t e n s i v e branched rhizome system makes Canada t h i s t l e d i f f i c u l t t o c o n t r o l , Cultural, mechanical and chemical c o n t r o l methods can be e f f e c t i v e in c u l t i v a t e d f i e l d s i f used p e r s i s t e n t l y in a long range c o n t r o l program (Hodgson, 1977). Where Canada t h i s t l e i s a p r e v a l e n t weed in range la n d , f a ll o w f i e l d s , waste l a n d s , r o a d s i d e s o r r a i l w a y s , b i o l o g i c a l c o n t r o l by i n ­ s e c t s can be a re a s o n a b l e a d j u n c t . These h o s t s p e c i f i c i n s e c t s are harmless t o n o n - t a r g e t p l a n t s and may be a b l e t o m u l t i p l y and d i s p e r s e to adjacent in f e s ta tio n s . B i o l o g i c a l Control of Canada T h i s t l e The p r i n c i p l e s and proc ed ures o f b i o l o g i c a l weed c o n t r o l have been well d e f i n e d and i l l u s t r a t e d with some s p e c t a c u l a r s u c c e s s e s (Be Bach, 1964; Van den Bosch, e t a l . , 1982). B i o l o g i c a l c o n t r o l s t r i v e s to reduce t h e abundance of a weed s p e c i e s by i n tr o d u c in g o r augmenting t h e weed's n a t u r a l enemies. Huffaker (1959) l i s t s examples in which 5 n a t u r a l l y o c c u r r i n g i n s e c t s have played an impor tant r o l e in a f f e c t i n g t h e abundance o f a p a r t i c u l a r p l a n t s p e c i e s . . The i n t r o d u c t i o n of ho s t s p e c i f i c phytophagous organisms has r e c e i v e d t h e most emphasis t o d a t e (Andres, e t a l . , 1976).. The s t e p s involved in such a t e c h n i q u e are d e s c r i b e d by H a r r i s (1971b). ■ Cirsium a rv e ns e ( L.) Scop, i s a prime c a n d i d a t e f o r b i o l o g i c a l con t r o l (Hume,.1982; H a r r i s , 1971b, A ndr e s , e t a l . , 1976) because: 1. The p l a n t has l i t t l e o r no va lu e t o people o r w i l d l i f e . 2. I t t e n d s t o grow in dense p o p u l a t i o n s , r e p r e s e n t i n g a dominant weed in p a s t u r e and waste a r e a s . 3. I t i s an in tr o d u c e d weed, and has very few p a r a s i t e s and predators. 4. I t i s not c l o s e l y r e l a t e d t o major economic crop; p l a n t s ; however a r t i c h o k e (Cynara scolymus I . ) and s a f f l o w e r ( C a r t h amus t i n c t o r i us L.) belong t o t h e same Cyhareae tribe. 5. Many of t h e t h i s t l e i n f e s t a t i o n s occu r in i n a c c e s s i b l e . a r e a s and th u s lend th em sel ve s t o b i o c o n t r o l e f f o r t s ( S t o r y , 1980). 6. The widespread d i s t r i b u t i o n of t h e weed w i l l r e s u l t in low c o s t p e r a c r e of c o n t r o l . T h is.c o st per acre will be lower tha n t h a t of o t h e r c o n t r o l methods ( H a r r i s , 1979). 6 B i o l o g i c a l c o n t r o l w i l l i n c r e a s e t h e environmental p r e s s u r e on Canada t h i s t l e . At. b e s t , t h e n a t u r a l enemies may e l i m i n a t e t h e need f o r o t h e r c o n t r o l methods o ve r much o f t h e p l a n t ' s range and form a sound b a s i s f o r f u t u r e weed management schemes. At l e a s t , t h e y would augment . e x i s t i n g c o n t r o l p r a c t i c e s ('Batra-, e t a l . , 1981) . The Commonwealth I n s t i t u t e of B i o l o g i c a l Control began work on t h e . c o n t r o l o f Canada t h i s t l e in 1961, with a stu dy of i t s p a r a s i t e s in Europe ( Pe sch ken , 1971). Eighty i n s e c t :s pe c ie s were found t o feed on t h e weed, A l t i c a c ar d u o ru m -Guer. ( C o l e o p t e r a : Chrysomeli d a e ) , Ceutorhynchus l i t u r a (F ab. ) ( C o le o p te ra : C u r c u li o n id a e ) and Urophora c a r d u i ( I . ) ( D i p t e r a : T e p h r i t i d a e ) were s e l e c t e d f o r f u r t h e r study b e ­ cause o f t h e i r a p p a r e n t h o s t s p e c i f i c i t y (Peschken, 1971; Z w o lf e r , 1964). Al I t h r e e i n s e c t s have been r e l e a s e d in Canada and t h e United S t a t e s . .. A l t i c a carduorum Guerin f a i l e d t o . e s t a b l i s h in a l l r e l e a s e s i t e s due t o c l i m a t i c s t r e s s and a t t a c k by i n s e c t p r e d a t o r s ( P es c hk en , e t a l . , 1970). Ceutorhynchus l i t u r a ( F .) i s e s t a b l i s h e d in a wide range o f c l i ­ mates in Canada (Peschken, e t a l . , 1980), Montana and Idaho; however, t h e range of i n f e s t a t i o n i s i n c r e a s i n g very slo wly . While t h i s weevil e x e r t s s t r e s s on i t s h o s t in t h e l a b o r a t o r y (Peschken and Beecher, 1973) t h e r e i s no ev ide nc e t h a t C,. l i t u r a c o n t r o l s Canada t h i s t l e in t h e f i e l d (Peschken, et, a l . , 19Q1). 7 Urophora c ar du i ( L . ) has. become e s t a b l i s h e d and i s sp re ad in g in e a s t e r n Canada ( P es c hke n, e t a l . , 1980). A microsporidian disease of LL c ar du i (Nosema s p . ) i s one o f t h e causes le adi ng t o f a i l u r e of e s t a b l i s h m e n t in t h e Western s t a t e s . Other r e a s o n s , such as i n f e r t i l i t y o f t h e f l i e s , o r s p r i n g f r o s t k i l l i n g o f t h e l a r v a e may have been a d d i ­ t i o n a l causes of m o r t a l i t y (Peschken, e t a l . , 1982). Ap pa rentl y t h e t w o . e s t a b l i s h e d in tr od uce d i n s e c t s w i l l not c o n t r o l Canada t h i s t l e . F u r t h e r s t r e s s f a c t o r s from o t h e r i n s e c t s , pathogens o r p l a n t c o m p e t it io n a r e needed t o c o n t r o l t h i s weed. Few p o t e n t i a l l y e f f e c t i v e and h o s t s p e c i f i c i n s e c t s from Z w o l f e r 's l i s t (1964) a r e s t i l l a v a i l a b l e . T i n g i s am plia ta H.-S. ( H e t e r o p t e r a: T in gi da e ) was r e c e n t l y t e s t e d f o r h o s t s p e c i f i c i t y in t h e l a b o r a t o r y . I t was c o n s id e r e d uns af e f o r i n t r o d u c t i o n in Canada, because i t s h o s ts i n c lu d e s a f f l o w e r and globe a r t i c h o k e . Lema c y a n e l l a ' ( I . ) The approval f o r r e l e a s e of ( C o le o p te r a : C u r c u l i o n i d a e ) has been w it h h e ld be­ cause i t a t t a c k s s e v e r a l Cirsium spp. indigenous t o North America (Peschken, e t a l . , 1980). The p r e s e n t concern f o r n a t i v e Cirsium s p e c ie s has become a c r i t i ­ cal issue. Had such s t r i n g e n t host s p e c i f i c i t y req u ir e m e n ts been a p p l i e d in t h e p a s t , a number of i n s e c t s would never have been r e l e a s e d . This f e a r f o r n a t i v e f l o r a could lengthe n t h e s c r e e n in g p r o c e s s con­ s i d e r a b l y and many promising agen ts could be r e j e c t e d . B i o l o g i c a l con­ t r o l could th e n become i m p r a c t i c a l a n d .v e r y ex p