Excerpt: The literature concerning the interactions of these insect groups is as interesting as it is extensive. It has generally been accepted that the relationship between ants and aphids has been one of mutualism in which ants derive all or a large part of their nutrients from aphid honeydew and perform special functions in return, primarily that of protecting aphids from their enemies. Every aspect of these associations has been covered by the excellent reviews of Nixon (1951) and Way (1963). It now appears that the nature and effectiveness of the protective value afforded the attended aphids by ants cannot be reduced to a simple statement of symbiosis. Nixon (1951) pointed out numerous situations that are responsible for fluctuations in the ant-aphid relationship. The emphasis in the present study was on the protection extended to Aphis rumicis Linn., a small aphid found on spirea shoots by the common tending ants. This restriction allowed constant surveillance of the protection supplied to a single aphid species by ants against the seasonal succession of predators. Aphid parasites were ignored in this study

Glass, Arthur W

Hamrum, Charles L

Rathcke, Beverly

English

Valparaiso University

The Great Lakes Entomologist Volume 1 Number 5 -- October 1967 Number 5 -- October 1967 Article 5 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 1 (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. 1967 THE MICHIGAN ENTOMOLOGIST 169 OBSERVATIONS ON THE INTERRELATIONSHIFS AMONG ANTS, APHIDS AND APHID PREDATORS B e v e r l y  Rathcke,  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 Biology,  Gus tavus  Adolphus College St. P e t e r ,  Minnesota  56082 The  l i t e r a t u r e  concern ing  the  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 extensive.  I t  h a s  genera l ly  b e e n  accepted tha t  the  re la t ionsh ip  between an ts  and aphids h a s  been  one  of mutua l i sm i n  which an ts  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  nu t r ien t s  f r o m  aphid honeydew and p e r f o r m  s p e c i a l  funct ions i n  r e t u r n ,  p r i m a r i l y  tha t  of p ro tec t ing  aphids 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  assoc ia -  t ions h a s  been  covered  by the exce l len t  r e v i e w s  of Nixon (1951) and Way (1963). I t  now a p p e a r s  tha t  the  n a t u r e  and e f fec t iveness  of the p ro tec t ive  value afforded the at tended aphids by an ts  cannot  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 out  n u m e r o u s  s i tua t ions  tha t  a r e  respons ib le  f o r  f luctuat ions i n  the  ant-aphid r e l a -  t ionship.  The  e m p h a s i s  in  the  p r e s e n t  s tudy was on the  protect ion extended to Aphis m m i c i s  Linn.,  a s m a l l  aphid found on s p i r e a  shoots  by t h e  common tending an ts .  T h i s  r e s t r i c t i o n  allowed cons tan t  s u r v e i l -  l a n c e  of the  protect ion supplied to  a s i n g l e  aphid s p e c i e s  by an ts  aga ins t  the  s e a s o n a l  success ion  of p r e d a t o r s .  Aphid p a r a s i t e s  w e r e  ignored  in th i s  study. T h e s e  observa t ions  w e r e  m a d e  f r o m  June  to 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  bushes .  Only two a n t s ,  Lasius neonigev E m e r y  and Fovmica fusca Linnaeus  w e r e  i n  r e g u l a r  a t tendance to  Aphis vumicis. The r e s p o n s e s  of t h e s e  an t s  to the aphid and to the f ive  p r e d a t o r  g roups  w e r e  s o  d i f fe ren t  t h a t  e a c h  an t  was  t r e a t e d  s e p a r a t e l y .  The  p r e d a t o r  s p e c i e s  w e r e  not  ident i f ied i n  al l  ins tances .  Spec ies  of Chrysopidae ,  Cocc ine l l idae ,  Syrphidae,  C h a m a e -  myiidae,  and Cecidomyiidae at tacked the tended aphids. The p r e d a t o r  protect ion provided by the  a n t s  was  the  p r i m e  ob jec t  of the study. T h e r e f o r e ,  oppor tun i t i es  to identify p r e d a c e o u s  l a r v a e  w e r e  frequent ly s a c r i f i c e d  f o r  potent ial  observa t ion  of an t -preda tor  encounte rs .  Lasius neonigev Even  though many s y r p h i d  l a r v a e  preyed  upon tended aphids ,  in  only one c a s e  was  a syrph id  l a r v a  at tacked by Lasius neonigev, and i n  th i s  ins tance  the  at tack m a y  have  been  prec ip i ta ted  by the  ac t ions  of the  o b s e r v e r  a s  t h e  l a r v a  was  placed 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  the aphid feeding ac t iv i t i es  of Aphidoletes (Cecidomyii-  dae)  and Leucopis (Chamaemyi idae)  was  even l e s s  than i n  the c a s e  of syrph ids .  