We investigated the effect of injections of four biogenic amines (serotonin, dopamine, octopamine and tyramine) on behavior patterns displayed by workers of the red wood ant Formica polyctena during dyadic confrontations with four types of opponents: a nestmate, an alien conspecific, an allospecific ant (Formica fusca), and a potential prey, a nymph of the house cricket (Acheta domesticus). Significant effects of biogenic amine administration were observed almost exclusively in the case of confrontations with allospecific opponents. Serotonin treatment exerted stimulatory effects on behavior patterns involving physical aggression (biting accompanied by gaster flexing, dragging and formic acid spraying), but these effects were relatively weak and/or documented by indirect evidence. Dopamine administration exerted a stimulatory effect on open-mandible threats directed by F. polyctena to F. fusca and to cricket nymphs, and on biting behavior directed to cricket nymphs. Surprisingly, octopamine treatment did not exert significant effects on aggressive behavior of the tested ants. Tyramine administration exerted a suppressing effect on threatening behavior directed to F. fusca, but led to shortening of the latencies to the first open-mandible threat during the tests with cricket nymphs. Biogenic amine administration also influenced non-aggressive behavior of the tested ants. Our findings confirmed the role of serotonin and dopamine in the mediation of ant aggressive behavior and documented for the first time significant effects of tyramine treatment on ant aggressive behavior. We also demonstrated that not only specific patterns of ant aggressive behavior, but also behavioral effects of biogenic amine treatments are as a rule strongly context-dependent.

Wood ant colonies that appear to consist of individuals representing different species are described in several previous papers. The present study is the first to elucidate the genetic basis of the spectacular morphological variability observed within such colonies. Two seemingly mixed colonies (FM-1 and FM-2) from southern Finland were investigated. On the basis of the morphology of their workers these colonies were comprised of individuals with phenotypes typical of Formica rufa L., F. polyctena F?rst., and F. aquilonia Yarr. The sequence of an mtDNA fragment (5' end of the cytochrome b gene) was used to examine the phylogenetic relationships among haplotypes of workers from homogeneous colonies of different wood ant species and the two supposedly mixed colonies, and to sort the individuals within the colonies into matrilines. Six microsatellite loci were used to analyse the genetic differentiation between colonies and among workers within colonies, and to detect putative hybrids. The results show that, independently of their phenotype, workers from the "mixed" colonies were genetically more similar to other individuals in their colony than to those in the homogeneous F. rufa, F. polyctena or F. aquilonia colonies. However, while colony FM-1 consisted of offspring of the same queen or more likely several maternally related queens, colony FM-2 consisted of the offspring of at least four unrelated queens. The data suggest hybridisation between F. polyctena ? F. aquilonia and F. polyctena ? F. rufa (and possibly subsequent mating between these hybrids) as the most probable mechanism leading to the existence of these two colonies, which implies that the hybrids are fertile. This study shows that colonies of wood ant hybrids can arise spontaneously and persist under natural conditions. The results also revealed that even some morphologically homogeneous colonies are genetically heterogeneous. In the case of closely related, morphologically similar species that interbreed, morphology can be a bad predictor of genetic differences between individuals.

Social insect workers usually start adult life from intranidal tasks and then switch to extranidal activities, but this process may be reversed: foragers may switch again to intranidal brood care. The transition forager – reverted nurse is known as the behavioral reversion. Ant foragers are known to avoid illuminated zones less strongly than intranidal workers, but illumination responses of reverted nurses were so far never investigated. We compared dark–light choice behavior of three classes of workers of the red wood ant Formica polyctena: nurses, foragers and reverted nurses. Sets of ten ants belonging to the same class were tested in “double nests” made of two interconnected test tubes, one kept in darkness and the other exposed to light. The number of ants present in the illuminated zone of each nest (n_i) was recorded on 10 sample points at 30 min intervals. The values of n_i were lower in nurses than in foragers and reverted nurses and decreased as a function of time in all three groups. Nurses differed from foragers with respect to the dynamics of dark–light choice behavior, but reverted nurses did not differ in that respect either from nurses, or from foragers. Reverted nurses and foragers did not differ significantly from each other with respect to the overall level of avoidance of illuminated zone, nor with respect to the dynamics of dark–light choice behavior. This implies that behavioral reversion is not accompanied by the return of illumination responses of workers of F. polyctena to the state characteristic for nurses.