en s iv e . Peschken (1982) 8 reviewed s u c c e s s f u l b i o l o g i c a l weed c o n t r o l p r o j e c t s and concluded t h a t no t a r g e t weed has e v e r become r a r e . He s t a t e d t h a t i t i s very u n l i k e l y t h a t n a t i v e p l a n t s , which a r e in e q u i l i b r i u m with t h e i r own i n s e c t fa u n a , would su p p o rt a d d i t i o n a l i n s e c t s p e c i e s . P o p u l a t i o n s of Canada t h i s t l e a r e a l s o a t t a c k e d by numerous in d i g e n ­ ous i n s e c t s and pa th o g en s . Nat ura l enemies, a s s o c i a t e d with Canada t h i s t l e in North America have been r e p o r t e d by Moore (197 5) , Maw (1976), Watson, e t a l . (198 0) , Detmers (1927), and Andres (1980). Among t h e most imp or tan t s p e c i e s were: a. The p a i n t e d lady b u t t e r f l y , Vanessa c a r d u i L. ( =Pyrameis cardui L . , =Cynthia c ar du i ( I . ) ) ( L e p i d o p t e r a : NymphaTidae). The l a r v a e o c c a s i o n a l l y cause s p e c t a c u l a r d e f o l i a t i o n o f t h i s t l e s in l o c a l a r e a s . . I t i s a m i g r a to r y b u t t e r f l y , and i t s numbers f l u c t u a t e a n n u a l l y , making i t u n r e l i a b l e as a n a t u r a l c o n t r o l agent. I t can be a p e s t of sunf lower and soybean (Morihara. and. Balsbaugh, 1976) and many o t h e r p l a n t s . b. The Canada t h i s t l e midge Dasyneura g ib s o n i F e l t ( D i p t e r a : Cecidomyii d a e ) , which a t t a c k s t h e de vel opin g seed c. heads. O r e l l i a r u f i c a u d a ( F . ) ( =Trypeta f l o r e s c e n t i a e L.) ( D i p t e r a : T e p h r i t i d a e ) which a t t a c k s up t o 70% o f t h e t h i s t l e he ads . This seed head f l y was probably a c c i d e n t a l l y in tr oduc e d from Europe ( H a r r i s , 1971a). 9 d. Cassida r u b i g i n o s a MuelI . ( C d le o p te ra : Chrysomeli d a e ) : This b e e t l e was a c c i d e n t a l l y in tr od uce d i n t o t h e United S t a t e s (Ward, 1976), and d e f o l i a t e s Canada t h i s t l e a t high po p u la ­ tions. This f l e a b e e t l e has become widely e s t a b l i s h e d in t h e E a s te r n United S t a t e s (Ward, 1976). e. The systemic a u to e c io u s r u s t P u c c in ia obtegens (Link) T u l . is an endemic patho gen , h o s t s p e c i f i c t o Canada t h i s t l e . N a tu r­ a l i n f e c t i o n i s no t high enough f o r economic c o n t r o l . Since host r e s i s t a n c e i s an im por tan t f a c t o r l i m i t i n g r u s t i n f e c t i o n , an a g g r e s s i v e s t r a i n may be more e f f e c t i v e as a b i o c o n t r o l agent o f Canada t h i s t l e (T u r n e r , 1981). None o f t h e l i s t e d s p e c i e s su pp re ss Canada t h i s t l e p o p u l a t i o n s below t h e economic l e v e l . However, i n v e s t i g a t i o n s of t h e bio lo g y o f t h e most d e s t r u c t i v e organisms could r e s u l t in t h e development o f approaches whereby p o p u l a t i o n s o f t h e s e n a t u r a l enemies a r e augmented t o i n c r e a s e t h e i r damage t o Canada t h i s t l e (Watson, e t a l . , 1980; Ward, 1976; Tu r n e r , 1981). B i o l o g i c a l Co ntrol of Canada T h i s t l e in Montana The b i o l o g i c a l c o n t r o l program o f Canada t h i s t l e in Montana began with t h e l i b e r a t i o n of A l t i c a carduorum Guerin in 1964. Sub se quen tl y, t h e stem weevil Ceutorhynchus l i t u r a ( F . ) was r e l e a s e d in 1973 and Urophora c ar du i (L.), in 1978 ( S t o r y , 1979) (Table I ) . Of t h e t h r e e , Table I . In f orm a tio n on t h e i n s e c t s r e l e a s e d in Montana f o r t h e b i o l o g i c a l c o n t r o l o f Canada t h i s t l e . a Insect A ltica carduorum Guerin Date Insects Released No. Insects Released County where Released Source of Insect Status . of Insect 1964 1966 200 200. R av a ll i Gallatin USDA USDA No recovery No rec overy Ceutorhynchus litu ra (F .) 1973 200 Gallatin . USDA Inc. slowly Urophora c ar du i ( I . ) 1978 92 R a v a ll i USDA No recovery ^Data from S t o r y , J . M., 1979. 11 onl y Ceutorhynchus l i t u r a ( F . ) has su rv iv e d and become e s t a b l i s h e d . Attempts t o r e d i s t r i b u t e t h e weevil were made in 1977. a t t h e Bozeman s i t e , when 95% of t h e s hoo ts were found t o be mined ( S t o r y , 1980). P l a n t s i n f e s t e d with C_. l i t u r a l a r v a e were c o l l e c t e d in June from t h e ■ r e l e a s e s i t e and t r a n s p l a n t e d in o t h e r a r e a s of Montana. The re a so n s f o r f a i l u r e of A I t i c a carduorum Guerin and Urophdra c a r d u i ( L . ) t o e s t a b l i s h in Montana a r e not known. t o e s t a b l i s h jJ. c a r d u i A d d i t i o n a l a tt e m pts w i l l be made as more i n s e c t s become a v a i l a b l e ( S t o r y , 1980). The b i o l o g i c a l weed c o n t r o l . p r o g r a m in Montana i s i n c r e a s i n g in momentum. I n t e r e s t and awareness by t h e p u b l i c and academic communities a re growing. Endemic p l a n t pathogen's, p a r t i c u l a r l y P u c c in ia obtegehs I (T ur ne r, 1981) and S c l e r o t i n i a s c l e r o t i o r u m (Simmonds, 1982) a re being i n v e s t i g a t e d as p o t e n t i a l b i o c o n t r o l ag en ts a g a i n s t Canada t h i s t l e in Montana. The use of p l a n t pat hogen s may extend t h e a p p l c i a t i o n of b i o ­ l o g i c a l weed c o n t r o l t o i n c lu d e c u l t i v a t e d a r e a s . MATERIALS AND METHODS Study S i t e s A t o t a l of 50 s i t e s were surveyed th ro ughou t t h e 1981 growing s e a ­ son, s t a r t i n g A p ri l 15 and ending September 30 (F ig ur e I).. This pe ri o d co ver s r o s e t t e , v e g e t a t i v e , f lo w e r in g and f r u i t i n g s t a g e s o f t h e t h i s t l e plant. Most o f t h e l o c a t i o n s r e p r e s e n t o c c a s i o n a l c o l l e c t i o n s i t e s . Seven were s e l e c t e d f o r more i n t e n s i v e weekly o r bi-m onthl y sampling (T able 2 ) . S e l e c t i o n o f t h e s e s i t e s was based on e x t e n t o f t h i s t l e i n f e s t a t i o n , h a b i t a t - t y p e , a c c e s s i b i l i t y and absence o f h e r b i c i d e and p e s t i c i d e u s e. C o l l e c t i n g Methods The samples were c o l l e c t e d by t h e f o ll o w i n g methods: 1. . Stand s .of Canada t h i s t l e p l a n t s were examined c a r e f u l l y f o r i n s e c t i n f e s t a t i o n s and e x t e r n a l symptoms of endophagy. Feeding o r o v i p o s i t i o n damage was recorded in t h e f i e l d and ' whenever p o s s i b l e c o r r e l a t e d with t h e i n s e c t s p e c i e s p r e s e n t . 