On one  s p i r e a  bush  t h e s e  p r e d a t o r s  comple te ly  ki l led the  aphid population being tended. Ants often o v e r r a n  the l a r v a e  and 1Rathcke et al.: Observations on the Interrelationships among Ants, Aphids, and ApPublished by ValpoScholar, 2067170 THE MICHIGAN ENTOMOLOGIST Vol. 1, No. 5 tapped them with t he i r  antennae. Only one Leucopis l a r v a  was  dis-  tu rbed  by these  at tent ions and lef t  the branch  while the o the r s  remained  undisturbed. Leucopis pupae we re  commonly found among tended aphids. These  an ts  were  never  observed  to d i s turb  any d ip te ran  eggs.  The r e sponse s  of Lasius neoniger to  coccinel l id l a rvae  s e e m e d  to depend upon the s i z e  of the l a rvae .  Smal l  l a r v a e ,  3 to 4 m m .  long, were  not at tacked.  The  ants  would, however,  invest igate  t h e i r  bodies  with antennae and palps.  These  s m a l l  l a r v a e  would r e m a i n  per fec t ly  quiet during these  at tent ions,  but would continue feeding o r  search ing  a s  soon a s  the an ts  would leave  them. T h e l a r g e r  coccinel l id l a r v a e  were  always attacked. One l a r v a ,  about 8 m m .  long,  was dragged  down a branch  by t h r e e  ants .  However ,  the s m a l l  Lasius worke r s  we re  usual ly unable to inf l ict  much damage  on  the l a r g e  l a rvae .  In a l l  observed  i n s t ances ,  the tending ants  did succeed  in driving l a r g e  coccinel l id l a rvae  away f r o m  aphids. Chrysopid  (Neuroptera)  l a r v a e  always exci ted L. neonigev whenever they we re  encountered n e a r  aphids. Some ants  succeeded  i n  biting a s m a l l  chrysopid l a r v a e  and c a r r i e d  the s t r ugg l i ng l a rva  away to the nest .  Many an ts  we re  r equ i r ed  to i n t e r f e r e  success fu l ly  with the search ing  and feeding ac t iv i t i es  of the l a r g e r  lacewing l a rvae .  Whenever the tending an ts  would a t tack ,  the l a r v a  would r a i s e  the t ip  of the abdomen and exude a drople t  of brown liquid. The l a r v a  would then twis t  about t ry ing  to touch an ant  with this  droplet .  Any an t  touched by the drople t  would f lee f r o m  the scene .  Soon all the  nearby  an ts  would ga the r  and t r y  to at tack the la rva .  Many we re  repe l led  before  the l a r v a w a s  finally dr iven  away. Whenever L. neoniger occu r r ed  i n  sufficient n u m b e r s ,  they s eemed  to b e  quite capable of protect ing t he i r  tended aphids against  the l a r g e r  chrysopid and coccinel l id la rvae .  However,  they did not s e e m  to recog- n ize  the Dip te ra  l a rvae  a s  i n t rude r s .  These  l a r v a e  we re  general ly quite pa s s ive  and ord inar i ly  did not make  exci ted movements  when touched by the ants .  These  an ts  appeared to t r e a t  them a s  they would t r e a t  an aphid, tapping them with t he i r  antennae and often palpat ing them. These  p r eda to r s  did not s e e m  to be  fore ign  to th i s  Lasius spec ies .  Pontin ( 1 9 5 9 )  l i s t s  two syrphid  spec i e s  and Chamaemyiidae and Cecidomyiidae l a rvae  among the p r eda to r s  adapted to at tack aphids tended by ants. Our  observa t ions  suppor t  Pontin 's  c l a im  that  c e r t a i n  p r eda to r s  a f e  to le ra ted  by Lasius and Formica spec i e s .  Lasius neoniger was observed  tending populations of a waxy coccid,  Planococcus citri Ris so  , on chrysanthemums.  Some s p i r e a  aphids we re  t r a n s f e r r e d  to the 'mum '  plants .  After  having se t t l ed  on the plant ,  they were  quickly s e i zed  and c a r r i e d  off to the ne s t  by the ants. Within a few minutes  al l  of the < ' immigrant"  aphids we re  removed.  Contradictory r e su l t s  we re  obtained when cocc ids  were  added to an aphid population on sp i r ea .  In this  ins tance  the " immigrants"  were  accepted by the an ts  which r egu l a r l y  sol ici ted honeydew f r o m  them. The cocc ids  remained  on the s p i r e a  fo r  s e v e r a l  days.  No  explanation i s  evident  f o r  the differ-  en t  r e sponse s  by L. neonigev toward these  two homopteran spec ies .  The r e sponse  of L. neo.nig2.v to the p r e sence  of p r eda to r s  among 2The Great Lakes Entomologist, Vol. 1, No. 5 [2067], Art. 5https://scholar.valpo.edu/tgle/vol1/iss5/51967 THE MICHIGAN ENTOMOLOGIST 171  tended aphids was not consistent  with the observations made on o ther  Lasius species.  Way (1963) did not cons ider  Lasius species  to be especial ly predaceous o r  host i le .  