Observations on the Interrelationships among Ants Aphids and Ap

The Great Lakes Entomologist Volume Number October 1967 Number October 1967 Article October 2067 Observations on the Interrelationships among Ants, Aphids, and Aphid Predators Beverly Rathcke Gustavus Adolphus College Charles L Hamrum Gustavus Adolphus College Arthur W Glass Gustavus Adolphus College Follow this and additional works at: https://scholar.valpo.edu/tgle Part of the Entomology Commons Recommended Citation Rathcke, Beverly; Hamrum, Charles L.; and Glass, Arthur W 2067 "Observations on the Interrelationships among Ants, Aphids, and Aphid Predators," The Great Lakes Entomologist, vol (5) Available at: https://scholar.valpo.edu/tgle/vol1/iss5/5 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar For more information, please contact a ValpoScholar staff member at scholar@valpo.edu Rathcke et al.: Observations on the Interrelationships among Ants, Aphids, and Ap 1967 THE MICHIGAN ENTOMOLOGIST 169 OBSERVATIONS ON THE INTERRELATIONSHIFS AMONG ANTS, APHIDS AND APHID PREDATORS B e v e r l y R a t h c k e , C h a r l e s L H a m r u m and A r t h u r W G l a s s D e p a r t m e n t of B i o l o g y , G u s t a v u s Adolphus C o l l e g e St P e t e r , M i n n e s o t a 56082 T h e l i t e r a t u r e c o n c e r n i n g t h e i n t e r a c t i o n s of t h e s e i n s e c t g r o u p s is a s i n t e r e s t i n g a s it is e x t e n s i v e I t h a s g e n e r a l l y b e e n a c c e p t e d t h a t t h e r e l a t i o n s h i p b e t w e e n a n t s and a p h i d s h a s b e e n o n e of m u t u a l i s m i n which a n t s d e r i v e a l l o r a l a r g e p a r t of t h e i r n u t r i e n t s f r o m aphid honeydew and p e r f o r m s p e c i a l f u n c t i o n s i n r e t u r n , p r i m a r i l y t h a t of p r o t e c t i n g a p h i d s f r o m t h e i r e n e m i e s E v e r y a s p e c t of t h e s e a s s o c i a t i o n s h a s b e e n c o v e r e d by t h e e x c e l l e n t r e v i e w s of Nixon (1951) and Way (1963) I t now a p p e a r s t h a t t h e n a t u r e and e f f e c t i v e n e s s of t h e p r o t e c t i v e value a f f o r d e d t h e a t t e n d e d a p h i d s by a n t s c a n n o t b e r e d u c e d to a s i m p l e s t a t e m e n t of s y m b i o s i s Nixon (1951) pointed o u t n u m e r o u s s i t u a t i o n s t h a t a r e r e s p o n s i b l e f o r f l u c t u a t i o n s i n t h e ant-aphid r e l a tionship The e m p h a s i s in the p r e s e n t study was on the protection e x t e n d e d t o Aphis m m i c i s Linn., a s m a l l aphid found on s p i r e a s h o o t s by t h e c o m m o n tending a n t s T h i s r e s t r i c t i o n allowed c o n s t a n t s u r v e i l l a n c e of t h e p r o t e c t i o n s u p p l i e d t o a s i n g l e aphid s p e c i e s by a n t s a g a i n s t t h e s e a s o n a l s u c c e s s i o n of p r e d a t o r s Aphid p a r a s i t e s w e r e i g n o r e d in t h i s study T h e s e o b s e r v a t i o n s w e r e m a d e f r o m J u n e t o mid-August, 1966, on f o u r d i s p e r s e d c l u m p s of aphid-infested s p i r e a b u s h e s Only two a n t s , Lasius neonigev E m e r y and Fovmica fusca L i n n a e u s w e r e i n r e g u l a r a t t e n d a n c e t o Aphis vumicis T h e r e s p o n s e s of t h e s e a n t s to the aphid and t o t h e f i v e p r e d a t o r g r o u p s w e r e s o d i f f e r e n t t h a t e a c h a n t w a s treated separately The predator s p e c i e s were not identified in all i n s t a n c e s S p e c i e s of C h r y s o p i d a e , C o c c i n e l l i d a e , S y r p h i d a e , C h a m a e m y i i d a e , and C e c i d o m y i i d a