Myrmecina Nipponica Research Articles

Geographic patterns in colonial reproductive strategy in Myrmecina nipponica: Links between biogeography and a key polymorphism in ants.

The ability to express different phenotypes can help define species distributions by allowing access to, and exploitation of, new environments. Social insects employ two markedly different reproductive strategies with contrasting cost/benefit characteristics: independent colony foundation (ICF), which is associated with high dispersal range and high risk, and dependent colony foundation (DCF), characterized by low risk but low dispersal. The ant Myrmecina nipponica employs both of these strategies, with the frequency of each apparently varying between populations. We combine molecular data with data on reproductive strategy from different populations of this species throughout Japan to explore how this polymorphism is linked to environmental factors and whether this relationship can help explain the current and historical biogeography of this species. Reproductive strategy exhibited a strong geographic pattern, with ICF predominant in southern populations and DCF more common in northern and southern highland populations. Molecular analyses clearly divided populations into broad geographic regions, with the southern lowland populations basal to (southern highland (+ northern)) populations. Intra-population polymorphism in colony-founding strategy was widespread, and polymorphism was reconstructed as the likely ancestral state. The frequency of different strategies was linked with climate, with DCF more common in colder areas. A recent inferred origin to the northern lineage suggests that colonization of northern Japan was a rapid event coincident with warming at the end of the Last Glacial Maxima, likely facilitated by the cold-adaptive advantages of DCF. We discuss how such polymorphisms could help explain the biogeography of this and other social insects.

An agent-based model of nest-site selection in a mass-recruiting ant

Complex systems are modular entities which can collectively generate sophisticated emergent solutions through interactions based on simple, local rules. In this study, I use an agent-based model to elucidate how numerous individual-level components contribute to the collective decision process during house-hunting in a mass-recruiting ant species. Myrmecina nipponica combines the use of pheromone trails with a quorum decision rule in collective decisions among nest sites when searching for a new home. The model employed only individual-level rules but accurately emulated group-level properties observed in empirical studies. Simulations suggest that in this system i) both social and private information are necessary for effective decision making, ii) decision making was effective even with very low numbers of 'discriminating' individuals, iii) individual acceptance thresholds were more influential than quorum thresholds in tuning decisions to emphasise speed or accuracy, and iv) acceptance thresholds could also help tune decisions to suit environmental complexity. Similar findings in species using one-to-one recruitment suggest that some individual parameters, such as acceptance thresholds, may hold key functions in collective decision making regardless of the form of recruitment.

Individual and Group Personalities Characterise Consensus Decision‐Making in an Ant

AbstractNon‐human animals can exhibit idiosyncratic behaviour across individuals in much in the same way as humans. Animals with specific personalities may have advantages in some environments, and this idiosyncrasy may thus be of considerable ecological and evolutionary importance. In group‐living organisms, personality can occur at the level of the group as well as that of the individual. However, at present, we have very little understanding of the possible benefits of group‐level personality, and how this is linked with individual personality. In this study, I examine the influence of individual and group personality during the process of colony migration in the Japanese ant, Myrmecina nipponica. These ants use a consensus decision process to decide among alternatives when searching for a new home. Individuals contribute to this process by scouting for new nest sites, recruiting nestmates by laying pheromone trails, and carrying brood to the new site, although whether these roles are consistent among individuals and how roles are distributed within and between colonies remain unclear. Individual contributions to the nest‐site selection process were quantified over five repeated relocations in five colonies. Results demonstrate that contributions to the relocation effort were highly skewed within the colonies and that individuals were consistent in their contributions over repeated relocation events. Furthermore, the distribution of effort differed between colonies, indicating that intercolony differences in composition of behavioural types resulted in colony‐level personality. While these differences did not lead to any detectable difference in relocation performance between colonies in the simple experimental arrangement used, colony personality could influence decision outcomes in more complex environments.

Thelytokous Parthenogenesis in the Ant Myrmecina nipponica (Hymenoptera: Formicidae).