However, El-Ziady and Kennedy (1956) showed that  Lasius n i g e ~ c a r r i e d  off o r  drove  away l a rvae  of Syrphidae and of the coccinellid Adalia bipunctata Linn. but did not h a r m  syrphid o r  coccinellid eggs. Adult coccinell ids were a lso  attacked by L. niger (Linn). Banks (1962) confirmed that  L. niger attacked syrphid and coccinel- lid la rvae  and drove  away chrysopid la rvae  a s  well a s  adult and im- mature  Anthocoridae (Hemiptera) .  Banks (1962) also observed the removal  of eggs of syrphids  and coccinell ids by this spec ies .  Conflicting with these  observations a r e  the investigations of Herzig (1938) and Pontin (1959). Herzig concluded that  ants r e a c t  to swift moving bodies a s  a threa t  to themselves and do not fight in t ruders  to protect  the i r  aphids. His exper iments  included observations of t h r ee  Lasius species.  Pontin found aphid p reda to r s  in the nes ts  of L. niger and L. ~ Z U V U S  (Fab.). Fomica fusca During the ea r ly  s u m m e r ,  syrphid,  Leucopis, chrysopid,  and coc- cinellid eggs were found on an aphid-infested s p i r e a  bush tended by a l a rge  colony of Formica f u ~ c a .  However,  during this period predators  were r a r e ly  seen.  When l a rvae  were t r ans fe r r ed  to the bush they were attacked a s  soon a s  discovered.  These l a r g e r  ants  swiftly b i t  and c a r r i e d  to the nes t  al l  introduced syrphid,  Leucopis, and Aphidoletes larvae.  All chrysopid and l a rge  coccinellid la rvae  were  also promptly attacked. The ants  were often repelled by l a r g e r  chrysopids in the s a m e  msnner  that  they discouraged L. neoniger. One Formica was repulsed by a secre t ion  f rom a wounded coccinellid la rva .  A s imi l a r  response  was noted when a Fomica which became agitated re t rea ted  f rom a wounded Leucopis. The coccinellid la rvae  usually escaped the attacking ants by dropping to the ground. In one instance severa l  sma l l  coccinellid l a rvae  were observed among the tended aphids and were not attacked. In this  instance F. fusca behaved a s  L. neoniger. Aside f rom the sma l l  coccinell ids,  F. f u ~ c a  had no difficulty recog- nizing and dealing with aphid predators .  During th is  period the aphid population was quite stable. The effect iveness of the i r  protection be- c a m e  apparent  when they abruptly ceased  tending the aphids nea r  the end of July,  s eve ra l  days before the winged ants appeared in August. This s p i r e a  bush was rapidly colonized by the Diptera p reda to r s ,  a coc- cinellid and a debris-collecting chrysopid. Thereaf te r ,  the aphid popu- lation was short ly decimated. In the only observed instance of egg predation,  a F. fusca was seen devouring a chrysopid egg f rom a bent  egg stalk. The ants  were also of benefit to the aphids by keeping them clean. In untended a r e a s  the aphids on s t em t ips became sticky and clogged with c a s t  skins and debris .  3Rathcke et al.: Observations on the Interrelationships among Ants, Aphids, and ApPublished by ValpoScholar, 2067172 THE MICHIGAN ENTOMOLOGIST Vol. 1, NO. 5 The  l i t e r a t u r e  does  not indicate  tha t  European  F o v m i c a a r e  formid-  able aphid pro tec tors .  Nixon (1951) does  not provide evidence of aggres -  s ive  protect ion by Fovmica.  Pontin (1959) c i t e s  to le rance  of Coccinel la  divaricata 01. by Fovmica  sanguinea La t r .  Fovmica  yufa Linn. i s  r e -  garded  by Way (1963) a s  only a par t ia l ly  success fu l  syrphid  preda tor .  Both ant spec i e s  observed  in th i s  study appeared to exhibi t  some  f o r m  of ownership toward the attended aphids. A L a s i u s  neonigev, cap- t u r ed  on  a bush and t r a n s f e r r e d  to another  bush 1 5  f ee t  away, was killed by the nat ive L. neonigev population. A L a s i u s  introduced onto the Fovmica-tended bush was a l so  ki l led with dispatch.  Col la te ra l  evidence of aphid "ownership" may  be  i n f e r r ed  f r o m  the reac t ions  of these  an ts  to aphid p r eda to r s  in  the absence  of aphids. A l a r g e  coccinel l id l a r v a  was placed on the ground n e a r  the Fovmica  fusca  nest .  I t  was completely ignored by the  ants  a s  i t  c rawled  on the ground. When t h i s  l a r v a  was placed on a tendedbranch  i t  was prompt ly  at tacked and forced  to d rop  to the apparen t  sa fe ty  of the  ground. Chrysopid l a rvae  on  the ground responded defensively t o w a r d L a s i u ~  which in  tu rn  ignored the l a rvae .  The  agg re s s ive  behavior  s e e m s  to be exhibi ted only under  conditions in  which ownership of t e r r i t o r y  ex is t s .  