e a t t a c k e d t h e t e n d e d aphids The p r e d a t o r p r o t e c t i o n p r o v i d e d by t h e a n t s w a s t h e p r i m e o b j e c t of t h e study T h e r e f o r e , o p p o r t u n i t i e s to identify p r e d a c e o u s l a r v a e w e r e f r e q u e n t l y s a c r i f i c e d f o r p o t e n t i a l o b s e r v a t i o n of a n t - p r e d a t o r e n c o u n t e r s Lasius neonigev E v e n though m a n y s y r p h i d l a r v a e p r e y e d upon t e n d e d a p h i d s , i n only o n e c a s e w a s a s y r p h i d l a r v a a t t a c k e d by Lasius neonigev, and i n t h i s i n s t a n c e t h e a t t a c k m a y h a v e b e e n p r e c i p i t a t e d by t h e a c t i o n s of t h e o b s e r v e r a s t h e l a r v a w a s p l a c e d on a s p i r e a twig T h e i n t e r f e r e n c e by L neonigev i n t h e aphid f e e d i n g a c t i v i t i e s of Aphidoletes ( C e c i d o m y i i d a e ) and Leucopis ( C h a m a e m y i i d a e ) w a s e v e n l e s s t h a n i n t h e c a s e of s y r p h i d s On o n e s p i r e a b u s h t h e s e p r e d a t o r s c o m p l e t e l y k i l l e d t h e aphid population b e i n g tended Ants often o v e r r a n t h e l a r v a e and Published by ValpoScholar, 2067 The Great Lakes Entomologist, Vol 1, No [2067], Art 170 THE MICHIGAN ENTOMOLOGIST Vol 1, No tapped t h e m with t h e i r antennae Only o n e Leucopis l a r v a w a s d i s t u r b e d by t h e s e a t t e n t i o n s and l e f t t h e b r a n c h while t h e o t h e r s r e m a i n e d undisturbed Leucopis pupae w e r e c o m m o n l y found among tended aphids T h e s e a n t s w e r e n e v e r o b s e r v e d t o d i s t u r b any d i p t e r a n e g g s T h e r e s p o n s e s of Lasius neoniger t o c o c c i n e l l i d l a r v a e s e e m e d t o depend upon t h e s i z e of t h e l a r v a e S m a l l l a r v a e , to m m long, w e r e not a t t a c k e d T h e a n t s would, h o w e v e r , i n v e s t i g a t e t h e i r b o d i e s with antennae and p a l p s T h e s e s m a l l l a r v a e would r e m a i n p e r f e c t l y quiet d u r i n g t h e s e a t t e n t i o n s , but would continue feeding o r s e a r c h i n g a s soon a s t h e a n t s would l e a v e t h e m T h e l a r g e r c o c c i n e l l i d l a r v a e w e r e always attacked One l a r v a , about m m l o n g , was d r a g g e d down a b r a n c h by t h r e e a n t s H o w e v e r , t h e s m a l l Lasius w o r k e r s w e r e u s u a l l y u n a b l e to i n f l i c t m u c h d a m a g e o n the l a r g e l a r v a e In a l l o b s e r v e d i n s t a n c e s , the tending a n t s did s u c c e e d in d r i v i n g l a r g e c o c c i n e l l i d l a r v a e away f r o m aphids C h r y s o p i d ( N e u r o p t e r a ) l a r v a e always e x c i t e d L neonigev w h e n e v e r they w e r e e n c o u n t e r e d n e a r aphids S o m e a n t s s u c c e e d e d i n biting a s m a l l c h r y s o p i d l a r v a e and c a r r i e d t h e s t r u g g l i n g l a r v a away to t h e n e s t Many a n t s w e r e r e q u i r e d t o i n t e r f e r e s u c c e s s f u l l y with t h e s e a r c h i n g and feeding a c t i v i t i e s of t h e l a r g e r l a c e w i n g l a r v a e Whenever t h e tending a n t s would a t t a c k , t h e l a r v a would r a i s e t h e t i p of t h e abdomen and exude a d r o p l e t of b r o w n liquid The l a r v a would t h e n t w i s t about t r y i n g to touch an a n t with t h i s d r o p l e t Any a n t touched by t h e d r o p l e t would f l e e f r o m t h e s c e n e Soon all t h e n e a r b y a n t s would g a t h e r and t r y to a t t a c k t h e l a r v a Many w e r e r