2. Most of t h e i n s e c t s were c o l l e c t e d by h a n d p i c k i n g , t h e only method a llo w ing c o r r e c t l o c a l i z a t i o n of t h e sampled specimens. Sweeping d i s t u r b e d t h e i n s e c t s and did not always d i s l o d g e i n d i v i d u a l s on upper and lower p a r t s o f t h e p l a n t . 3. O c c a s i o n a l l y , a simple p o l y e t h y l e n e - b a g s a m p l in g .t e c h n iq u e was used (Trumble, e t a l . , 1975). A l a r g e p o l y e t h y l e n e bag M ' W O u I* # : occas ional c o l l e c t i o n s i t e s O : s e le c te d c o l l e c t i o n s i t e s Fi gu re I . C o l l e c t i o n s i t e s of t h e i n s e c t survey on Canada t h i s t l e , 1981. 14 Table 2. Site S e l e c t e d c o l l e c t i o n s i t e s o f t h e 1981 i n s e c t survey on Canada thistle. Location Habitat. I Agronomy Farm, Bozeman ( G a l l a t i n County) Mowed g r a s s l a n d with n a t u r a l r u s t i n f e s t a t i o n (P. ob te ge ns ) o f 52%. (T u rn e r, 198IT. 2 Fort E l l i s ( G a l l a t i n County) Fallow f i e l d p l a n t e d with wheat and b a r l e y in 1980 3. Southern Research Cent er Fallow f i e l d , b o r d e r in g b a r l e y Huntley (Yellowstone County) f i e l d . 4, Waste a r e a , shaded by high t r e e s Southern. Research C en te r Huntley (Yellowstone County) 5 Billings (Yellowstone County) H ills id e , natural h a b ita t 6 Columbus ( S t i l l w a t e r County) D is tu rb e d a r e a , r o a d s i d e 7 Park C it y ( S t i l l w a t e r County) Roadside, bord e ri n g p a s t u r e l a n d 15 was i n v e r t e d o ve r t h e t a r g e t p l a n t s and f a s t e n e d a t t h e open end. P la n ts , were then upr o o te d , la b e l e d and t r a n s p o r t e d , t o t h e l a b o r a t o r y ' f o r exami nat ion . 4. Two t o f i v e t h i s t l e p l a n t s were s e l e c t e d a t random and uprooted . a t each s i t e . Roots, crowns and stems were d i s s e c t e d arid ex­ amined f o r endophagous i n s e c t s . 5. Flower heads and buds were d i s s e c t e d in t h e f i e l d . By t h e end o f t h e growing s ea s on , c o l l e c t i o n s o f seed heads were made a t' different site s. Approximately 50% o f t h e seed heads were ex­ amined in. t h e l a b o r a t o r y and t h e r e s t were s t o r e d in po ly e th y Iene bags t o r e c o v e r emerging a d u l t s o f endophagous s p e c i e s . Experimental Rearings . Immature i n s e c t s were r e a r e d t o t h e a d u l t s t a g e on p o t t e d p l a n t s o r f r e s h c u t Canada t h i s t l e f o l i a g e in the. i n s e c t a r y . Conspicuous i n s e c t s such as B a r is s p . , p os s. c i r s i i G i j b e r t 1 and Corythucha d i s t i n c t a Osborn and Drake were c o l l e c t e d in c o n t a i n e r s and c on fi ne d t o p o t t e d p l a n t s in the. i n s e c t a r y . Canada t h i s t l e p l a n t s was observed d a i l y . T h e i r impact on p o t t e d Cages used t o c o n f i n e i n s e c t s were c y l i n d r i c a l in shape (22 x 38 cm), and had wooden frames covered by a f i n e mesh c l o t h . ---------------- - r — --------- :------------------------ :----------------------------- Whitehead, D. R. (S yst em at ic Entomological L a b o r a to r y , USDA) . could not supply a p o s i t i v e i d e n t i f i c a t i o n . . We w i l l r e f e r t o i t . as Baris sp. - 16 Field Studies S i t e s 5 a n d . 6 (Table 2) were s e l e c t e d f o r o b s e r v a t i o n s of Baris sp. ( C o le o p te r a : C u r c u l i o n i d a e ) on Canada t h i s t l e . Both s i t e s were h e a v i l y i n f e s t e d with t h e weevil and i t s h o s t . 1. He ights of e i g h t damaged t h i s t l e p l a n t s were rec ord ed in mid-June a t each s i t e and compared with h e i g h t s o f e i g h t u n i n f e s t e d p l a n t s a t t h e same s i t e . D i f f e r e n c e s were t e s t e d f o r s i g n i f i c a n c e with t h e t - t e s t (P < 0 . 0 1 ) . 2. At t h e end of t h e growing sea so n, 17 damaged t h i s t l e p l a n t s were u p ro o te d , d i s s e c t e d , and pupal counts were made. A regression a n a l y s i s was performed o f t h e p l a n t h e t g h t over t h e l a r v a l f r e ­ quency ( P<- 0 . 0 1 ) . 3. Emergence t r a p s , each co verin g ap proxim ately two p l a n t s , were p la ce d in s i t e 5, in e a r l y September, 1981. These c a g e s , ma de,., o f f i n e w i r e - n e t t i n g , were c o n i c a l in shape with a dia m e te r of 80 cm. A small t r a p was a t t a c h e d on t h e top ( F ig u r e 2 ) . These t r a p s made i t p o s s i b l e t o c o n t r o l seaso nal a c t i v i t y of B aris a d u l t s and t o c o l l e c t o t h e r i n s e c t s p e c i e s emerging from t h e same t h i s t l e p l a n t s . Insect Id e n tifica tio n I n s e c t specimens were s o r t e d and s e n t t o taxonomic a u t h o r i t i e s for identification. 17 Fi gu re 2. Emergence t r a p in s i t e 5. RESULTS AND DISCUSSION The i n s e c t s c o l l e c t e d on Canada t h i s t l e were grouped as I) phyto­ phagous i n s e c t s (Table 3 ) ; 2) i n s e c t s c o l l e c t i n g p o l l e n o r n e c t a r , i n ­ s e c t s which a r e i n c i d e n t a l v i s i t o r s , p r e d a t o r s and p a r a s i t o i d s o f o t h e r insects. I n s e c t s c o l l e c t e d a r e l i s t e d by o r d e r acc ord ing t o B o rr o r , De Long and T r i p l e h o r n (1976). Phytophagous I n s e c t s Table 3 l i s t s 58 s p e c i e s , r e p r e s e n t i n g s i x o r d e r s , 22 f a m i l i e s and 51 g e n e r a . food p l a n t . tiv e host. All of t h e s e s p e c i e s have adopted Canada t h i s t l e as a Twenty-six s p e c i e s a l s o use t h i s a l i e n weed as a repro duc ­ However, none of t h e s e i n s e c t s a re r e s t r i c t e d s o l e l y t o Cirsium a r v e n s e , w ith t h e e x c e p ti o n o f Ceutorhynchus l i t u r a ( F a b r i c i u s ) , which has been p u rp o s e ly in tr oduce d f o r t h e c o n t r o l of Canada t h i s t l e . On t h e c o n t r a r y , one t h i r d o f t h e i n s e c t s found f e e d in g on Canada t h i s t l e in so u th er n Montana a r e e i t h e r minor o r major p e s t s of economic crops. Four s p e c i e s , Corythucha d i s t i n c t a Osborn and Drake (Hemiptera: T i n g i d a e ) , B a r is s p . p o s s . c i r s i i G i l b e r t ( C o l e o p t e r a: C u r c u l i o n i d a e ) , Vanessa c ar du i L. ( L e p id o p t e r a : Nymphali d a e ) , and O r e l l i a ru f ic a u d a ( F a b r ic iu s )(Diptera: T e p h r i t i d a e ) , were c onsi de re d c onsp ic uous , because of t h e damage th e y i n f l i c t e d t o t h e t h i s t l e p l a n t . They can be very 19 Table 3 Phytophagous i n s e c t s c o l l e c t e d from Canada t h i s t l e , Cirsium a rv e ns e ( L . ) S c o p . , in so u th er n Montana, 1981. Insects Orthoptera Acndidae Chortophaga v ir ld if a s c ia