The multiple recruitment systems of the african weaver ant Oecophylla longinoda (Latreille) (Hymenoptera: Formicidae)

1. African weaver ants (Oecophylla longinoda) utilize no less than five recruitment systems to draw nestmates from the leaf nests to the remainder of the nest tree and to the foraging areas beyond: (a) recruitment to new food sources, mediated by odor trails produced from the rectal gland, a newly discovered exocrine organ, together with tactile stimuli presented during mouth-opening, antennation, and head-waggling; (b) recruitment to new terrain, entailing odor trails released from the rectal gland and tactile stimulation through antennation: (c) emigration to new sites; (d) short-range recruitment to territorial intruders, during which the terminal abdominal sternite is maximally exposed and dragged for short distances over the ground to release an attractant from the sternal gland, a second newly discovered structure; and (e) long-range recruitment to intruders, mediated by odor trails from the rectal gland and by antennation and intense body jerking. These systems exist in addition to the elaborate pheromone-mediated alarm communication previously described by Bradshaw et al. (1975). In aggregate, the alarm and recruitment systems of O. longinoda constitute the most complex of such repertories thus far discovered in ants. 2. Weaver ants recognize new terrain by means of both visual and olfactory cues, with the latter being the more effective. When major workers cannot cross gaps to the terrain by walking, they attempt to make the traverse by building bridges with their bodies. Individuals are attracted to the bridge site visually, but when the bridge is complete, they recruit nestmates to the new terrain with rectal-gland odor trails. 3. Workers mark newly acquired home range with randomly placed drops of fluid extruded from the rectal vesicle. They distinguish their own domain from that of alien conspecific colonies in part by means of the odor of the anal spots. When a section of terrain is found unmarked, the rate of anal-drop deposition is accelerated, even when adjacent areas are already heavily marked. 4. The anal substance is a true territorial pheromone: workers respond to alien spots initially with hostility and aversion, then by recruiting nestmates to the vicinity. In laboratory experiments, workers entering an arena simultaneously with workers from alien colonies always gained the initial advantage in the ensuing conflict if they had previously been allowed to mark the arena. When the arena was placed in a spatial position familiar to one colony but possessed a floor previously marked by the second colony, the second colony still won. To our knowledge these results represent the first demonstration of a true territorial pheromone in the social insects. 5. During foraging the Oecophylla workers move independently of one another and are distributed at random (Poisson distributed) or with slight temporary clumping of no more than two or three workers. Short-range recruitment of intruders causes the ants to shift to a more distinctly clumped pattern, involving as many as ten or more workers, at the same time that long-range recruitment brings more defenders into the vicinity. Together the two forms of response result in a more efficient capture of intruders that are too large to be immobilized by only one or two workers. 6. The complex recruitment and territorial behavior displayed by O. longinoda is considered to be part of the adaptation of these relatively large ants to a strongly arboreal existence. The similarity of four of the recruitment systems to each other (1 a, b, c, and e above) is interpreted as an example of signal economy in the evolution of social insect communication systems. The parallel evolution has been enhanced by the lack of any strong functional distinction between territorial defense and predation (see Discussion). 6. Signal ritualization appears to have occurred in at least two contexts: the modification of body thrusting during territorial battles into the jerking signal used in long-range recruitment of nestmates to enemies; and the adoption of anal excrement in the chemical marking of territories.