Undoubtedly some  populations of Aphis vumic is  p ro spe red  through the at tent ions of F. fusca and L. neonigev. Fovmica  fu sca  was v e r y  effect ive in protect ing the  aphids and kept  a s tab le  population of aphids until  August. L a s i u s  neonigev appeared  to be  m o r e  e r r a t i c  in  tending and would d e s e r t  the aphids f o r  days. They we re  not nea r l y  s o  effective a s  p ro t ec to r s  against  p reda tors .  The  na tu r e  of th i s  protect ion may  o r  may  not be  a defense of "owned" aphids. The  variabi l i ty  of the  da t a  p r e sen t ed  by respons ib le  o b s e r v e r s  cer ta in ly  ind ica tes  tha t  these  in te r -  re la t ionsh ips  among an t s ,  aphids,  and aphid p r eda to r s  fluctuate. More s tudy i s  needed to define the r e a s o n s  fo r  t he se  fluctuations. P e r h a p s  the ant  spec i e s  involved i s  chiefly respons ib le  f o r  the var ie ty  of e f f ec t s  observed  in  these  in te rspec i f ic  relat ionships.  ACKNOWLEDGMENTS The au thors  a r e  indebted to Edmund Olson,  Depar tment  of Entomol- ogy, Universi ty of Minnesota,  St. Pau l ,  Minnesota,  fo r  identifying the homopteran  spec i e s  utilized in  this  study. We a l so  thank Violet Neal f o r  h e r  aid in  the final p repara t ion  of the manuscr ip t .  This  s tudy was suppor ted  by the Gustavus Resea r ch  Fund. LITERATURE CITED Banks ,  C.J .  1962. Ef fec t s  of the ant  Lasius nigev(L.)  on i n sec t s  preying on s m a l l  populations of Aphis fabae Scop. on bean plants. Ann. Appl. Biol.  50:669-679. El-Ziady,  S. and J.S. Kennedy. 1956. Beneficial  e f fec t s  of the common garden  an t ,  L a s i u s  n i g e r  L . ,  on the black bean aphid,  Aphis fabae Scopoli. Roy. Entomol. Soc. London P roc .  (A) 31:61-65. Herz ig ,  J. 1938. Ameisen und Blattl'duse. Z. Angew. Entomol. 24:367- 435. 4The Great Lakes Entomologist, Vol. 1, No. 5 [2067], Art. 5https://scholar.valpo.edu/tgle/vol1/iss5/51967 THE MICHIGAN ENTOMOLOGIST 173 Nixon, G.E.J. 1951. The  assoc ia t ion  of an t s  with aphids and coccids.  London. Pont in ,  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 ack  aphids at tended by ants .  Entomol.  Monthly Mag. 95: 154-155. Way, M.J. 1963. Mutual ism between an ts  and honeydew-producing Homoptera.  Ann. Rev. Entomol.  8:307-344. REVIEWS OF RECENT LITERATURE TRAP-NESTING WASPS AND BEES: L I F E  HISTORIES, NESTS. AND ASSOCIATES. K a r l  V. Krombein.  Washington,  D.C.: The  Smithsonian Inst i tut ion,  1957. Smithsonian Publ. 4670. vi ,  570 pp. $12.50. The  technique of "trap-nesting" f o r  wasps  and b e e s  by putting out  s t r i p s  of wood having a hole  b o r e d  in one end i s  not  exact ly new, but  only within the  l a s t  1 5  y e a r s  h a s  i t  been widely employed in th i s  coun- t ry .  This  new book by K a r l  V. Krombein ,  c h a i r m a n  of the Depar tment  of Entomology a t  the U.S. National Museum,  r e v e a l s  how enormous ly  product ive the technique can  be. I t  i s ,  of c o u r s e ,  useful  only f o r  s p e c i e s  tha t  normal ly  n e s t  in  hollow twigs.  Spec ies  tha t  b o r e  in  pith (such a s  many  c r a b r o n i n e  wasps)  do not usual ly accep t  the  t r a p s ,  nor  do ground- n e s t e r s  (which m a k e  up  the m a j o r i t y  of wasps  and b e e s ) .  Despi te  i t s  l imi ta t ions ,  the t rap-nes t ing  technique i s  e x t r e m e l y  valuable,  a s  i t  a l lows one to follow the development  of the  c e l l  con- t e n t s  c lose ly  once the n e s t  h a s  been s p l i t  open,  and a l s o  b e c a u s e  the n e s t s  a r e  s o  s i m p l e  and inexpensive tha t  i t  i s  p rac t icab le  to u s e  l a r g e  n u m b e r s  of t h e m ,  thus obtaining in format ion  on ecology and population phenomena  not avai lable  f r o m  i so la ted  n e s t s .  The tech- nique is a l so  useful  in  s u m m e r  p r o g r a m s  f o r  young people;  t h e r e  i s  probably no equally s i m p l e  method f o r  demons t ra t ing  s o m e  of the complex i t i es  in  na tura l  env i ronments .  Krombein ' s  s tudy is based  on d a t a  gained f r o m  about 3400 t r a p - n e s t s  put ou t  o v e r  a 12-year  per iod  in s e v e r a l  e a s t e r n  s t a t e s  and Arizona.  In  a l l ,  h e  t r e a t s  75 s p e c i e s  of wasps  tha t  accepted 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 di-  v e r s e  g roups .  P r e v i o u s  knowledge of t h e s e  i n s e c t s  i s  rev iewed,  but in  a g r e a t  many  c a s e s  nothing a t  a l l  was  known about them.  In fac t ,  5 s p e c i e s  of wasps  had to b e  d e s c r i b e d  a s  new f r o m  the Washington a r e a  a lone ,  and the symbiot ic  m i t e s  included 2 new g e n e r a  and 17 new spec ies !  If t h e r e  a r e  p e r s o n s  who f e e l  tha t  descr ip t ive  n a t u r a l  h i s t o r y  h a s  < ' r u n  i t s  course"  (and t h e r e  a r e ) ,  they should b e  c lose ted  with th i s  book f o r  a few days.  Krombein  i s  known f o r  h i s  met icu lous  attention to de ta i l ,  and th i s  book i s  no except ion.  I t  i s  a g r e a t  compendium of f a c t s ,  g a t h e r e d  with g r e a t  5Rathcke et al.: Observations on the Interrelationships among Ants, Aphids, and ApPublished by ValpoScholar, 2067

