Fertility Signaling and Partitioning of Reproduction in the Ant Neoponera apicalis.

All individuals in social insect colonies benefit from being informed about the presence and fertility state of reproducers. This allows the established reproductive individuals to maintain their reproductive monopoly without the need for physical control, and the non-reproductive individuals to make appropriate reproductive choices. Here, we studied whether fertility signaling is responsible for the partitioning of reproduction in the ant Neoponera apicalis. This species forms small colonies from one single-mated queen, with workers establishing reproductive hierarchies when hopelessly queenless. Previous studies identified putative fertility signals, particularly the hydrocarbon 13-methylpentacosane (13-MeC25), and have shown that precise status discrimination based on these signals could be involved in the regulation of reproductive activities. Here, we extend these findings and reveal that all individuals, be they queens or workers, differ in their cuticular hydrocarbon profile according to fertility state. Proportions of 13-MeC25 were a strong predictor of an individual's ovarian activity, and could, thus, advertise the established reproducer(s) in both queenright and queenless conditions. Furthermore, this compound might play a key role in the establishment of the reproductive hierarchy, since workers with low fertility at the onset of hierarchy formation already have relatively high amounts of 13-MeC25. Dyadic encounters showed that individuals with experimentally increased amounts of 13-MeC25 triggered less agonistic interactions from top rankers, in accord with them "advertising" higher status. Thus, these bioassays supported the use of 13-MeC25 by competing ants. This simple recognition system potentially allows permanent regulation of partitioning of reproduction in this species.

Chemical signature and reproductive status in the facultatively polygynous ant Pachycondyla verenae.

In insects, cuticular hydrocarbons (CHCs) generally are used as cues and signals for within colony processes, such as signaling reproductive status, and between colony processes, such as colony membership. We examined CHC profiles of the facultatively polygynous ant Pachycondyla verenae in order to identify chemical signals of reproductive queens within colonies containing many gynes. Colonies of P. verenae, belonging to two different members of a complex of morphospecies, were collected from three geographic localities within South America. We also tested whether CHC profiles differed between geographic localities and morphospecies. We found three alkenes, two isomers of pentacosene and heptacosene, which were more abundant in CHC profiles of reproductive queens of this morphospecies complex. When we tested whether these differences were consistent across geographic localities, we found the abundance of these alkenes differed according to morphospecies, with the isomers of pentacosene being more abundant in queens from morph one, and heptacosene being more abundant in queens from morph two. Our study has given further insight into the mechanisms behind maintenance of reproductive dominance, and has demonstrated that chemical signatures associated with reproductive status in Pachycondyla verenae are not conserved within this species complex.

Enriched topological learning for cluster detection and visualization.

The exponential growth of data generates terabytes of very large databases. The growing number of data dimensions and data objects presents tremendous challenges for effective data analysis and data exploration methods and tools. Thus, it becomes crucial to have methods able to construct a condensed description of the properties and structure of data, as well as visualization tools capable of representing the data structure from these condensed descriptions. The purpose of our work described in this paper is to develop a method of describing data from enriched and segmented prototypes using a topological clustering algorithm. We then introduce a visualization tool that can enhance the structure within and between groups in data. We show, using some artificial and real databases, the relevance of the proposed approach.

The role of microgynes in the reproductive strategy of the neotropical ant Ectatomma ruidum.

Miniaturized queens, microgynes, are regarded as an alternative reproductive strategy sparsely present through the ant world. The described roles of miniaturized queens include alternative short-distance dispersal morphs, an adaptation to polygyny and inquiline parasites. Some of these inquiline parasite microgynes have been described as a separate species from their host. In the poneromorph group, miniaturized queens are only reported in two Mexican populations of two Ectatomminae: Ectatomma tuberculatum, in which small queens represent an inquiline species (Ectatomma parasiticum) and Ectatomma ruidum. E. ruidum presents apparently facultative polygyny with microgynes. We used mitochondrial DNA markers and newly developed microsatellite loci to investigate the status as well as the role of microgynes in E. ruidum. We confirmed that microgynes and macrogynes are from the same species. This species is almost exclusively monogynous and monoandrous, supernumerary dealate queens of both types being actually daughters of the mother queen. An apparently polygynous nest was more often headed by a macrogyne than a microgyne. We didn't find any inbreeding or isolation by distance in the studied population, indicating that new gynes are inseminated by unrelated males and can establish a new nest far from their natal nest. However, re-adoption of daughter queens seems to be the rule and rate of microgyny appears to be linked to nest density and environmental factors.

Permanent genetic resources added to Molecular Ecology Resources Database 1 October 2010-30 November 2010.