Myrmecina nipponica Wheeler is a terrestrial ant nesting chiefly in the soil in forest. It is a specialized predator of oribatid mites, but also scavenges on a broad spectrum of other arthropods. In the studied population at Cape Manazuru in central Japan, M. nipponica colonies are typically monogynous, and previous dissections of queens suggested that these individuals were not inseminated, thus suggesting these ants can reproduce via thelytokous parthenogenesis. To test for thelytokous parthenogenesis in M. nipponica the spermathecae of queens (dealate gynes) from worker-containing colonies were histologically examined in detail. All specimens examined (n=5) had no spermatozoa in the spermatheca. In addition, a total of four colony-founding queens were reared in isolation in the laboratory to test whether non-inseminated females were capable of egg laying and to test whether female offspring emerged from this brood. In all of four culture replicates, only new workers were produced from the eggs those queens had laid and male offspring was absent. After the breeding experiment, the queens' spermathecae were histologically examined and no sperm were detected in their spermathecae. These results reveal that M. nipponica queens of the Manazuru population are capable of producing female offspring thelytokously. Sexual reproduction by typical gynes and also by intermorphs has been known from other local populations of M. nipponica; therefore, this species shows geographical polymorphism in sexuality.

Ratio-dependent quantity discrimination in quorum sensing ants

To optimise behaviour, organisms require information on the quantity of various components of their environment, and the ability of animals to discriminate quantity has been a subject of considerable recent interest. This body of research hints at generalised mechanisms of quantity discrimination in vertebrates, but data on invertebrates are still relatively scarce. In this study, I present data on the quantification abilities of an invertebrate in a novel context: quorum sensing. Quorum sensing generates a behavioural response in group-living animals once a threshold number of individuals, a 'quorum', is detected performing some key action. This process forms the basis for consensus decision-making in many species and allows group-living organisms to decide among mutually exclusive alternatives without compromising group integrity. To determine when a quorum is achieved, individuals must assess the number of group members performing the key action. Social insects employ quorum decisions to decide among potential nest sites when searching for a new home. In the Japanese ant, Myrmecina nipponica, quorum thresholds increase with colony size, providing an opportunity to assess the accuracy of quantity discrimination at different stimulus magnitudes. In this study, I demonstrate that the variation in individual quorum thresholds around the mean increases with increasing colony size. This indicates that the quantity discrimination ability of ants decreases with stimulus magnitude, and thus exhibits ratio dependence in the manner of Weber's Law. This may have implications for the accuracy of consensus decision-making and other collective actions in a range of group-living organisms.

Synergy between pheromone trails and quorum thresholds underlies consensus decisions in the ant Myrmecina nipponica

Coordination of group actions in social organisms is often a self-organised process lacking central control. These collective behaviours are driven by mechanisms of positive feedback generated through information exchange. Understanding how different methods of communication generate positive feedback is an essential step in comprehending the functional mechanisms underlying complex systems. The Japanese small-colony ant, Myrmecina nipponica uses both pheromone trails and an apparent quorum response during consensus decisions over a new home. Both of these mechanisms have been shown to generate positive feedback and are effective means of selecting among mutually exclusive courses of action. In this study, I investigate how pheromone trails and quorum thresholds contribute to consensus decisions during house-hunting in this species through experimental manipulations of pheromone trails, colony size and environmental context. Results demonstrate that (1) providing colonies with pre-established pheromone trails increased the number of ants finding the new site and led to higher quorum thresholds and more rapid relocations, (2) experimentally halving colony size resulted in a proportional decrease in quorum thresholds and (3) colonies relocating long distances had higher quorums than those relocating short distances. Taken together, these data indicate that pheromone trails are important for recruitment and navigation during nest site selection, but that decision making is contingent on a quorum response. Such synergy between mechanisms of positive feedback may be a common means of optimising collective behaviours.