e p e l l e d b e f o r e t h e l a r v a w a s finally d r i v e n away Whenever L neoniger o c c u r r e d i n sufficient n u m b e r s , they s e e m e d to b e quite c a p a b l e of p r o t e c t i n g t h e i r tended aphids a g a i n s t t h e l a r g e r c h r y s o p i d and c o c c i n e l l i d l a r v a e H o w e v e r , t h e y did not s e e m to r e c o g n i z e the D i p t e r a l a r v a e a s i n t r u d e r s T h e s e l a r v a e w e r e g e n e r a l l y quite p a s s i v e and o r d i n a r i l y did not m a k e e x c i t e d m o v e m e n t s when touched by t h e a n t s T h e s e a n t s a p p e a r e d t o t r e a t t h e m a s they would t r e a t an aphid, tapping t h e m with t h e i r antennae and often p a l p a t i n g t h e m T h e s e p r e d a t o r s did not s e e m to b e f o r e i g n to t h i s Lasius s p e c i e s Pontin ( 9 ) l i s t s two s y r p h i d s p e c i e s and C h a m a e m y i i d a e and C e c i d o m y i i d a e l a r v a e a m o n g t h e p r e d a t o r s adapted t o a t t a c k a p h i d s tended by ants Our observations support Pontin's claim that certain predators a f e t o l e r a t e d by Lasius and Formica s p e c i e s Lasius neoniger w a s o b s e r v e d tending populations of a waxy c o c c i d , Planococcus citri R i s s o , on c h r y s a n t h e m u m s S o m e s p i r e a aphids w e r e t r a n s f e r r e d to t h e ' m u m ' p l a n t s A f t e r having s e t t l e d on t h e p l a n t , they w e r e quickly s e i z e d and c a r r i e d off to t h e n e s t by the ants Within a few m i n u t e s a l l of t h e < ' i m m i g r a n t " aphids w e r e r e m o v e d C o n t r a d i c t o r y r e s u l t s w e r e obtained when c o c c i d s w e r e added to an aphid population on s p i r e a In t h i s i n s t a n c e t h e " i m m i g r a n t s " w e r e a c c e p t e d by t h e a n t s which r e g u l a r l y s o l i c i t e d honeydew f r o m t h e m T h e c o c c i d s r e m a i n e d on t h e s p i r e a f o r s e v e r a l d a y s N o explanation i s e v i d e n t f o r t h e d i f f e r e n t r e s p o n s e s by L neonigev t o w a r d t h e s e two h o m o p t e r a n s p e c i e s T h e r e s p o n s e of L neo.nig2.v to t h e p r e s e n c e of p r e d a t o r s a m o n g https://scholar.valpo.edu/tgle/vol1/iss5/5 Rathcke et al.: Observations on the Interrelationships among Ants, Aphids, and Ap 1967 THE MICHIGAN ENTOMOLOGIST 171 tended aphids was not c o n s i s t e n t with the o b s e r v a t i o n s made on o t h e r Lasius s p e c i e s Way (1963) did not c o n s i d e r Lasius s p e c i e s to be e s p e c i a l l y p r e d a c e o u s o r h o s t i l e However, El-Ziady and Kennedy (1956) showed t h a t Lasius n i g e ~ c a r r i e doff o r d r o v e away l a r v a e of Syrphidae and of the coccinellid Adalia bipunctata Linn but did not h a r m syrphid o r coccinellid eggs Adult c o c c i n e l l i d s were a l s o attacked by L niger (Linn) B a n k s (1962) c o n f i r m e d t h a t L niger attacked s y r p h i d and coccinellid l a r v a e and d r o v e away c h r y s o p i d l a r v a e a s well a s adult and i m m a t u r e Anthocoridae ( H e m i p t e r a ) B a n k s (1962) a l s o o b s e r v e d the r e m o v a l of e g g s of s y r p h i d s and c o c c i n e l l i d s by this s p e c i e s Conflicting with t h e s e o b s e r v a t i o n s a r e the investigations of H e r z i g (1938) and Pontin (1959) H e r z i g concluded t h a t ants r e a c t to swift moving bodies a s a t h r e a t to t h e m s e l v e s and not fight i n t r u d e r s to p r o t e c t t h e i r aphids His e x p e r i m e n t s included o b s e r v a t i o n s of