This article documents the addition of 277 microsatellite marker loci to the Molecular Ecology Resources Database. Loci were developed for the following species: Ascochyta rabiei, Cambarellus chapalanus, Chionodraco hamatus, Coptis omeiensis, Cynoscion nebulosus, Daphnia magna, Gerbillus nigeriae, Isurus oxyrinchus, Lates calcarifer, Metacarcinus magister, Oplegnathus fasciatus, Pachycondyla verenae, Phaethon lepturus, Pimelodus grosskopfii, Rotylenchulus reniformis, Scomberomorus niphonius, Sepia esculenta, Terapon jarbua, Teratosphaeria cryptica and Thunnus obesus. These loci were cross-tested on the following species: Austropotamobius italicus, Cambarellus montezumae, Cambarellus puer, Cambarellus shufeldtii, Cambarellus texanus, Chionodraco myersi, Chionodraco rastrospinosus, Coptis chinensis, Coptis chinensis var. brevisepala, Coptis deltoidea, Coptis teeta, Orconectes virilis, Pacifastacus leniusculus, Pimelodus bochii, Procambarus clarkii, Pseudopimelodus bufonius, Rhamdia quelen, Sepia andreana, Sepiella maindroni, Thunnus alalunga, Thunnus albacares, Thunnus maccoyii, Thunnus orientalis, Thunnus thynnus and Thunnus tonggol.

Worldwide invasion by the little fire ant: routes of introduction and eco-evolutionary pathways.

Biological invasions are generally thought to occur after human aided migration to a new range. However, human activities prior to migration may also play a role. We studied here the evolutionary genetics of introduced populations of the invasive ant Wasmannia auropunctata at a worldwide scale. Using microsatellite markers, we reconstructed the main routes of introduction of the species. We found three main routes of introduction, each of them strongly associated to human history and trading routes. We also demonstrate the overwhelming occurrence of male and female clonality in introduced populations of W. auropunctata, and suggest that this particular reproduction system is under selection in human-modified habitats. Together with previous researches focused on native populations, our results suggest that invasive clonal populations may have evolved within human modified habitats in the native range, and spread further from there. The evolutionarily most parsimonious scenario for the emergence of invasive populations of the little fire ant might thus be a two-step process. The W. auropunctata case illustrates the central role of humans in biological change, not only due to changes in migration patterns, but also in selective pressures over species.

Stridulations reveal cryptic speciation in neotropical sympatric ants.

The taxonomic challenge posed by cryptic species underlines the importance of using multiple criteria in species delimitation. In the current paper we tested the use of acoustic analysis as a tool to assess the real diversity in a cryptic species complex of Neotropical ants. In order to understand the potential of acoustics and to improve consistency in the conclusions by comparing different approaches, phylogenetic relationships of all the morphs considered were assessed by the analysis of a fragment of the mitochondrial DNA cytochrome b. We observed that each of the cryptic morph studied presents a morphologically distinct stridulatory organ and that all sympatric morphs produce distinctive stridulations. This is the first evidence of such a degree of specialization in the acoustic organ and signals in ants, which suggests that stridulations may be among the cues used by these ants during inter-specific interactions. Mitochondrial DNA variation corroborated the acoustic differences observed, confirming acoustics as a helpful tool to determine cryptic species in this group of ants, and possibly in stridulating ants in general. Congruent morphological, acoustic and genetic results constitute sufficient evidence to propose each morph studied here as a valid new species, suggesting that P. apicalis is a complex of at least 6 to 9 species, even if they present different levels of divergence. Finally, our results highlight that ant stridulations may be much more informative than hitherto thought, as much for ant communication as for integrative taxonomists.

Effect of social and environmental factors on ant aggregation: a general response?

In social species, the interactions that occur between individuals and also between individuals and their environment can lead to various spatial distributions. Our goal was to understand how both factors (social and environmental) can affect such spatial distribution of individuals. Aggregation experiments were conducted on the ant Crematogaster scutellaris: the level of inter-attraction between individuals was characterized and the influence of an environmental factor (light intensity) was studied. Two ethological castes (brood-tenders and foragers) were tested, showing a strong and similar aggregation pattern-although faster for foragers. This difference could be explained by a higher degree of inter-attraction for brood-tenders, resulting in longer interaction time and therefore a slower aggregation rate. We also showed the influence of light conditions by carrying out experiments in total darkness and under red light. C. scutellaris appeared highly sensitive to the red light, aggregating just underneath it. Nevertheless, the level of aggregation is not affected by the luminosity condition. These results are discussed in relation to the ecology of the species, and are compared with results obtained in another ant species, Lasius niger, which exhibits a similar response to aggregation. We propose, therefore, a general link between the level/pattern of aggregation and polyethism in ants.