Observations on the Interrelationships among Ants, Aphids, and Aphid Predators

ValpoScholar

The Great Lakes Entomologist 
Volume 1 
Number 5 -- October 1967 Number 5 -- October 
1967 
Article 5 
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 1 (5) 
Available at: https://scholar.valpo.edu/tgle/vol1/iss5/5 
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1967 THE MICHIGAN ENTOMOLOGIST 169 
OBSERVATIONS ON THE INTERRELATIONSHIFS AMONG 
ANTS, APHIDS AND APHID PREDATORS 
B e v e r l y  Rathcke,  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 Biology,  Gus tavus  Adolphus College 
St. P e t e r ,  Minnesota  56082 
The  l i t e r a t u r e  concern ing  the  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 extensive.  I t  h a s  genera l ly  b e e n  accepted tha t  
the  re la t ionsh ip  between an ts  and aphids h a s  been  one  of mutua l i sm i n  
which an ts  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  nu t r ien t s  f r o m  aphid 
honeydew and p e r f o r m  s p e c i a l  funct ions i n  r e t u r n ,  p r i m a r i l y  tha t  of 
p ro tec t ing  aphids 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  assoc ia -  
t ions h a s  been  covered  by the exce l len t  r e v i e w s  of Nixon (1951) and 
Way (1963). I t  now a p p e a r s  tha t  the  n a t u r e  and e f fec t iveness  of the 
p ro tec t ive  value afforded the at tended aphids by an ts  cannot  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 out  n u m e r o u s  
s i tua t ions  tha t  a r e  respons ib le  f o r  f luctuat ions i n  the  ant-aphid r e l a -  
t ionship.  The  e m p h a s i s  in  the  p r e s e n t  s tudy was on the  protect ion 
extended to Aphis m m i c i s  Linn.,  a s m a l l  aphid found on s p i r e a  shoots  
by t h e  common tending an ts .  T h i s  r e s t r i c t i o n  allowed cons tan t  s u r v e i l -  
l a n c e  of the  protect ion supplied to  a s i n g l e  aphid s p e c i e s  by an ts  aga ins t  
the  s e a s o n a l  success ion  of p r e d a t o r s .  Aphid p a r a s i t e s  w e r e  ignored  
in th i s  study. 
T h e s e  observa t ions  w e r e  m a d e  f r o m  June  to 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  bushes .  Only two a n t s ,  
Lasius neonigev E m e r y  and Fovmica fusca Linnaeus  w e r e  i n  r e g u l a r  
a t tendance to  Aphis vumicis. The r e s p o n s e s  of t h e s e  an t s  to the aphid 
and to the f ive  p r e d a t o r  g roups  w e r e  s o  d i f fe ren t  t h a t  e a c h  an t  was  
t r e a t e d  s e p a r a t e l y .  The  p r e d a t o r  s p e c i e s  w e r e  not  ident i f ied i n  al l  
ins tances .  Spec ies  of Chrysopidae ,  Cocc ine l l idae ,  Syrphidae,  C h a m a e -  
myiidae,  and Cecidomyiidae at tacked the tended aphids. The p r e d a t o r  
protect ion provided by the  a n t s  was  the  p r i m e  ob jec t  of the study. 
T h e r e f o r e ,  oppor tun i t i es  to identify p r e d a c e o u s  l a r v a e  w e r e  frequent ly 
s a c r i f i c e d  f o r  potent ial  observa t ion  of an t -preda tor  encounte rs .  
Lasius neonigev 
Even  though many s y r p h i d  l a r v a e  preyed  upon tended aphids ,  in  only 
one c a s e  was  a syrph id  l a r v a  at tacked by Lasius neonigev, and i n  th i s  
ins tance  the  at tack m a y  have  been  prec ip i ta ted  by the  ac t ions  of the  
o b s e r v e r  a s  t h e  l a r v a  was  placed 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  the aphid feeding ac t iv i t i es  of Aphidoletes (Cecidomyii-  
dae)  and Leucopis (Chamaemyi idae)  was  even l e s s  than i n  the c a s e  of 
syrph ids .  On one  s p i r e a  bush  t h e s e  p r e d a t o r s  comple te ly  ki l led the  
aphid population being tended. Ants often o v e r r a n  the l a r v a e  and 
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170 THE MICHIGAN ENTOMOLOGIST Vol. 1, No. 5 
tapped them with t he i r  antennae. Only one Leucopis l a r v a  was  dis-  
tu rbed  by these  at tent ions and lef t  the branch  while the o the r s  remained  
undisturbed. Leucopis pupae we re  commonly found among tended aphids. 
These  an ts  were  never  observed  to d i s turb  any d ip te ran  eggs.  