ta (DeGeer) E ri t e t t i x simplex (Thomas) Melanoplus b i v itta tu s (Say) M. femur-rubrum (DeGeer) M. packardii Scudder M. sanguinipes (Fabricius) Gryllidae Allonemobius allardi (Alexander and Thomas) Hemiptera Miridae Chlamydatus associatus (Uhler) Lygus lineolaris (Palisot de Beauvois) L. robustus Uhler L. schulli Knight Tingidae Corythucha d istin cta Osbom and Drake Pentatomidae Euschistus euschistoides (Vollenmolen) Euschistus sp . Rhytidolomia sp. I unidentified sp. Homoptera Membracidae Ceresini sp. Publilia modesta (Uhler) T ortistilu s wlckhami (Van Duzee) Cercopidae Aphrophora permutata Uhler Philaenus spumarius (L.) Cicadellidae Aceratagallia sp. Agallia sp. Cuerna prob. s t r i a t a (Walker) Empoasca sp. Euscelidius variegatus (Kirschbaum) Macrosteles fascifrons (Stall Xerophloea v i r id ls (FabricIus) I unidentified sp. Frequency in Collection6 Plant Association Plant Growth Stage)s r Stages6 Feeding0 Part(S)6 Recorded Literature Hosts' Source A A N1A N1A A A ECT ECT ECT ECT ECT ECT L L L L L L R R F F F F A ECT L F A A N,A A ECT ECT ECT ECT F F V1F F C F F F P (29) (29) (29) (29) C E.N.A ECT L R1V1F P (13,15) O O O O A N A E ECT ECT ECT - L L L L R F F F P (8.15) O C R N A A ECT ECT ECT S S L F R1V F P - (15) O C A N.A ECT ECT S L.S V R1V1F P P (15) (18) R O O R R LC R R A A N.A A A A N E ECT ECT ECT ECT ECT ECT ECT L L L L L L L S F V R.V R F F F F P (43) P P P (6.44) (6,44) (6,15) LC O LC C C C R R R LC R - P P P P P (15,38) (15) (15.38) (15.38) (15) - P C - 20 Table 3. (con ti n u e d ) Insects Aphididae Aphis fabae Scopoli Aphis SP. Brachycauduscardul (L.) Capitophorus carduinus (Walker) DdCtynotus sp. Pseudococcidae Chnaurococcus t r i f o l l i i (Forbes) Phenacoccus solani Ferris Coleoptera Mordel Iidae Mordellistena sp. Mordellistena sp. "A" Mordellistena sp. "B" Chrysomelidae Criocens duodecimpunctatus ( I.) Deloyala guttata (Olivier) Oiachus auratus (Fabncius) Pachybrachys melanostictus Suffnan Systena blanda Melsheimer Tnrhabda prob. convergens LeConte Curculionidae Baris s p . , poss. c i r s i i Gilbert Ceutorhynchus l itu ra (Fabncius) Dysticheus sp. Macrorhoptus sp. Notaris bimaculatus (Fabricius) Otiorhynchus ovatus (L.) Tychlus p i c i r o s t r is (FabrIcius) Rhinocyllus conicus Froelich Lepidoptera Pterophoridae P la ty p tilia carduidactyla (Riley) , Relative Frequency in ColIectiona Plant Association P T a n l “ Plant Growth Stage(s)e Stages6 Feeding0 P art(s)d Recorded Literature Hosts' Source O O O R R N1A N1A N1A N1A N1A ECT ECT ECT ECT ECT S S S S S F F F F F P t t (46) R R N1A N1A ECT ECT R S F V P P C O O L A A END ECT ECT S L L F V V P - (53) R R O O R R A A A A A A ECT ECT ECT ECT ECT ECT L L L L L L V R R R1V F F P P P P P C (23) (23) (23) (23) (15) (23) C LC R R R O R C L1A L.A A A A A A L.A END1ECT END1ECT ECT ECT ECT ECT ECT END1ECT R1S S1L L L L L L F1L R1V1F R1V F F R V1F R R1V1F t t (17) (48) O L END S V (45) (45) (16) (15,16) P C t (15,23) (23) (5) C (15) 21 Table 3. (c o n ti n u e d ) Insects Tortrlcldae I unidentified sp. I unidentified sp. Arctlidae Apantesiswilliamsii (Dodge) Noctuidae I unidentified sp. Nymphalidae Vanessa (Cynthia) cardul L. Diptera Sciaridae I unidentified sp. Cecidomyiidae I unidentified sp. Tephritidae Orellia ruficauda (Fabricius) Lauxaniidae Camptoproscopella sp. „ Rela tive. Frequency in Col lection® Association Plant Growth Stage(s)e Stages6 Feedlngc Recorded Literature Hosts' Source R R L L ECT ECT L L V V - R L ECT L V C O L END R F - O L ECT L V P O L END R V1F O L END F F _ C L1A END,ECT F F C A ECT L.F V.F - LC aNumber of sites in which the species appears/50 si t e s ; R * Rare (species found in I collection s i t e ) , 0 = Occasional (species found in 2-5 collection s i t e s ) , LC = Locally coimon (species found in 2-5 collection s i t e s , and present in high density), C = Common (species found in more than 5 collection s i t e s ) . bE = eggs, L = larvae, N * nymphs. A = adults. cECT = ectophagous, END = endophagous. bR = roots, S = stems, L = leaves, F = flower heads or buds. eR = rosette stage, V = ver tical growth stage, F * flowering and fru itin g stages. f t - t h i s t l e s (host plants apparently restricted to closely related genera as Carduus, Clrsiun, and Silybun) c = Composltae (host plants apparently restricted to the Compositae), p * poIyphagous (attacking plants belonging to d iffe rent fam ilies ), - * no information. (60) (15,60) (39) 22 e f f e c t i v e in lo c a l a r e a s , although d e n s i t i e s were too low t o a f f e c t t h e weed p o p u l a t i o n s . Grasshoppers were c o l l e c t e d in most of t h e s a m p l e s , with t h e h e a v i e s t i n f e s t a t i o n d u ri n g f lo w e ri n g and f r u i t i n g s t a g e s o f t h e p l a n t . These i n s e c t s a r e g e n e r a l f e e d e r s and t h u s have l i t t l e i n f l u e n c e in t h e n a t u r a l c o n t r o l of t h i s t l e s . pests. In a d d i t i o n , they a r e c o n s i d e r e d ; economic T h i s t l e r o s e t t e s were examined f o r fe e d in g damage in s i t e I (Table.2). The g ra s sh o p p e rs p r e f e r r e d r u s t i n f e s t e d over h e a l t h y t h i s t l e plants. Lewis (1979) s u g g e s ts t h a t such p r e f e r e n c e s a re due t o f a v o r a b l e a l t e r a t i o n s in t h e n u t r i t i o n a l a n d / o r d e f e n s i v e chem is tr y of the plant. The most common Hemiptera c o l l e c t e d from Canada t h i s t l e p l a n t s were Corythucha d i s t i n c t a Osborn and Drake and s e v e r a l Lygus s p e c i e s . The l a t t e r a r e common s a p f e e d e r s on g r a s s e s , weeds and a wide v a r i e t y of economic p l a n t s ( K e l t o n , 1975). C_. d i s t i n c t a appeared in lo c a l popu­ l a t i o n s , p a r t i c u l a r l y in mo is t h a b i t a t s . This lace bug was found f e e d ­ ing on t h e t h i s t l e le av e s from e a r l y May th r oug hout t h e growing season of t h e p l a n t (F ig ure 3 ) . Adults and nymphs were h ig h ly aggregated on t h e t h i s t l e le aves and caused brown o r bla ck fee din g s c a r s (Fig ure 4),. When l a r g e numbers o c cu r re d p e r p l a n t , t h e le av e s became n e c r o t i c . A du lts .w er e c o l l e c t e d in e a r l y May from s i t e s 4, 5 and 7 (Table 2) and co nf in ed t o p o t t e d t h i s t l e p l a n t s in t h e i n s e c t a r y . A f t e r two weeks, most o f t h e p l a n t s showed a g g r e g a ti o n o f nymphs, c l u s t e r e d ne ar t h e s p o t 23 Fi gu re 3. Corythucha d i s t i n c t a a d u l t s f e e d i n g on Canada t h i s t l e l e a v e s . Fig ur e 4. Feeding damage of Corythucha d i s t i n c t a on Canada t h i s t l e . 