60 
Psyche 
[Vol. 91 
Forward Jerking and Mandibular Pecking — a rapid movement 
forward and a biting at the object of intrusion. Pecking was also 
demonstrated in the absence of forward jerking. Although pecking 
behavior was easily elicited in a young colony, it appeared to be 
demonstrated in such cases almost entirely by the queen. Pecking is 
a behavior commonly used by adult females toward small hymenop- 
terous parasitoid intruders. In P. annularis, forward jerking and 
pecking have been observed by us to chase and/or discard intruding 
ants from the nest. We witnessed such behavior in older colonies 
observed in August, 1983. 
Backward Jerking — best demonstrated under strong provocation. 
Abdominal Pumping — although abdominal pumping, a rhythy- 
mic extension and retraction of the gaster, is seen as normal 
behavior in all vespids, it appears to be exaggerated during 
provocation. 
Abdominal twisting — this behavior was not evident on a young 
nest except under very strong provocation. However, when strongly 
provoked, most females of a young nest demonstrated this behavior 
as well as exaggerated abdominal pumping. 
Movement to back of nest (retreat) — a behavior that was 
demonstrated upon continuous provocation. Even females that 
demonstrate warning behaviors move to the back of the nest upon 
continuous provocation. 
Escape — this is readily demonstrated in young colonies under 
provocation. In fact, it is difficult to elicit defensive behaviors in 
young colonies and collect all of the adults because of their 
readiness to escape. Nest departure by adults of more mature 
colonies is expressed more as pseudoattack in which case adults 
return to the nest after a short erratic flight. Adults in escape flights 
do not return for a longer period of time. 
Leg waving — this behavior was demonstrated on occasion in 
young colonies and readily in mature ones. Although females of 
young colonies are reluctant to leg wave unless strongly provoked, 
this behavior was less difficult to elicit in a young colony than were 
some of the other behaviors. Leg waving frequencies may be found 
in Table III. 
None of the behaviors appeared to be demonstrated in a 
particular sequence, and many of the behaviors were demonstrated 
by some individuals and not by others. Escape and retreat were the 
most dependable behaviors in young (preemergence) colonies. 