The r e sponse s  of Lasius neoniger to  coccinel l id l a rvae  s e e m e d  to 
depend upon the s i z e  of the l a rvae .  Smal l  l a r v a e ,  3 to 4 m m .  long, were  
not at tacked.  The  ants  would, however,  invest igate  t h e i r  bodies  with 
antennae and palps.  These  s m a l l  l a r v a e  would r e m a i n  per fec t ly  quiet 
during these  at tent ions,  but would continue feeding o r  search ing  a s  soon 
a s  the an ts  would leave  them. T h e l a r g e r  coccinel l id l a r v a e  were  always 
attacked. One l a r v a ,  about 8 m m .  long,  was dragged  down a branch  by 
t h r e e  ants .  However ,  the s m a l l  Lasius worke r s  we re  usual ly unable to 
inf l ict  much damage  on  the l a r g e  l a rvae .  In a l l  observed  i n s t ances ,  the 
tending ants  did succeed  in driving l a r g e  coccinel l id l a rvae  away f r o m  
aphids. 
Chrysopid  (Neuroptera)  l a r v a e  always exci ted L. neonigev whenever 
they we re  encountered n e a r  aphids. Some ants  succeeded  i n  biting a 
s m a l l  chrysopid l a r v a e  and c a r r i e d  the s t r ugg l i ng l a rva  away to the nest .  
Many an ts  we re  r equ i r ed  to i n t e r f e r e  success fu l ly  with the search ing  and 
feeding ac t iv i t i es  of the l a r g e r  lacewing l a rvae .  Whenever the tending 
an ts  would a t tack ,  the l a r v a  would r a i s e  the t ip  of the abdomen and exude 
a drople t  of brown liquid. The l a r v a  would then twis t  about t ry ing  to 
touch an ant  with this  droplet .  Any an t  touched by the drople t  would f lee 
f r o m  the scene .  Soon all the  nearby  an ts  would ga the r  and t r y  to at tack 
the la rva .  Many we re  repe l led  before  the l a r v a w a s  finally dr iven  away. 
Whenever L. neoniger occu r r ed  i n  sufficient n u m b e r s ,  they s eemed  
to b e  quite capable of protect ing t he i r  tended aphids against  the l a r g e r  
chrysopid and coccinel l id la rvae .  However,  they did not s e e m  to recog- 
n ize  the Dip te ra  l a rvae  a s  i n t rude r s .  These  l a r v a e  we re  general ly 
quite pa s s ive  and ord inar i ly  did not make  exci ted movements  when 
touched by the ants .  These  an ts  appeared to t r e a t  them a s  they would 
t r e a t  an aphid, tapping them with t he i r  antennae and often palpat ing them. 
These  p r eda to r s  did not s e e m  to be  fore ign  to th i s  Lasius spec ies .  Pontin 
( 1 9 5 9 )  l i s t s  two syrphid  spec i e s  and Chamaemyiidae and Cecidomyiidae 
l a rvae  among the p r eda to r s  adapted to at tack aphids tended by ants. 
Our  observa t ions  suppor t  Pontin 's  c l a im  that  c e r t a i n  p r eda to r s  a f e  
to le ra ted  by Lasius and Formica spec i e s .  
Lasius neoniger was observed  tending populations of a waxy coccid,  
Planococcus citri Ris so  , on chrysanthemums.  Some s p i r e a  aphids we re  
t r a n s f e r r e d  to the 'mum '  plants .  After  having se t t l ed  on the plant ,  they 
were  quickly s e i zed  and c a r r i e d  off to the ne s t  by the ants. Within a few 
minutes  al l  of the < ' immigrant"  aphids we re  removed.  Contradictory 
r e su l t s  we re  obtained when cocc ids  were  added to an aphid population 
on sp i r ea .  In this  ins tance  the " immigrants"  were  accepted by the an ts  
which r egu l a r l y  sol ici ted honeydew f r o m  them. The cocc ids  remained  
on the s p i r e a  fo r  s e v e r a l  days.  No  explanation i s  evident  f o r  the differ-  
en t  r e sponse s  by L. neonigev toward these  two homopteran spec ies .  
The r e sponse  of L. neo.nig2.v to the p r e sence  of p r eda to r s  among 
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1967 THE MICHIGAN ENTOMOLOGIST 171  
tended aphids was not consistent  with the observations made on o ther  
Lasius species.  Way (1963) did not cons ider  Lasius species  to be 
especial ly predaceous o r  host i le .  However, El-Ziady and Kennedy (1956) 
showed that  Lasius n i g e ~ c a r r i e d  off o r  drove  away l a rvae  of Syrphidae 
and of the coccinellid Adalia bipunctata Linn. but did not h a r m  syrphid 
o r  coccinellid eggs. Adult coccinell ids were a lso  attacked by L. niger 
(Linn). 
Banks (1962) confirmed that  L. niger attacked syrphid and coccinel- 
lid la rvae  and drove  away chrysopid la rvae  a s  well a s  adult and im- 
mature  Anthocoridae (Hemiptera) .  Banks (1962) also observed the 
removal  of eggs of syrphids  and coccinell ids by this spec ies .  
Conflicting with these  observations a r e  the investigations of Herzig 
(1938) and Pontin (1959). Herzig concluded that  ants r e a c t  to swift 
moving bodies a s  a threa t  to themselves and do not fight in t ruders  to 
protect  the i r  aphids. His exper iments  included observations of t h r ee  
Lasius species.  Pontin found aphid p reda to r s  in the nes ts  of L. niger 
and L. ~ Z U V U S  (Fab.). 