24 where t h e eggs were l a i d , on t h e u n d e r - s i d e o f t h e l e a f (F ig u re 5 ) . A f t e r a p e r io d of s i x week s, t h e la ce bugs were p r e s e n t in such d e n s i t y t h a t a l l t h e le av e s were c u r l e d and n e c r o t i c . produce fl o w e r buds. These p l a n t s f a i l e d t o £ . d i s t i n c t a i s m u l t i v o l t i n e and has two o r more g e n e r a t i o n s p e r y e a r , depending on t h e c l i m a t i c c o n d i t i o n s (Drake and R u h o ff , 1960). However, n a t u r a l Corythucha p o p u l a t i o n s were too small t o cause s u b s t a n t i a l d e f o l i a t i o n o f t h e t h i s t l e p l a n t s . Lamp and McCarthy (1982) r e p o r t e d t h a t nymphaI p o p u l a t i o n s o f (X d i s t i n c t a on Cirsium canescens N u t t , caused n e c r o s i s o f t h e l e a v e s , but did not reduce t h e seed p ro d u c ti o n o f t h e p l a n t s . Hosto p l a n t s of C_. d i s t i n c t a a r e rec ord ed as being Carduus l a n c e o l a t u s , Cnicus s p . , Cirsium pulc herrimum, £ . c a n e s c e n s , Lathyrus n u t t a l l i i , and Alth a ea s p . , In a d d i t i o n , Ess ig (1958) r e p o r t e d i t s o c c u r re n c e on balsam r o o t , beans,, c o r n , l e t t u c e , l u p i n e , p a r s n i p , squash, and t u r n i p . The s p i t t l e b u g , P h il a e n u s spumarius (L .) was found t o be very abundant in a wide v a r i e t y of h a b i t a t s . The nymphs were observed on t h e t h i s t l e r o s e t t e s in e a r l y May, surrounded by a mass o f s p i t t l e - l i k e froth. The a d u l t s fe d on t h e upper p a r t of t h e t h i s t l e stem th rou ghout t h e r e s t of t h e summer. FX spumarius i s c onsi de re d a n . i m p o r t a n t eco­ nomic p e s t o f f o r a g e crops in e a s t e r n United S t a t e s (Halkka, e t a l . , 1967). plants. However, th e y appeared t o have l i t t l e e f f e c t on t h e t h i s t l e A c o in c i d e n c e o f high r u s t (£. o b t e g e n s ) i n f e s t a t i o n (52%) and a high d e n s i t y o f s p i t t l e b u g nymphs was observed in s i t e I . In June, 25 F ig ure 5. Aggregation of Corythucha d i s t i n c t a nymphs on th e u n d e r - s i d e of Canada t h i s t l e , leaves. 26 1981, a t o t a l of 425 t h i s t l e p l a n t s were examined in s i t e I f o r s p i t t l e bug a t t a c k . An average o f 2 % p l a n t s were found t o be i n f e s t e d with spittlebugs. ' - P u b l i l i a modesta (Uhler) was c o l l e c t e d only as an a d u l t on Cirsium a r v e n s e . Like t h e s p i t t l e b u g s , t h e y fed on t h e upper p a r t of t h e stem from June t o September and were a t t e n d e d by s e v e r a l an t s p e c i e s The aphid c o l o n i e s on Canada t h i s t l e were common in l a t e J u l y and August I n d i v i d u a l c o l o n i e s did not reach damaging d e n s i t i e s . The most numerous phytophagous group in t h e Montana survey was r e p r e s e n t e d by t h e o r d e r o f t h e C o l e o p t e r a . As in t h e European survey ( Z w o l f e r , 1964), t h e l a r g e s t b e e t l e f a m il y c o l l e c t e d in south Montana was t h e C u r c u l i o n i d a e . Four of t h e 15 b e e t l e s p e c i e s fe e d i n g on Canada t h i s t l e a r e endophagous in t h e i r l a r v a l s t a g e s . Of t h e s e , M o r d e l l i s t e n a sp. ( M o r d e l l i d a e ) , B a r i s s p . , po s s. c i r s i i G i l b e r t and Rhynocyllus conicu s F r o e l i c h ( C u r c u li o n id a e ) occu rr ed in a t l e a s t e i g h t , d i f f e r e n t c o l l e c t i o n s i t e s , with a r e l a t i v e high fr equ en cy a t each site. The f i r s t M o r d e l l i s t e n a sp. l a r v a e mining in t h e t h i s t l e stems were observed by m i d - J u l y . These l a r v a e o v e r w in te r in old t h i s t l e s t a l k s , and pupate t h e fo ll o w i n g s p r i n g . The a d u l t s , pro ba bl y emerging in l a t e May, were found f e e d in g on Canada t h i s t l e in Ju ne . The l a r v a l f e e d i n g th ro u g h o u t t h e summer a p p a r e n t l y did not a f f e c t t h e normal growth and r e p r o d u c t i o n of t h e t h i s t l e p l a n t . 27 A ro o t- b o r in g , w e e v i l , B ar is sp. p o s s . c i r s i i G i l b e r t , n a t i v e t o . North America, a t t a c k e d Canada t h i s t l e in s o u t h e a s t e r n Montana. This w e e v i l , a l s o rec ord ed on o t h e r C i r s i u m .spp. ( G i l b e r t , 1964), i s probably t h e only indigenous i n s e c t approaching t h e monophagous h a b i t . Gilbert (1964) c l a s s i f i e s j3. f u t i l i s , j3. c i r s i i , B_. bru nnei pe s and Bv monticola as f o u r s p e c i e s of t h e same subgroup, comprising a. complex o f c l o s e l y r e l a t e d s p e c i e s which a r e a l l r e s t r i c t e d t o h o s t s in t h e genus C ir s iu m . B a r is c i r s i i has been recorded on s i x d i f f e r e n t Cirsium h o s t s in C a l i f o r n i a : Cv q u e r c e to r u m , Cv o c c i d e n t a l e , Cv c o u l t e r ! , Cv c a l i f o r n i c a , C_. cymosum and £ . f o l i o s u m . In our su rv e y, B a r is sp. po s s. c i r s i i was o nly observed on Cirsium a rv e nse and was found e s p e c i a l l y in d i s t u r b e d a r e a s , r o a d s i d e s and g r a s s l a n d s . Both a d u l t and l a r v a l feeding, were observed in s i t e s 5 and 6 and in t h e i n s e c t a r y . ■ B a r is a d u l t s o v e r w i n t e r as unemerged a d u l t s in t h e r o o t s o f t h e i r h o s t s and emerged in e a r l y s p r i n g by making t h e i r way through old l a r v a l g a l l e r i e s t o t h e base of t h e stem. There was no i n d i c a t i o n of a c t i v i t y in t h e c e l l s of o v e r w i n t e r i n g . a d u l t s . t h e t h i s t l e s were about f i v e t o 20 cm t a l l . At t h e time of emergence, The w e e v il s s t a r t e d f e e d ­ ing between t h e newly formed d i s t a l l e a f l e t s of t h e young t h i s t l e s (F ig u re 6 ) ; l a t e r in middle and lower l e a f a x i l s of t a l l e r p l a n t s . G e n e r a l l y , two specimens p e r p l a n t were obser ve d, up t o f o u r on t a l l e r plants. The f e e d i n g on t h e young p l a n t s u s u a l l y damaged t h e primary v e r t i c a l shoot. Damaged p l a n t s produced new s i d e s h o o t s , r e s u l t i n g in Fig ur e 6. B ar is s p . a d u l t fee din g on th is tle rosette. 