t h r e e Lasius s p e c i e s Pontin found aphid p r e d a t o r s i n the n e s t s of L niger and L ~ Z U V U S(Fab.) F o m i c a fusca During the e a r l y s u m m e r , s y r p h i d , Leucopis, c h r y s o p i d , and coccinellid eggs w e r e found on an aphid-infested s p i r e a b u s h tended by a l a r g e colony of Formica f u ~ c a H o w e v e r , during this p e r i o d p r e d a t o r s w e r e r a r e l y s e e n When l a r v a e w e r e t r a n s f e r r e d to the bush they were attacked a s soon a s d i s c o v e r e d T h e s e l a r g e r a n t s swiftly b i t and c a r r i e d to the n e s t a l l introduced s y r p h i d , Leucopis, and Aphidoletes l a r v a e All chrysopid and l a r g e coccinellid l a r v a e w e r e a l s o promptly attacked The a n t s w e r e often r e p e l l e d by l a r g e r c h r y s o p i d s i n the s a m e m s n n e r t h a t they discouraged L neoniger One Formica was r e p u l s e d by a s e c r e t i o n f r o m a wounded coccinellid l a r v a A s i m i l a r r e s p o n s e was noted when a F o m i c a which b e c a m e agitated r e t r e a t e d f r o m a wounded Leucopis The coccinellid l a r v a e usually e s c a p e d t h e attacking ants by dropping to the ground In one i n s t a n c e s e v e r a l s m a l l coccinellid l a r v a e w e r e o b s e r v e d among the tended aphids and w e r e not attacked In t h i s i n s t a n c e F fusca behaved a s L neoniger Aside f r o m the s m a l l c o c c i n e l l i d s , F f u ~ c ahad no difficulty r e c o g nizing and dealing with aphid p r e d a t o r s During t h i s p e r i o d the aphid population was quite stable The e f f e c t i v e n e s s of t h e i r protection bec a m e a p p a r e n t when they abruptly c e a s e d tending the aphids n e a r the end of J u l y , s e v e r a l days b e f o r e t h e winged ants a p p e a r e d i n August T h i s s p i r e a bush was r a p i d l y colonized by the D i p t e r a p r e d a t o r s , a coccinellid and a debris-collecting chrysopid T h e r e a f t e r , the aphid population was s h o r t l y decimated In the only o b s e r v e d i n s t a n c e of egg p r e d a t i o n , a F fusca was s e e n devouring a c h r y s o p i d egg f r o m a b e n t egg stalk The a n t s were a l s o of benefit to the aphids by keeping t h e m clean In untended a r e a s the aphids on s t e m t i p s b e c a m e sticky and clogged with c a s t s k i n s and d e b r i s Published by ValpoScholar, 2067 The Great Lakes Entomologist, Vol 1, No [2067], Art 172 THE MICHIGAN ENTOMOLOGIST Vol 1, NO T h e l i t e r a t u r e d o e s not i n d i c a t e t h a t E u r o p e a n F o v m i c a a r e f o r m i d able aphid p r o t e c t o r s Nixon (1951) d o e s not p r o v i d e e v i d e n c e of a g g r e s s i v e p r o t e c t i o n by F o v m i c a Pontin (1959) c i t e s t o l e r a n c e of C o c c i n e l l a divaricata 01 by F o v m i c a s a n g u i n e a L a t r F o v m i c a yufa Linn i s r e g a r d e d by Way (1963) a s only a p a r t i a l l y s u c c e s s f u l s y r p h i d p r e d a t o r Both ant s p e c i e s o b s e r v e d in t h i s study a p p e a r e d t o e x h i b i t s o m e f o r m of o w n e r s h i p t o w a r d t h e attended aphids A L a s i u s neonigev, capt u r e d o n a b u s h and t r a n s f e r r e d to a n o t h e r b u s h f e e t away, was killed by the n a t i v e L n e o n i g e v population A L a s i u s i n t r o d u c e d onto t h e F o v m i c a - t e n d e d b u s h was a l s o k i l l e d with d i s p a t c h C o l l a t e r a l e v i d e n c e of aphid "ownership" m a y b e i n f e r r e d f r o m the r e a c t i o n s of t h e s e a n t s to aphid