Consensus decision making in the ant Myrmecina nipponica: house-hunters combine pheromone trails with quorum responses

Consensus decisions enable group-living organisms to coordinate collective tasks without the need for complex cognitive abilities or centralized leadership. Systems of nonlinear positive feedback, such as quorum responses, are thought to be integral to consensus decision processes that seek to optimize decision speed and accuracy while maintaining group cohesion, requirements typical of the house-hunting process in social insects. Studies of nest site selection in honeybees, Apis mellifera, and Temnothorax ants have revealed strong similarities in the manner in which consensus decisions are achieved, and imply a central role for quorum responses, but treatments of other species are scarce. I investigated the process of consensus decision making during colony relocation in the small colony ant Myrmecina nipponica. Colonies of individually marked ants were forced to relocate in experimental laboratory arenas, and given a choice of one or two alternative new sites. This species differs from other well-studied systems in using pheromone trails to navigate between old and new sites, and does not use obvious means of recruitment such as tandem running or adult transport. A quorum threshold apparently governed the switch to brood transport, which marked the final phase of the relocation. Furthermore, there was a strong relationship between inferred quorum threshold and colony size. A demonstration of the use of a quorum rule in M. nipponica would underline the central role of quorum responses in consensus decision making, despite interspecific variation in colony size and differing modes of recruitment and navigation.

Ergatoid queen development in the ant Myrmecina nipponica : modular and heterochronic regulation of caste differentiation

Caste polyphenism in social insects provides us with excellent opportunities to examine the plasticity and robustness underlying developmental pathways. Several ant species have evolved unusual castes showing intermediate morphologies between alate queens and wingless workers. In some low-temperature habitats, the ant Myrmecina nipponica produces such intermediate reproductives (i.e. ergatoids), which can mate and store sperm but cannot fly. To gain insight into the developmental and evolutionary aspects associated with ergatoid production, we conducted morphological and histological examinations of the post-embryonic development of compound eyes, gonads and wings during the process of caste differentiation. In compound eyes, both the queen-worker and ergatoid-worker differences were already recognized at the third larval instar. In gonads, queen-worker differentiation began at the larval stage, and ergatoid-worker differentiation began between the prepupal and pupal stages. Wing development in ergatoids was generally similar to that in workers throughout post-embryonic development. Our results showed that the developmental rate and timing of differentiation in body parts differed among castes and among body parts. These differences suggest that the rearrangement of modular body parts by heterochronic developmental regulation is responsible for the origination of novel castes, which are considered to be adaptations to specific ecological niches.

Discussions of worker ants’ rule-based CHC dealing with changing environments

Contrary to popular belief, biologists discovered that worker ants are really not all hardworking. It has been found that in three separate 30-strong colonies of black Japanese ants ( Myrmecina nipponica), about 20% of worker ants are diligent, 60% are ordinary, and 20% are lazy. That is called 20:60:20 rule. Though they are lazy, biologists suggested that lazy worker ants could be contributing something to the colony that is yet to be determined. In our last research, we used CHC (cross generational elitist selection, heterogeneous recombination, and cataclysmic mutation) with the worker ants’ rule (WACHC) aiming at solving optimization problems in changing environments. CHC is a nontraditional genetic algorithm (GA) which combines a conservative selection strategy that always preserves the best individuals found so far with a radical (highly disruptive) recombination operator. In our last research, we verified that WACHC performs better than CHC in only one case of fully changing environment. In this paper, we further discuss our proposed WACHC dealing with changing environment problems with varying degree of difficulty, compare our proposal with hypermutation GA which is also proposed for dealing with changing environment problems, and discuss the difference between our proposal and ant colony optimization algorithms.

