NESTMATE RECOGNITION AND KIN RECOGNITION IN ANTS

Abstract  All ants (Hymenoptera, Formicidae) are highly eusocial insects that are characterized by reproductive division of labor with sterile castes (worker and soldier) helping fertile castes (queen and male) to reproduce.
Ant societies, like other complex animal societies, have developed a sophisticated communication system, in which recognition behaviors are frequently involved Recognition abilities allow individuals to orient and modulate their behaviors effectively and appropriately in response to the characteristics andlor signals expressed by other organisms. Among recognition behaviors, nestmate recognition and kin recognition mechanisms have attracted great attention of sociobiologists, ecologists, insect physiologists and biochemists since 1970's. This is parallel with the popularization of Hamilton's kin selection theory. The present paper aims at reviewing the current understanding on nestmate/kin recognition in ants. This review consists of three parts. The first part concerns the diversity of recognition behaviors and their ecological implications with emphasis on nestmatelkin recognition; in the second part, the current understandings on the mechanism of nestmatelkin recognition are outlined; and in the third part, we discuss the ontogenetic development of nestmate recognition behavior and naturally mixed colonies. The study of the integration mechanism of social parasite may provide heuristic clues to the understanding of kin/nestmate recognition system.     

Winged leaf-cutting ants on nuptial flights used as transport by Attacobius spiders for dispersal

Abstract.  1. Sexuals of a leaf-cutting ant, Atta bisphaerica Forel, left their nest for nuptial flights in October to December.
2. When leaving a nest, 53 of the 479 winged sexuals (or alates) observed (11.1%) carried up to three inquiline spiders of Attacobius luederwaldti .
3. Spiders exclusively selected winged sexuals, not workers, and preferred females, indicating their expectation of the stronger flight ability of females. Neither these sexuals nor workers that appeared out of the nest on flight days attempted to remove or attack spiders on the body of alates.
4. New qucens landing from their nuptial flight did not carry spiders, indicating that the spiders had left the ants in the sky to be dispersed by wind.
5. No spiders were found in more than 100 incipient nests, which were estimated to be 2–3 months old. This suggests that the spiders jumped off the alate during mid-flight and dispersed on the wind to inhabit larger nests.     

Tree killing by herbicide producing ants for the establishment of pureTococa occidentalis populations in the Peruvian Amazon

Populations ofTococa occidentalis (Melastomataceae) and the inhabiting ants (Myrmelachista sp.) were observed for more than eight months in the Peruvian Amazon (Sira mountains). They represent a complex coevolutionary system: the plants offer shelter (leaf domatia, hollow stems) and food (leaf glands), whereas the ants kill all surrounding plants, including large trees up to 10 m, by chemical weapons. Experiments with exposed plants revealed a highly specialized way to attack meristematic tissue and leaf nervature, which leads to a quick decay of the plant individuals. The clearing of the vegetation by the ants allows theTococa population to expand mostly by vegetative shoots to large monocultures (up to 30 m in diameter) free from any other plant species. Artificially introduced plant individuals, from differentT. occidentalis populations, are regarded as a foreign species by the ants.The succession of such aTococa-Myrmelachista system begins with one or a few founder plants on a light place in the midst of the vegetation.Myrmelachista soon inhabits their host plants which otherwise would not survive and begin to clear the place from all foreign plant species.Tococa expands quickly, forming circle shaped populations. Distantly situated canopy trees shade theTococa population after a number of years and cause their decay. The whole place appears contaminated for years and no other plant can establish itself. Some of the consequences of these open places are erosion and a severe influence on the regeneration of the forest.     

On the mimetic association between nymphs of Hyalymenus spp. (Hemiptera: Alydidae) and ants

Nymphs of Hyalymenus , unlike adults, have a highly differentiated ant-like morphology. Both H. tarsatus and H. limbativentris feed mainly on reproductive parts of composites and solanaceous plants, respectively. Mimetic nymphs were observed on plants, together with ants, both day and night; adult Hyalymenus , however, are predominantly nocturnal. Ant-resemblance in nymphs is achieved by several structural adaptations which, when coupled with the mimic's zig-zag locomotion and constantly agitated antennae, produces a striking visual deception. Experiments in captivity showed that mimetic nymphs, but not adult Hyalymenus , are somewhat protected against attacks from the praying mantid Oxyopsis media found on their host plant. Colour and size changes through different nymphal instars of Hyalymenus allow the immature bugs to resemble, during their development, differently sized and coloured ant models. Similar-looking ant species seem to act as Mullerian mimics toward insectivorous vertebrates and invertebrates that avoid ants. It is suggested that nymphs of Hyalymenus gain Batesian protection by resembling available ant models of different Mullerian complexes. Density-dependent selection is thought to be responsible for the observed differences in mimetic morph proportions between populations of mimics, as well as for the mimetic strategy itself employed by nymphs of Hyalymenus.     