Fomica fusca 
During the ea r ly  s u m m e r ,  syrphid,  Leucopis, chrysopid,  and coc- 
cinellid eggs were found on an aphid-infested s p i r e a  bush tended by a 
l a rge  colony of Formica f u ~ c a .  However,  during this period predators  
were r a r e ly  seen.  When l a rvae  were t r ans fe r r ed  to the bush they were 
attacked a s  soon a s  discovered.  These l a r g e r  ants  swiftly b i t  and 
c a r r i e d  to the nes t  al l  introduced syrphid,  Leucopis, and Aphidoletes 
larvae.  All chrysopid and l a rge  coccinellid la rvae  were  also promptly 
attacked. The ants  were often repelled by l a r g e r  chrysopids in the 
s a m e  msnner  that  they discouraged L. neoniger. One Formica was 
repulsed by a secre t ion  f rom a wounded coccinellid la rva .  A s imi l a r  
response  was noted when a Fomica which became agitated re t rea ted  
f rom a wounded Leucopis. The coccinellid la rvae  usually escaped 
the attacking ants by dropping to the ground. In one instance severa l  
sma l l  coccinellid l a rvae  were observed among the tended aphids and 
were not attacked. In this  instance F. fusca behaved a s  L. neoniger. 
Aside f rom the sma l l  coccinell ids,  F. f u ~ c a  had no difficulty recog- 
nizing and dealing with aphid predators .  During th is  period the aphid 
population was quite stable. The effect iveness of the i r  protection be- 
c a m e  apparent  when they abruptly ceased  tending the aphids nea r  the 
end of July,  s eve ra l  days before the winged ants appeared in August. 
This s p i r e a  bush was rapidly colonized by the Diptera p reda to r s ,  a coc- 
cinellid and a debris-collecting chrysopid. Thereaf te r ,  the aphid popu- 
lation was short ly decimated. 
In the only observed instance of egg predation,  a F. fusca was seen 
devouring a chrysopid egg f rom a bent  egg stalk. 
The ants  were also of benefit to the aphids by keeping them clean. 
In untended a r e a s  the aphids on s t em t ips became sticky and clogged 
with c a s t  skins and debris .  
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172 THE MICHIGAN ENTOMOLOGIST Vol. 1, NO. 5 
The  l i t e r a t u r e  does  not indicate  tha t  European  F o v m i c a a r e  formid-  
able aphid pro tec tors .  Nixon (1951) does  not provide evidence of aggres -  
s ive  protect ion by Fovmica.  Pontin (1959) c i t e s  to le rance  of Coccinel la  
divaricata 01. by Fovmica  sanguinea La t r .  Fovmica  yufa Linn. i s  r e -  
garded  by Way (1963) a s  only a par t ia l ly  success fu l  syrphid  preda tor .  
Both ant spec i e s  observed  in th i s  study appeared to exhibi t  some  
f o r m  of ownership toward the attended aphids. A L a s i u s  neonigev, cap- 
t u r ed  on  a bush and t r a n s f e r r e d  to another  bush 1 5  f ee t  away, was killed 
by the nat ive L. neonigev population. A L a s i u s  introduced onto the 
Fovmica-tended bush was a l so  ki l led with dispatch.  
Col la te ra l  evidence of aphid "ownership" may  be  i n f e r r ed  f r o m  the 
reac t ions  of these  an ts  to aphid p r eda to r s  in  the absence  of aphids. A 
l a r g e  coccinel l id l a r v a  was placed on the ground n e a r  the Fovmica  fusca  
nest .  I t  was completely ignored by the  ants  a s  i t  c rawled  on the ground. 
When t h i s  l a r v a  was placed on a tendedbranch  i t  was prompt ly  at tacked 
and forced  to d rop  to the apparen t  sa fe ty  of the  ground. Chrysopid l a rvae  
on  the ground responded defensively t o w a r d L a s i u ~  which in  tu rn  ignored 
the l a rvae .  The  agg re s s ive  behavior  s e e m s  to be exhibi ted only under  
conditions in  which ownership of t e r r i t o r y  ex is t s .  
Undoubtedly some  populations of Aphis vumic is  p ro spe red  through 
the at tent ions of F. fusca and L. neonigev. Fovmica  fu sca  was v e r y  
effect ive in protect ing the  aphids and kept  a s tab le  population of aphids 
until  August. L a s i u s  neonigev appeared  to be  m o r e  e r r a t i c  in  tending 
and would d e s e r t  the aphids f o r  days. They we re  not nea r l y  s o  effective 
a s  p ro t ec to r s  against  p reda tors .  The  na tu r e  of th i s  protect ion may  o r  
may  not be  a defense of "owned" aphids. The  variabi l i ty  of the  da t a  
p r e sen t ed  by respons ib le  o b s e r v e r s  cer ta in ly  ind ica tes  tha t  these  in te r -  
re la t ionsh ips  among an t s ,  aphids,  and aphid p r eda to r s  fluctuate. More 
s tudy i s  needed to define the r e a s o n s  fo r  t he se  fluctuations. P e r h a p s  
the ant  spec i e s  involved i s  chiefly respons ib le  f o r  the var ie ty  of e f f ec t s  
observed  in  these  in te rspec i f ic  relat ionships.  