29 a g e n e r a l bushy a ppea rance , which were d i s t i n c t i v e in t h e f i e l d . Heights of e i g h t damaged t h i s t l e p l a n t s were recorded a t s i t e s 5 and 6 and compared with h e i g h t of e i g h t u n i n f e s t e d p l a n t s a t t h e same s i t e . Adult fe e d i n g reduced t h e v e r t i c a l growth o f t h e t h i s t l e p l a n t s i g n i f i ­ c a n t l y (Table 4 ) . In e a r l y May, B a r i s a d u l t s were t r a n s f e r r e d from s i t e 5 t o t h e i n s e c t a r y . a n d confine d t o p o t t e d t h i s t l e p l a n t s , where t h e i r impact on t h e p l a n t s was ob s er ved . Most o f t h e s e p l a n t s , a lr e a d y under heavy s t r e s s due t o low l i g h t i n t e n s i t y , did not s u r v i v e t h e a t t a c k o f t h e weevil (F ig u re 7) and t h e p l a n t s died w i t h i n two weeks. F i e l d o b s e r v a t i o n s in Montana i n d i c a t e d t h a t emerged a d u l t s l iv e d from two t o f o u r months, were h i g h l y l o c a l i z e d in d i s t r i b u t i o n and d i s p e r s e d by ambulation o r o c c a s i o n a l l y by f l i g h t . G i l b e r t (1964) r e p o r t s t h a t t h e r e i s a p e r i o d o f about t h r e e weeks between emergence and o v i p o s t i o n O v i p o s i t i o n s i t e s , always in t h e lower h a l f o f t h e p l a n t , vary during t h e s e a s on , becoming p r o g r e s s i v e l y lower on t h e main stem. The f i r s t B a r i s l a r v a e were d e t e c t e d in s i t e 5 by mid J u l y . The l a r v a e were f e e d i n g a c t i v e l y in t h e t h i s t l e r o o t s , about 15 cm below ground l e v e l . The c e n t r a l v a s c u l a r c y l i n d e r , as well as t h e c o r t i c a l t i s s u e of t h e r o o t s , were e i t h e r consumed o r used f o r t h e c o n s t r u c t i o n o f t h e pupal chamber (F ig u re 8 ) . By t h e end o f J u l y , t h e small bushy p l a n t s w i l t e d and f a i l e d t o produce fl ow e r buds (F igu re 9 ) . A t o t a l o f 17 w i l t e d p l a n t s were uprooted by t h e end of t h e growing season and unemerged a d u l t s were c ounte d. I found an av erage of Table 4. Site Average p l a n t h e i g h t s (cm) o f e i g h t B ar is i n f e s t e d and e i g h t uni n f e s t e d ' t h i s t l e p l a n t s in s i t e s 5 and 6, June 19, 1981. * P l a n t H eight, cm Weevil P r e s e n t Weevil Absent 5 . 29 . op 6 32.25 58.10 • 71.00 * P l a n t h e i g h t s were s i g n i f i c a n t l y d i f f e r e n t , t - t e s t (P < 0 . 0 1 ) . 31 F ig ure 7. Feeding damage of B aris s p. a d u l t s on p o t t e d t h i s t l e p l a n t s in t h e insectary. Fi g u r e 9. F ig ur e 8 . B ar is sp. l a r v a in Canada t h i s t l e r o o t , Wilted t h i s t l e p l a n t in s i t e 5, i n f e s t e d with B ar is sp. l a r v a e . 32 1.6 unemerged a d u l t s p e r Canada t h i s t l e p l a n t and d is c o v e r e d a s i g n i f i ­ c a n t c o r r e l a t i o n between p l a n t h e i g h t and l a r v a l fr equ en cy ( r = 0 . 7 6 6 , P < 0.01 ) (F ig u re 10). This could be due t o a h i g h e r l a r v a l s u r v i v a l r a t e in t a l l e r p l a n t s o r t o a h i g h e r imaginal freque ncy on t h e l a r g e r p l a n t s e a r l y in t h e seaso n. The damage i n f l i c t e d by t h e weevil i s both d i r e c t and i n d i r e c t , c ov er in g t h e e n t i r e growing season o f t h e p l a n t . Adults s t r e s s t h e p l a n t in e a r l y s p r i n g by d e s t r o y i n g t h e m e r i s t e m a t i c t i p and a f f e c t i n g t h e v e r t i c a l growth o f t h e p l a n t . Larvae burrow i n t o t h e v a s c u l a r t i s s u e of t h e r o o t and block t r a n s p o r t , s t u n t i n g t h e o v e r a l l growth of. t h e p l a n t and p r e v e n t i n g seed p r o d u c t i o n . Both a d u l t and l a r v a l f e e d ­ ing r e n d e r t h e p l a n t s u s c e p t i b l e t o inv asi on by o t h e r org anisms. e a s e s a s s o c i a t e d with weevil damage have not been i d e n t i f i e d . o b s e r v a t i o n s showed t h a t o t h e r i n s e c t s invade damaged r o o t s . i n s e c t s , Chnaurococcus t r i f o l i i D is ­ Field, Two s c a l e (Forbes) and Phenococcus s o l a n i F e r r i s (Homoptera: P s e u d o c o c c i d a e ) , and u n i d e n t i f i e d S c i a r i d a e (D ip te r a ) l a r v a e were c o l l e c t e d from t h i s t l e r o o t s damaged by B ar is l a r v a l f e e d i n g . U n i d e n t i f i e d f a c t o r s a p p a r e n t l y p r e v e n t t h e b u ild up o f high den­ s i t i e s of Baris populations. p a r a s i t e s of t h e genus B a r i s . Very l i t t l e i s known about p r e d a t o r s and None have been observed in t h e s u r v e y ,. with t h e e x c e p ti o n of one C a n t h a r i s sp. l a r v a found a s s o c i a t e d with a B a r i s l a r v a in t h e t h i s t l e r o o t . The r o l e of i n q u i l i n e i n s e c t s i s not y e t known, b u t i t i s a p p a r e n t t h a t t h e s e i n s e c t s could be c o m p e t it o rs o r 33 1 2 3 N o. .Figure 10. 4 OF LARVAE Li near r e g r e s s i o n of p l a n t h e i g h t (cm) v e rs us l a r v a l f r e q u e n c y , August 1981. The b va lu e ( b = l I .183, S. E .= 2.43) is s i g n i f i c a n t l y d i f f e r e n t from zero (P < 0 . 0 1 ) . 34 may i n t e r r u p t o r impede o v i p o s i t i o n . In a l l c o l l e c t i o n s i t e s i n f e s t e d . with B a r i s , M o r d e l l i s t e n a l a r v a e were found fe e d in g in t h e stem p i t h of t h e same p l a n t ; In s i t e 6, R hin oc yll us conicus and B a r i s were p r e s e n t on t h e same p l a n t . G i l b e r t (1964) r e p o r t e d t h a t w h e n .la r v ae of O r e l l i a a r e abundant in C i r s i u m , l a r v a e of B a r is a re s c a r c e o r wa ntin g. This was su pported by my o b s e r v a t i o n s . R hin oc yll us c on ic us F r o e l i c h i s a t h i s t l e seed he ad - fe e din g w e e v i l , in tr od uc e d from France f o r t h e c o n t r o l of Carduus nutans L. (musk t h i s t l e ) . This weevil a l s o a t t a c k s Canada t h i s t l e . Larvae of R:, c on icu s in Canada t h i s t l e seed heads f r e q u e n t l y e a t through, t h e wall of t h e bud, and t h e r e b y become v u l n e r a b l e t o pre d a ci o u s i n s e c t s and s p i d e r s (Rees, 1982). observed in our su rv e y. A maximum of t h r e e l a r v a e p e r fl o w e r bud were Growth c r a c k s in i n f e s t e d Canada t h i s t l e seed- heads and stems were very common. Two s p e c i e s o f Le pid op te ra were found t o d e f o l i a t e t h i s t l e p l a n t s . The most common, Vanessa c ardui L . , t h e p a i n t e d la dy, s t a r t e d fe e d in g on t h e t h i s t l e le ave s in l a t e May, causing c o n s i d e r a b l e d e f o l i a t i o n in local a reas. Because i t i s a m ig r a to r y b u t t e r f l y , i t s numbers f l u c t u a t e widely from y e a r t o y e a r and make i t u n r e l i a b l e as a n a t u r a l c o n tr o l agent. I t can a l s o be a p e s t of sun flowe r and soybean (Morihara B al sb au gh, 1976). and The a r t i c h o k e plume moth, P l a t y p t i l i a c a r d u i d a c t y l a R i l e y , was r e a r e d on f i e l d c o l l e c t e d t h i s t l e stems. I t i s c onsid ered a troubl eso me p e s t o f globe a r t i c o k e in C a l i f o r n i a ( E s s i g , 1958). 