1974] 
Moglich & Holldobler \ — Nest Moving of Ants 
225 
grasp the females at the mandibles and pull them to the target area 
(Fig. 5). The first behavioral steps that lead to this pulling be- 
havior are usually similar to those which initiate carrying behavior 
in workers. 
The methods by which males are transported are notably different, 
however. These individuals are sometimes picked up at any part of 
the body and dragged or carried to the target area. In a few species 
workers apply specific stereotyped carrying methods for males. In 
several Camponotus species we observed that most of the males are 
grasped at the “neck” and lifted into an oblique position in which 
they are carried away (Fig. 6a). During transportation the males 
remain motionless with the antennae and legs folded to the body. As 
Fig. 6 b and c shows, there are occasionally exceptions of this carry- 
ing technique. 
In some other species, as in N ovomessor cocker elli and Aphaeno- 
gaster floridanus , workers grasp the males between the thorax and 
gaster and carry them beneath their body between their legs (Fig. 7). 
Division of Labor During Nest Movings 
Social carrying behavior in ants can serve many purposes; it is, 
however, most frequently employed during nest emigrations. To- 
gether with the tandem running technique it can be considered to be 
a rather primitive recruitment method. In both cases, each recruiting 
ant can only recruit one nestmate at one time. This leads to the 
question: Do all workers of a colony have the same carrier or tan- 
dem-leader potential, or is there instead a group of specialists, who 
organize nest movings. To investigate this matter we chose two 
formicine species, one which moves almost exclusively by using the 
adult transport method ( Formica sanguinea) and another which 
primarily employs the tandem running technique ( Camponotus seri- 
ceus) . 
Nest emigrations can be induced in the laboratory by keeping the 
Fig. 3. The behavioral sequences that initiate carrying behavior. 1. The 
recruiter ant (black) approaches a nestmate (white) and conducts the 
jerking response for 2-3 seconds. 2. The recruiter grasps the nestmate at 
the mandibles and pulls it back for about 2-20 cm. 3. When the recruiter 
turns, it holds the nestmate with a firm grip. The nestmate is thereby 
slightly lifted. 4. The nestmate folds its legs and antennae tightly to the 
body and rolls its gaster inward. 5. In this posture it is carried to the 
target area. The arrows indicate the direction of the movements. These 
sequences were based on a film analysis. (After Holldobler, Moglich, 
Maschwitz 1974; illustrations by Turid Holldobler). 