p r e d a t o r s i n t h e a b s e n c e of aphids A l a r g e c o c c i n e l l i d l a r v a was p l a c e d on t h e ground n e a r the F o v m i c a f u s c a n e s t I t was c o m p l e t e l y i g n o r e d by t h e a n t s a s i t c r a w l e d on t h e ground When t h i s l a r v a w a s p l a c e d on a t e n d e d b r a n c h i t w a s p r o m p t l y a t t a c k e d and f o r c e d t o d r o p t o t h e a p p a r e n t s a f e t y of t h e ground C h r y s o p i d l a r v a e o n t h e g r o u n d r e s p o n d e d d e f e n s i v e l y t o w a r d L a s i u ~which i n t u r n i g n o r e d the l a r v a e T h e a g g r e s s i v e b e h a v i o r s e e m s t o b e e x h i b i t e d only u n d e r conditions i n which o w n e r s h i p of t e r r i t o r y e x i s t s Undoubtedly s o m e populations of Aphis v u m i c i s p r o s p e r e d through the a t t e n t i o n s of F fusca and L neonigev F o v m i c a f u s c a was v e r y e f f e c t i v e in p r o t e c t i n g t h e aphids and k e p t a s t a b l e population of aphids until August L a s i u s n e o n i g e v a p p e a r e d t o b e m o r e e r r a t i c i n tending and would d e s e r t t h e aphids f o r days They w e r e not n e a r l y s o effective a s p r o t e c t o r s a g a i n s t p r e d a t o r s T h e n a t u r e of t h i s p r o t e c t i o n m a y o r m a y not b e a d e f e n s e of "owned" aphids T h e v a r i a b i l i t y of t h e d a t a p r e s e n t e d by r e s p o n s i b l e o b s e r v e r s c e r t a i n l y i n d i c a t e s t h a t t h e s e i n t e r r e l a t i o n s h i p s a m o n g a n t s , a p h i d s , and aphid p r e d a t o r s fluctuate M o r e s t u d y i s needed to define t h e r e a s o n s f o r t h e s e fluctuations P e r h a p s the a n t s p e c i e s involved i s chiefly r e s p o n s i b l e f o r t h e v a r i e t y of e f f e c t s observed in these interspecific relationships ACKNOWLEDGMENTS T h e a u t h o r s a r e indebted to Edmund O l s o n , D e p a r t m e n t of Entomology, U n i v e r s i t y of M i n n e s o t a , St P a u l , M i n n e s o t a , f o r identifying the h o m o p t e r a n s p e c i e s utilized i n t h i s study We a l s o thank Violet Neal f o r h e r aid i n t h e final p r e p a r a t i o n of the m a n u s c r i p t T h i s s t u d y was s u p p o r t e d by t h e G u s t a v u s R e s e a r c h Fund LITERATURE CITED B a n k s , C J 1962 E f f e c t s of the a n t Lasius n i g e v ( L ) on i n s e c t s p r e y i n g on s m a l l populations of Aphis f a b a e Scop on b e a n plants Ann Appl Biol 50:669-679 E l - Z i a d y , S and J.S Kennedy 1956 B e n e f i c i a l e f f e c t s of t h e c o m m o n g a r d e n a n t , L a s i u s n i g e r L , on t h e b l a c k b e a n a p h i d , Aphis f a b a e Scopoli Roy Entomol Soc London P r o c (A) 31:61-65 H e r z i g , J 1938 A m e i s e n und Blattl'duse Z Angew Entomol 24:367435 https://scholar.valpo.edu/tgle/vol1/iss5/5 Rathcke et al.: Observations on the Interrelationships among Ants, Aphids, and Ap 1967 Nixon, G.E.J London THE MICHIGAN ENTOMOLOGIST 173 1951 T h e a s s o c i a t i o n of a n t s with a p h i d s and c o c c i d s P o n t i n , A.J 1959 S o m e r e c o r d s of p r e d a t o r s and p a r a s i t e s adapted to a t t a c k a p h i d s a t t e n d e d by a n t s E n t o m o l Monthly Mag 95: 154-155 Way, M.J 1963 M u t u a l i s m b e t w e e n a n t s and honeydew-producing H o m o p t e r a Ann Rev E n t o m o l 8:307-344 REVIEWS OF RECENT LITERATURE TRAP-NESTING WASPS AND B E E S : L I F E HISTORIES, NESTS AND ASSOCIATES K a r l V K r o m b e i n W a s h i n g t o n , D.C.: T h e S m i t h s o n i a n I n