Observations on the Interrelationships among Ants Aphids and Ap

Number 5 -- October 1967 Number 5 -- October October 2067 Observations on the Interrelationships among Ants, Aphids, and Aphid Predators Beverly Rathcke Gustavus Adolphus College Cha

Number 5 October 1967 Number 5 October

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

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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 1 (5)

Available at: https://scholar.valpo.edu/tgle/vol1/iss5/5

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1967 T H E 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 -

t i o n s h i p T h e e m p h a s i s i n t h e p r e s e n t s t u d y w a s o n t h e p r o t e c t i o n

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

t r e a t e d s e p a r a t e l y T h e p r e d a t o r s p e c i e s w e r e n o t i d e n t i f i e d i n a l l

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

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 , 3 to 4 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 8 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

( 1 9 5 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

O u r o b s e r v a t i o n s s u p p o r t P o n t i n ' s c l a i m t h a t c e r t a i n p r e d a t o r s 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

1967 THE MICHIGAN ENTOMOLOGIST 1 7 1

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 d off 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 coccinel-

lid 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 do 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 coc-

cinellid 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 a had 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 be-

c 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 coc-

cinellid and a debris-collecting chrysopid T h e r e a f t e r , the aphid popu-

lation 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

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, cap-

t 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 1 5 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

o b s e r v e d i n t h e s e i n t e r s p e c i f i c r e l a t i o n s h i p s

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 Entomol- ogy, 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 24:367-

435

1967 T H E MICHIGAN ENTOMOLOGIST 1 7 3

Nixon, G.E.J 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

London

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 1 5 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 coun-

t 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 do ground-

n 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 con-

t 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 tech-

nique 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

c o m p l e x i t i e s i n n a t u r a l e n v i r o n m e n t s

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 1 2 - 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 , 5

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 2 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