ACKNOWLEDGMENTS 
The au thors  a r e  indebted to Edmund Olson,  Depar tment  of Entomol- 
ogy, Universi ty of Minnesota,  St. Pau l ,  Minnesota,  fo r  identifying the 
homopteran  spec i e s  utilized in  this  study. We a l so  thank Violet Neal 
f o r  h e r  aid in  the final p repara t ion  of the manuscr ip t .  This  s tudy was 
suppor ted  by the Gustavus Resea r ch  Fund. 
LITERATURE CITED 
Banks ,  C.J .  1962. Ef fec t s  of the ant  Lasius nigev(L.)  on i n sec t s  preying 
on s m a l l  populations of Aphis fabae Scop. on bean plants. Ann. Appl. 
Biol.  50:669-679. 
El-Ziady,  S. and J.S. Kennedy. 1956. Beneficial  e f fec t s  of the common 
garden  an t ,  L a s i u s  n i g e r  L . ,  on the black bean aphid,  Aphis fabae 
Scopoli. Roy. Entomol. Soc. London P roc .  (A) 31:61-65. 
Herz ig ,  J. 1938. Ameisen und Blattl'duse. Z. Angew. Entomol. 24:367- 
435. 
4
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1967 THE MICHIGAN ENTOMOLOGIST 173 
Nixon, G.E.J. 1951. The  assoc ia t ion  of an t s  with aphids and coccids.  
London. 
Pont in ,  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 ack  aphids at tended by ants .  Entomol.  Monthly Mag. 95: 154-155. 
Way, M.J. 1963. Mutual ism between an ts  and honeydew-producing 
Homoptera.  Ann. Rev. Entomol.  8:307-344. 
REVIEWS OF RECENT LITERATURE 
TRAP-NESTING WASPS AND BEES: L I F E  HISTORIES, NESTS. AND 
ASSOCIATES. K a r l  V. Krombein.  Washington,  D.C.: The  Smithsonian 
Inst i tut ion,  1957. Smithsonian Publ. 4670. vi ,  570 pp. $12.50. 
The  technique of "trap-nesting" f o r  wasps  and b e e s  by putting out  
s t r i p s  of wood having a hole  b o r e d  in one end i s  not  exact ly new, but  
only within the  l a s t  1 5  y e a r s  h a s  i t  been widely employed in th i s  coun- 
t ry .  This  new book by K a r l  V. Krombein ,  c h a i r m a n  of the Depar tment  
of Entomology a t  the U.S. National Museum,  r e v e a l s  how enormous ly  
product ive the technique can  be. I t  i s ,  of c o u r s e ,  useful  only f o r  s p e c i e s  
tha t  normal ly  n e s t  in  hollow twigs.  Spec ies  tha t  b o r e  in  pith (such a s  
many  c r a b r o n i n e  wasps)  do not usual ly accep t  the  t r a p s ,  nor  do ground- 
n e s t e r s  (which m a k e  up  the m a j o r i t y  of wasps  and b e e s ) .  
Despi te  i t s  l imi ta t ions ,  the t rap-nes t ing  technique i s  e x t r e m e l y  
valuable,  a s  i t  a l lows one to follow the development  of the  c e l l  con- 
t e n t s  c lose ly  once the n e s t  h a s  been s p l i t  open,  and a l s o  b e c a u s e  the 
n e s t s  a r e  s o  s i m p l e  and inexpensive tha t  i t  i s  p rac t icab le  to u s e  
l a r g e  n u m b e r s  of t h e m ,  thus obtaining in format ion  on ecology and 
population phenomena  not avai lable  f r o m  i so la ted  n e s t s .  The tech- 
nique is a l so  useful  in  s u m m e r  p r o g r a m s  f o r  young people;  t h e r e  i s  
probably no equally s i m p l e  method f o r  demons t ra t ing  s o m e  of the 
complex i t i es  in  na tura l  env i ronments .  
Krombein ' s  s tudy is based  on d a t a  gained f r o m  about 3400 t r a p - n e s t s  
put ou t  o v e r  a 12-year  per iod  in s e v e r a l  e a s t e r n  s t a t e s  and Arizona.  
In  a l l ,  h e  t r e a t s  75 s p e c i e s  of wasps  tha t  accepted 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 di-  
v e r s e  g roups .  P r e v i o u s  knowledge of t h e s e  i n s e c t s  i s  rev iewed,  but 
in  a g r e a t  many  c a s e s  nothing a t  a l l  was  known about them.  In fac t ,  5 
s p e c i e s  of wasps  had to b e  d e s c r i b e d  a s  new f r o m  the Washington a r e a  
a lone ,  and the symbiot ic  m i t e s  included 2 new g e n e r a  and 17 new 
spec ies !  If t h e r e  a r e  p e r s o n s  who f e e l  tha t  descr ip t ive  n a t u r a l  h i s t o r y  
h a s  < ' r u n  i t s  course"  (and t h e r e  a r e ) ,  they should b e  c lose ted  with th i s  
book f o r  a few days.  
Krombein  i s  known f o r  h i s  met icu lous  attention to de ta i l ,  and th i s  book 
i s  no except ion.  I t  i s  a g r e a t  compendium of f a c t s ,  g a t h e r e d  with g r e a t  
5
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Observations on the Interrelationships among Ants, Aphids, and Aphid Predators

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