s t i t u t i o n , 1957 S m i t h s o n i a n Publ 4670 v i , 570 pp $12.50 T h e t e c h n i q u e of "trap-nesting" f o r w a s p s and b e e s by putting o u t s t r i p s of wood h a v i n g a h o l e b o r e d i n o n e e n d i s n o t e x a c t l y n e w , b u t only within t h e l a s t y e a r s h a s i t b e e n widely e m p l o y e d i n t h i s count r y T h i s new book by K a r l V K r o m b e i n , c h a i r m a n of t h e D e p a r t m e n t of Entomology a t t h e U.S National M u s e u m , r e v e a l s how e n o r m o u s l y p r o d u c t i v e t h e t e c h n i q u e c a n be I t i s , of c o u r s e , u s e f u l only f o r s p e c i e s t h a t n o r m a l l y n e s t i n hollow t w i g s S p e c i e s t h a t b o r e i n pith ( s u c h a s m a n y c r a b r o n i n e w a s p s ) d o not u s u a l l y a c c e p t t h e t r a p s , n o r groundn e s t e r s (which m a k e u p t h e m a j o r i t y of w a s p s and b e e s ) D e s p i t e i t s l i m i t a t i o n s , t h e t r a p - n e s t i n g technique i s e x t r e m e l y v a l u a b l e , a s i t a l l o w s o n e to follow t h e d e v e l o p m e n t of t h e c e l l cont e n t s c l o s e l y o n c e t h e n e s t h a s b e e n s p l i t o p e n , and a l s o b e c a u s e t h e n e s t s a r e s o s i m p l e and i n e x p e n s i v e t h a t i t i s p r a c t i c a b l e to u s e l a r g e n u m b e r s of t h e m , t h u s obtaining i n f o r m a t i o n on ecology and population p h e n o m e n a not a v a i l a b l e f r o m i s o l a t e d n e s t s T h e technique is a l s o u s e f u l i n s u m m e r p r o g r a m s f o r young p e o p l e ; t h e r e i s p r o b a b l y no equally s i m p l e m e t h o d f o r d e m o n s t r a t i n g s o m e of t h e complexities in natural environments K r o m b e i n ' s s t u d y is b a s e d on d a t a g a i n e d f r o m about 3400 t r a p - n e s t s put o u t o v e r a - y e a r p e r i o d i n s e v e r a l e a s t e r n s t a t e s and A r i z o n a I n a l l , h e t r e a t s 75 s p e c i e s of w a s p s t h a t a c c e p t e d h i s t r a p s , a s well a s 43 s p e c i e s of b e e s and 83 s p e c i e s of p a r a s i t e s and p r e d a t o r s of d i v e r s e g r o u p s P r e v i o u s knowledge of t h e s e i n s e c t s i s r e v i e w e d , but i n a g r e a t m a n y c a s e s nothing a t a l l w a s known about t h e m In f a c t , s p e c i e s of w a s p s h a d to b e d e s c r i b e d a s new f r o m t h e Washington a r e a a l o n e , and t h e s y m b i o t i c m i t e s included new g e n e r a and 17 new s p e c i e s ! If t h e r e a r e p e r s o n s who f e e l t h a t d e s c r i p t i v e n a t u r a l h i s t o r y h a s < ' r u n i t s c o u r s e " (and t h e r e a r e ) , they s h o u l d b e c l o s e t e d with t h i s book f o r a few d a y s K r o m b e i n i s known f o r h i s m e t i c u l o u s attention t o d e t a i l , and t h i s book i s no e x c e p t i o n I t i s a g r e a t c o m p e n d i u m of f a c t s , g a t h e r e d with g r e a t Published by ValpoScholar, 2067 ...Rathcke et al.: Observations on the Interrelationships among Ants, Aphids, and Ap 1967 THE MICHIGAN ENTOMOLOGIST 169 OBSERVATIONS ON THE INTERRELATIONSHIFS AMONG ANTS, APHIDS AND APHID PREDATORS... al.: Observations on the Interrelationships among Ants, Aphids, and Ap 1967 THE MICHIGAN ENTOMOLOGIST 171 tended aphids was not c o n s i s t e n t with the o b s e r v a t i o n s made on o... al.: Observations on the Interrelationships among Ants, Aphids, and Ap 1967 Nixon, G.E.J London THE MICHIGAN ENTOMOLOGIST 173 1951 T h e a s s o c i a t i o n of a n t s with a p h i d s and c