Effects of worker manipulation on the sex ratio of a Japanese ant species,Myrmecina nipponica

In order to test the effects of colony size and nutritional condition on the survivorship and sex ratio of ants,Myrmecina nipponicacolonies were housed in a laboratory in colony sizes of 10 or 30 individuals and fed either daily or weekly. Under all conditions, most of the larvae successfully grew into adults, which suggests that survivorship was not significantly affected by either colony size or nutritional condition. However, the number of new queens was significantly higher in colonies that were fed daily. These results indicate that workers do not control the proportion of diploid and haploid broods by eliminating some larvae and that nutritional condition exerts a significant effect on sex ratio.

Morphology and Developmental Plasticity of Reproductive Females in <I>Myrmecina nipponica</I> (Hymenoptera: Formicidae)

Abstract We examined the morphology and developmental plasticity of alate/dealate queens and permanently wingless reproductive females (hereafter referred to as “intermorphic queens”) in Myrmecina nipponica Wheeler. Alate/dealate queens and intermorphic queens had nearly equal reproductive capacities. The number of ovarioles was almost identical between virgin and inseminated alate/dealate queens, but inseminated intermorphic queens had significantly more ovarioles than virgin females. To determine if intermorphic queens in M. nipponica are ‘ergatoid queens’ or ‘intercaste,’ we compared seven external traits of each female caste. The external traits of intermorphic queens, in particular pronotal width and Weber’s length of the mesosoma, were more similar to workers than to alate/dealate queens. Data on ovarioles suggest that the intermorphic queens have a reproductive ability similar to alate/dealate queens. The absence of a reproductive female in a colony and low temperature stimulated the production of ...

Mating frequency, genetic structure, and sex ratio in the intermorphic female producing ant speciesMyrmecina nipponica

Summary1.Myrmecina nipponicahas two types of colonies: a queen colony type, in which the reproductive females are queens and new colonies are made by independent founding, and an intermorphic female colony type, in which reproductive females belong to a wingless intermediate morphology between queen and worker, and where colonies multiply through colonial budding.2. The mating frequencies of reproductive females in both types indicate monoandry. The relatedness among nestmates in both types was almost 0.75, however relatedness between mother and daughter in intermorphic female colonies was slightly higher than that of queen colonies.3. The sex ratio (corrected investment female ratio) was 0.70 at the population level, suggesting that the sex ratio is controlled by workers in this species, however the ratio differed greatly between the two types of colonies. Queen colonies (n = 37) had a female‐biased sex ratio of 0.77 while intermorphic female colonies (n = 33) had a ratio of 0.56.4. Each reproductive intermorphic female was accompanied by an average of 2.9 workers (including virgin intermorphic females) in the colonial budding, and when the investment to those workers was added to the female investment, the sex ratio reached 0.81.5. The frequency distribution of sex ratio was bimodal, with many colonies producing exclusively males or females, however mean estimated relatedness within colonies was almost 0.75. These data are inconsistent with the genetic variation hypothesis, which is one of the predominant hypotheses accounting for the between‐colony variation in sex ratio.

Sex–ratio regulation: the economics of fratricide in ants

In many insect societies, workers can manipulate the reproductive output of their colony by killing kin of lesser value to them. For instance, workers of the mound-building Formica exsecta eliminate male brood in colonies headed by a single-mated queen. By combining an inclusive fitness model and empirical data, we investigated the selective causes underlying these fratricides. Our model examines until which threshold stage in male brood development do the workers benefit from eliminating males to rear extra females instead. We then determined the minimal developmental stage reached by male larvae before elimination in F. exsecta field colonies. Surprisingly, many male larvae were kept until they were close to pupation, and only then eliminated. According to our model, part of the eliminated males were so large that workers would not benefit from replacing them with new females. Moreover, males were eliminated late in the season, so that new females could no longer be initiated, because matings take place synchronously during a short period. Together, these results indicate that workers did not replace male brood with new females, but rather reduced total brood size during late larval development. Male destruction was probably triggered by resource limitation, and the timing of brood elimination suggests that males may have been fed to females when these start to grow exponentially during the final larval stage. Hence, the evolution of fratricides in ants is best explained by a combination of ecological, demographic and genetic parameters.