Evolution of nesting strategies of ants: genetic evidence from different population types of Formica ants

Genetic population structure was studied in two types of populations in the ants Formica exsecta and F. pressilabris: populations consisting of single-nest colonies (monodomy) and populations consisting of multi-nest colonies (polydomy). These characteristics seem to be associated with the number of egg-laying females (gynes) in a nest, mating structure of the population, sex ratio and male size variation. The monodomous populations are characterized by single-gyne nests, the population sex ratio is either I:1 or female-biased, males are mainly large-sized, and there is slight inbreeding in the population. The polydomous populations have multi-gyne nests with gynes related to each other, sex ratio is strongly male-biased, most males are small-sized, and there is slight genetic microdifferentiation within the populations. Diploid males found in a polydomous F. pressilabris population suggest that the population is inbred and isolated. Habitat localization is presented as a plausible explanation for the evolution of the polygynous and polydomous population structure.     

“Wall-papering” and elaborate nest architecture in the ponerine antHarpegnathos saltator

Summary Excavation of 18 nests ofHarpegnathos saltator from southern India revealed an unusually complex architecture for a ponerine ant. The inhabited chambers are not deep in the ground. The uppermost chamber is protected by a thick vaulted roof, on the outside of which is an intervening space serving as isolation from the surrounding soil. In large colonies, the vaulted roof is extended into a shell which encloses several superimposed chambers. Little openings, which may be encircled by moulded flanges, occur in the upper region of the shell. The inside of the chambers is partly or completely lined with strips of empty cocoons. A refuse chamber is always found deeper than the inhabited chambers; live dipteran larvae (family Milichiidae) are typically present. These elaborate nests represent a large energetic investment, and we speculate therefore that nest emigration is unlikely in this species. Consequently, colony fission may never occur, unlike other ants where gamergates reproduce.     

Queen-worker conflict over sex ratio: A comparison of primary and secondary sex ratios in the Argentine ant,Iridomyrmex humilis

We compare the primary sex ratio (proportion of haploid eggs laid by queens) and the secondary sex ratio (proportion of male pupae produced) in the Argentine ant Iridomyrmex humilis with the aim of investigating whether workers control the secondary sex ratio by selectively eliminating male brood. The proportion of haploid eggs produced by queens was close to 0.5 in late winter, decreased to less than 0.3 in spring and summer, and increased again to a value close to 0.5 in fall. Laboratory experiments indicate that temperture is a proximate factor influencing the primary sex ratio with a higher proportion of haploid eggs being laid at colder temperatures. Production of queen pupae ceased in mid-June, about three weeks before that of male pupae. After this time only worker pupae were produced. During the period of production of sexuals, the proportion of male pupae ranged from 0.30 to 0.38. Outside this period no males were reared although haploid eggs were produced all the year round by queens. Workers thus exert a control on the secondary sex ratio by eliminating a proportion of the male brood during the period of sexual production and eliminating all the males during the remainder of the cycle. These data are consistent with workers preferring a more female-biased sex ratio than queens. The evolutionary significance of the production of male eggs by queens all the year round is as yet unclear. It may be a mechanism allowing queen replacement in the case of the death of the queens in the colony.     

Attraction of the sexes inFormica lugubris Zett (Hymenoptera: Formicidae)

Summary Sexuals ofFormica lugubris fly to mating places, where females attract males by using a sex pheromone. Females collected on the nest surface before departing on a mating flight are much less attractive than those collected on the mating place after the mating flight, suggesting that the mating flight triggers the release of the sex pheromone. Olfactory cues are essential for males to locate females while they patrol. Males probably use visual cues to locate females once they have alighted nearby them. Males are also attracted by aggregations of other males on the ground, probably because one or several females are likely to be close to male aggregations.