86 
Psyche 
[Vol. 95 
in large numbers by mandibular-gland secretions that other males 
discharge upon arrival at the mating arena. Queens are subsequently 
“lured” to these arenas by the collectively-secreted male pheromone 
(Holldobler, 1976, 1984). In the formicine ant Camponotus hercu- 
leanus, mass flights of winged queens from the nest are also stimu- 
lated by secretions from the males’ mandibular glands (Holldobler 
& Maschwitz, 1965). Although queen honey bees secrete a sex 
pheromone (trans-9-keto-2-decenoic acid) from their mandibular 
glands (Butler, 1971), Polyergus breviceps is the first species of ant 
in which the queen’s sex-attractant pheromone has been traced to 
the mandibular glands. 
In an extensive review of flight activities, Kannowski (1963) 
reported a distinct tendency for ground mating in formicine ants. In 
Formica rufa and F. subnuda, for example, ground mating occurs 
after a nuptial flight. But in Formica subintegra and F. montana, 
winged females typically remain near their nest, with approaching 
males forming a “ground swarm” that terminates in mating. Alate 
queens of Polyergus breviceps have apparently adapted this behav- 
ioral pattern to a parasitic mode of life, especially their inability for 
independent colony foundation (Topoff et al., in press). By running 
and mating in slave-raid swarms, mated queens of P. breviceps 
arrive at colonies of Formica whose workers and queen are scat- 
tered across the substrate. Such disorganization in the target colony 
could facilitate the usurpation of Formica colonies by Polyergus 
queens. 
References 
Buschinger, A. 
1975. Sexual pheromones in ants. In: Pheromones and defensive secretions in 
social insects (Ch. Noirot, P. Howse, and G. Le Masne, eds.). Proc. Symp. 
IUSSI. Pp. 225-233. 
Buschinger, A. and Alloway, T. M. 
1979. Sexual behavior in the slave-making ant, Harpagoxenus canadensis M. 
R. Smith, and sexual pheromone experiments with H. canadensis, H. 
americanus (Emery), and H. sublaevis (Nylander) (Hymenoptera; Formicidae). 
Z. Tierpsychol. 49: 113-119. 
Butler, C. G. 
1971. The mating behavior of the honeybee ( Apis mellifera L.). J. Entomol. 46: 
1 - 11 . 
HOlldobler, B. 
1976. The behavioral ecology of mating in harvester ants (Hymenoptera: 
Formicidae: Polyergus). Behav. Ecol. Sociobiol. 1: 405-423. 