35 Most, a d u l t D i p t e r a c o l l e c t e d in t h e Montana survey were c onsid ered o c c a s i o n a l v i s i t o r s , with the. e x c e p ti o n of t h e seed-head f l y , O r e I l i a r u f i c a u d a ( F a b r i c i u s ) , r e p r e s e n t i n g one of t h e most common i n s e c t s in our su rv e y. A dul ts appeared on t h e fl o w e r buds from e a r l y summer th ro u g h o u t J u l y . The l a r v a e fee d on t h e t h i s t l e seeds (F ig u re 11) and weave a coccoon of pappus h a i r s , in which th e y o v e r w i n t e r . By t h e end of t h e summer a t o t a l o f 350 seed-h ead s were c o l l e c t e d a t f o u r d i f f e r e n t sites. Of t h e s e , 170 were d i s s e c t e d in t h e l a b o r a t o r y and 0. r u f icauda pupae were cou nte d. The remaining seed-he ads were s t o r e d t o re c ov er emerging £ . r u f icauda a d u l t s and p o s s i b l e p a r a s i t e s . Our stud y showed an upper l i m i t o f 38% heads a t t a c k e d by £ . r u f icauda with an average of 1.5 l a r v a e per. head (Table 5 ) . This i s 32% l e s s th a n V i r l y and Watson (1977) r e p o r t e d from s t u d i e s a t Macdonald Col le ge (Quebec). A high . oc cu r re n c e of jR. c on ic us in s i t e s A a n d . B, and B ar is s p . in s i t e s . C and .D, a r e pro ba bly reducing o r i n h i b i t i n g t h e 0. r u f icauda i n f e s t a t i o n . An u n i d e n t i f i e d Cecidomyiidae l a r v a c o l l e c t i o n s th ro u g h o u t t h e summer. young f lo w e r buds. f r e q u e n t l y o c cu r re d in my The l a r v a e were.found f e e d in g in t h e All a t t e m p t s t o . r e a r t h i s i n s e c t f a i l e d . This could p o s s i b l y be Dasyneura g ib s o n i F e l t , as r e p o r t e d by Detmers (1927) i n ­ f e s t i n g both s t a m i n a t e and c a r p e l l a t e t h i s t l e heads. 36 Fi g u r e 11. Feeding damage o f O r e l l i a r u f i c a u d a , l a r v a e on Canada t h i s t l e se e ds . 37 Table 5. I n f e s t a t i o n of Canada t h i s t l e heads by O r e I l i a ru f i c a u d a ( F a b r i c i u s ) , September 3, 1981. % . Heads Attacked Mean Number of Pupae/Head Site A (n=60) 38.30 2.04 Site B . (n=37) 33.30 2.16 Site C . ( n=52) . 7.70 1.00 9.50 1.00 S ite D. (n=21) . . n = t o t a l number o f seed heads examined f o r each c o l l e c t i o n s i t e . 38 V i s i t o r s , P r e d a t o r s and P a r a s i t o i ds The v i s i t o r s included in Table 6- a r e p o l l e n c o l l e c t o r s , n e c t a r f e e d e r s , o r i n s e c t s t e n d i n g on aphids and membracids. Most o f t h e predaceous i n s e c t s c o l l e c t e d in t h e Montana survey a r e common p r e d a t o r s p re yi ng on immature o r small i n s e c t s , such as aphids o r t h r i p s . Larvae of Phyllobaenus s p . were observed fe e di ng on 0. r u f i c a u d a l a r v a e in Canada t h i s t l e s e e d -h e a d s . Formica po d z o lica Fr ancoeur was found t e n d i n g aphids on Canada t h i s t l e . This s p e c i e s i s known t o h a r v e s t R. c onicu s l a r v a e on Canada t h i s t l e (Rees, 1982). Of t h e p a r a s i t i c Hymenoptera, l i s t e d in Table 6, two s p e c i e s , Pteromalus s p . and Eurytoma sp. were c o l l e c t e d from pupal chambers of Rv c on icu s in t h i s t l e seed he ads . In e a r l y August, a t o t a l of 615 seed heads were c o l l e c t e d in s i t e 2 and examined in t h e l a b o r a t o r y . c on icu s i n f e s t a t i o n was based on pupal chamber c o u n t s . Rv One hundred twenty-two (20%) s eed -he ads were found t o be i n f e s t e d with Rv c o n i c u s . Of t h e s e , 20% c o n ta i n e d a d u l t P t e r o m a l i d s , i d e n t i f i e d as Pteromalus sp. None of t h e s e were r e a r e d from Rv c onic us l a r v a e and have not been r e ­ corded as p a r a s i t o i d s of Rv conicu s in t h e United S t a t e s (Dowd and Kok, 1982; Rees, 1982). S u r l e s (1974) r e p o r t e d Eurytoma sp. (Eurytomidae) as a l a r v a l p a r a s i t o i d o f Rv conicus in Europe. The p a r a s i t o i d s r e a r e d from O r e l l i a r u f icauda pupae and l a r v a e in t h e i r i s e c t a r y were i d e n t i f i e d as Eulophidae (Hymenoptera). At t h e end o f t h e summer, Canada t h i s t l e , s e e d - h e a d s , c o l l e c t e d a t f o u r d i f f e r e n t 39 Table 6 V i s i t o r s , p r e d a t o r s , and p a r a s i t o i d s c o l l e c t e d from Canada t h i s t l e , Cirsium a rv e nse (L .) Scop, in so ut her n Montana, 1981. Relative Frequency in Collection^ Stages Collectedb Plant P a rt(s)c Plant Growth Stage(s )d Hemiptera Anthocoridae Orius t r is tic o lo r (White) O N1A F F Nabidae Nabis altern ates Parshley LC N1A L R.V.F Insects predaceous on mites aphids, th rip s , other small insects and eggs R A L F Lygaeidae Geocoris sp. R N1A L V1F O A L F O A L A L Coleoptera E lateridae Ctenicera gIauca (Germar) Cantharidae Cantharis sp. Drob. Canthans sp. Anobiidae Tricorvnus productus White Dermestidae Attagenus canadensis Casey Cleridae Phyllobaenus or lsohydnocera sp. Phyllobaenus sp. Trichodes ornatus Say R Source (1,30) predaceous on aphids,(30) th rip s and other small insects Reduviidae Sinea diadema (Fabncius) Neuroptera Hemerobiidae Micromus vanolosus Hagen Chrysopidae Chrysopa carnea Stephens Habits predaceous on so ft bodied insects (8,30) F pollen and nectar feeder (7) R flower v is ito r (15) predaceous on Aphis (15,23) O R A L L R R F R A L R R A L R flower v is ito r (23) O L F F Feeding on 0. ru fIcauda leaves predaceous on woodboring insects Pers. observ R R A A L F V F - flower v isito rs larvae predaceous on bees and wasps (23) (15.23) 40 Table 6. (c o n ti n u e d ) Insects Melyndae Collops bipunctatus Say Collops tric o lo r (Say) Malachius aeneus L. Coccinel Iidae Brachyacantha ursina (Fabricius) Coccinella transversoguttata transversoguttata Falderman Hippodamia convergens Guerin H. parenthesis (Say) Keiative Frequency in Collection* Stages Collectedb Plant P a rt(s)c O R . R A A A L L L V.F V.f V R A L V A E.L.A L L F F A L F O LC R H. quinquesignata qumqueslgnata (Kirby) LC Hyperaspis undulata (Say) Scymnus (Pullus) postpinctus Casey Lepidoptera PyralIdae I unidentified sp. Blastobasidae I unidentified sp. Diptera Chironomidae Cricotopus sp. Sciaridae Bradysia sp. Bombyllidae Systoechus vulgaris Loew Syrphidae Sphaerophoria p hilanthus (Meigen) S y rltta piplens (L .) Anthomyiidae Delia platura (Meigen) Plant Growth Stage(s)d E.L.A L R.V.F R A L F R A L R LC A L V R A L V R A L R R A F F LC A F F LC O A A F F F F O A F F Habits Source flower v isito rs (15.23) - . predaceous on Aphis s p ., eggs and larvae predaceous on Aphis sp. predaceous on Aphis sp predaceous on various unarmored scales (15) (15) (15) (15) . predaceous on grasshopper egg pods (9) - larva is a pest of vegetables (9) 41 Table 6. (con ti n u e d ) Insects Keiative Frequency in Collectiona Stages Collectedb Plant P a r tts lc Scathophagidae Scathophagastercorarla (L.) O A L Tachinidae Hyalomya a ld rich it Townsend P eleteria sp. R R A A R Plant Growth Stage(s)d Habits Source V predaceous on blow­ fly and house fly F F p arasite of Hemiptera (9) F A F F - R* A F F - R A L V I unidentified sp. I unidentified sp. Eulophidae I unidentified sp. R R A A L F V F O L F F I unidentified sp. C L F F L.A F F - A F F - Hymenoptera Xyelidae Xyela obscura (Strobl) Braconidae Bracon sp. Ichneumonidae Thyrateles sp .. poss. Iugubrator (Gravenhorst) Pteromalidae Pteromalus sp. EuryLomidae Eurytoma sp. Chrysidldae Hedychrum sp iloventer French Formicidae Formica neoclara Emery F. o b scunventris c l Ivia Creighton F. podzolica Francoeur Formica sp. Hyrmica Incompleta Provancher Pompilidae PomplIus sp. Apidae Apis m ellifera L. Bombus sp. LC O F (15) - p arasite of several Nymphalldae (31) - p arasite of 0. ruficauda larvae p arasite of 0. ruficauda larvae (pers. observ) (pers. observ) R A F F O LC R R S.L S1L S.L S.L S.L R.V.F R.V V.F O A A A A A R A F F predaceous on spiders (31) LC A F F O A F F feeding on nectar and (31) pollen feeding on nectar and (31) pollen F V p arasite of Cecerine wasp (Bohart, R.M. pers. comm.) Tending Tending Tending Tending (66) aphids aphids aphids Membracids (6 6 ) (pers. observ) (pers. observ) - aNumber of s ite s in which the species appears/50 s ite s ; R « Rare (species found in I co llectio n s i t e ) , O = Occasional (species found in ?-5 collection s ite s ) . LC = Locally common (species found in 2-5 collection s ite s , and present in high d en sity ). C = Common (species found in more than 5 collection s ite s ) . bE = eggs, L = larvae. N = nymphs. A = adults. cR = roots, S = stems. L = leaves, F = flower heads or flower buds. dR = ro sette stage. V = v ertical growth stage, F * flowering and fru itin g stages. 42 s i t e s were examined f o r 0. r u f i c a u d a i n f e s t a t i o n (Table 5 ) . At t h e same time p a r a s i t i z e d 0. r u f icauda pupae were counted (Table 7, Figure 12). An average in c id e n c e o f 49% p a r a s i t i s m was obser ved . Summary Whenever Canada t h i s t l e i s f o u n d , i t has a l a r g e number of i n s e c t s a s s o c i a t e d with i t . Although,many o f t h o s e i n s e c t s a r e s t r a y s from o t h e r p l a n t s , many o f them can be c o n s id e re d as i n c i d e n t a l v i s i t o r s and about one t h i r d of t h e l i s t e d phytophagous s p e c i e s a l s o a t t a c k economic plants. Only a few i n s e c t s were c o n s id e re d conspicuous because of t h e i r damage i n f l i c t e d t o t h e t h i s t l e p l a n t and because o f t h e i r commonness to the c o lle c tio n s . The i n s e c t survey in so uth ern Montana i n d i c a t e s t h a t Cirsium a rv e ns e has been e x p l o i t e d by t h r e e seed he ad - fe e d in g i n s e c t s , t h r e e s tem -bo rin g i n s e c t s , one r o o t - b o r i n g i n s e c t , and two d e f o l i a t i n g a g e n t s . C on sid erin g t h e i r in c id e n c e of a t t a c k , more i n s e c t s were a s s o c i a t e d with t h e . d e v e l o p i n g seed heads than with f o l i a g e , stems o r f o o t s . Although some i n s e c t s caused c o n s i d e r a b l e p l a n t s t r e s s , f u r t h e r s t r e s s f a c t o r s from o t h e r i n s e c t s and pathogens a re needed t o c o n t r o l ■ t h i s weed. Few p o t e n t i a l l y e f f e c t i v e and h o s t s p e c i f i c i n s e c t s from Zwolfer's l i s t . a r e s t i l l a v ailab le f o r fu tu re introd u ctio n s. Thus, t h e in fo r m a t io n g a th e r e d in t h i s survey could form t h e f o u n d a t i o n f o r f ol lo w - up s t u d i e s on endemic s p e c i e s . Stenophagous i n s e c t s , as 43 Table 7. P a r a s i t i s m of O r e l I l a r u f i c a u d a ( F a b r i c i u s ) pupae in Canada t h i s t l e seed he ad s , September 3, 1981. Tot al # Heads Attacked Site A (n=60). Site B (n=37) ■ 23 ■ To ta l # Pupae Tot al # P a r a s i t i z e d Pupae 47 21 26 7 . . 12 - S i t e C(n=52) 4 4 3 S i t e D. (n=21) 2 2 I h = To ta l number of seed heads examined f o r each c o l l e c t i o n s i t e . 44 45 Ceuthorynchus I i t u r a and B a r is s p . , which a r e e s t a b l i s h e d , b u t a re sp re ad in g s lo w ly , could be augmented and r e d i s t r i b u t e d over t h e s t a t e . The p o t e n t i a l o f t h e combination o f two s t r e s s - f a c t o r s , t o i n ­ c r e a s e t h e impact on t h e t h i s t l e p l a n t s in t h e f i e l d , should be i n v e s t ! gated. I n s e c t damage and a pathogen form a p e r f e c t comb ina tion. In t h e i r stu dy of i n s e c t involvement in pathogen t r a n s m i s s i o n , H a r r i s and Maramorosch (1980) s t a t e d : "We o f t e n ove rlo ok t h e f a c t t h a t i n s e c t i n ­ vad ers of p l a n t s a r e a l w a y s , w it h o u t any e x c e p t i o n , accompanied or f o l ­ lowed by fun gi and b a c t e r i a . " Endemic pathogens of Canada t h i s t l e a r e p r e s e n t ; two fung al d i s e a s e s , P u c c in ia obte gen s and S c l e r o t i n i a . s c l e r o t i o r u m and an a s t e r yellow caused by a mycoplasma-1 ike organism. The w e ev ils Ceuthorynchus l i t u r a , Rh in oc yllu s c onic us and B ar is s p . a re p o t e n t i a l v e c t o r s of b o t h , v i r a l and fung al d i s e a s e s . Transmission, exp erimen ts a r e n e c e s s a r y t o e v a l u a t e t h e i r p o t e n t i a l as t h i s t l e p a th o ­ gen v e c t o r s . LITERATURE CITED 1. Anderson, N. H. 1962. Anthocorldae of t h e P a c i f i c Northwest with n o te s on d i s t r i b u t i o n s , l i f e h i s t o r i e s , and h a b i t s ( H e t e r o p t e r a ) . Can. E n t . 94:1325-1334. 2. Andres, L. A. 1980. The b i o l o g i c a l . c o n t r o l o f Canada t h i s t l e ( Cirsium a r v e nse ( L . ) Scop, in t h e United S t a t e s . Proceedings o f Can. t h . Symposium. March 1980. A g r i c u l t u r e C anada:112-127. 3. Andres, L. A ., C. J . D a v i s , P. H a r r i s , and A. J . Wapshere. 1976. ■ B i o l o g i c a l c o n t r o l o f weeds. In "Theory and P r a c t i c e of B i o l o g i ­ c a l c o n t r o l (Huff a k e r , C. B. and P. S. Messenger, e d . ) . Academic P r e s s , New Y o r k . . 481-497. 4. Andres, L. A., and R. D. Goeden. 1971. The b i o l o g i c a l c o n t r o l of weeds by in tr o d u c e d n a t u r a l enemies, in " B i o l o g i c a l Control" (C. B. H u f f a k e r , e d . ) . Plenum, NY. 143-164. 5. B a t r a , S. W. T . , J . R. Coulson, P. 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