Postmating changes in cuticular chemistry and visual appearance in Ectatomma tuberculatum queens (Formicidae: Ectatomminae).

In the ectatommine ant Ectatomma tuberculatum, the visual appearance of queens changes after mating and ovarian development in that their cuticle turns from shiny to matte. In this study, we have shown that this change seems to be caused by 15-fold accumulation of hydrocarbons, in particular heptacosane that covers the multiple grooves present on the cuticular surface creating a wax coat in mated fully fertile queens. Analyses of the scrapped wax revealed that it is composed largely of heptacosane. Peak-by-peak comparison of the cuticular hydrocarbon (CHC) composition of mated, virgin with developed ovaries and virgin with nondeveloped ovaries revealed significant differences between the queen groups. Although the total amount of the CHC of virgin queens with developed ovaries was not higher than virgin queens that did not have developed ovaries, the composition showed a shift toward the mated queen. While it is possible that the large accumulation of hydrocarbons may give extra physical and chemical protection to queens, we propose that the switch in the relative abundance of heptacosane and nonacosane and perhaps of other components is indicative of being a mating and fertility cue. This is the first report in social insects where external chemical changes are accompanied by changes in visual appearance.

How an ant manages to display individual and colonial signals by using the same channel.

Cuticular hydrocarbons are used by some ants to discriminate nestmates from nonnestmates. Every member of the colony bears the same pattern because they are continuously exchanged among nestmates. The postpharyngeal gland (PPG) stores the blend of hydrocarbons and is involved in the distribution of this common mixture. However, some individuals might display individual information on the cuticle (such as a chemical signal of fertility) that must not be mixed within the common pool. We investigated how this paradox is solved in the ant Pachycondyla goeldii by analyzing the nature and localization of colonial and fertility signals. Workers in a queenless condition showed a dominance hierarchy that was correlated with ovarian development. Hydrocarbons from the cuticle and the PPG analyzed by gas chromatography (GC) and identified by GC-mass spectrometry showed a clear discrimination among colonies, supporting the involvement of the PPG in the colonial identity signal. We identified and selected 11 cuticular hydrocarbons that permitted us to discriminate ovarian development classes and that might function as a fertility signal. They allowed clear colony discrimination as well, which suggests that the two signals (the individual signal of fertility and the common signal of colony identity) can be conveyed by the same compounds. However, the hydrocarbons in the PPG did not discriminate among ovarian developmental classes, suggesting that the portion of variation in the cuticular hydrocarbons constituting the fertility signal is superimposed on the signal of colony identity.

Queen movement during colony emigration in the facultatively polygynous ant Pachycondyla obscuricornis.

In ants, nest relocations are frequent but nevertheless perilous, especially for the reproductive caste. During emigrations, queens are exposed to predation and face the risk of becoming lost. Therefore the optimal strategy should be to move the queen(s) swiftly to a better location, while maintaining maximum worker protection at all times in the new and old nests. The timing of that event is a crucial strategic issue for the colony and may depend on queen number. In monogynous colonies, the queen is vital for colony survival, whereas in polygynous colonies a queen is less essential, if not dispensable. We tested the null hypothesis that queen movement occurs at random within the sequence of emigration events in both monogynous and polygynous colonies of the ponerine ant Pachycondyla obscuricornis. Our study, based on 16 monogynous and 16 polygynous colony emigrations, demonstrates for the first time that regardless of the number of queens per colony, the emigration serial number of a queen occurs in the middle of all emigration events and adult ant emigration events, but not during brood transport events. It therefore appears that the number of workers in both nests plays an essential role in the timing of queen movement. Our results correspond to a robust colony-level strategy since queen emigration is related neither to colony size nor to queen number. Such an optimal strategy is characteristic of ant societies working as highly integrated units and represents a new instance of group-level adaptive behaviors in social insect colonies.

The influence of red light on the aggregation of two castes of the ant, Lasius niger.

Insects are generally described as essentially blind to red wavelengths but sensitive to ultraviolet. Therefore, ants are generally reared in nests where "obscurity" is obtained by the use of a red filter. Yet, no study has been done in order to confirm this idea and to verify whether ants behave differently with or without red light. We performed aggregation experiments with the ant Lasius niger under red light and in total darkness to control the sensitivity of workers. Aggregation is a basic behavior associated with the origin of sociality and with the spatial organization of the colony. We demonstrate that L. niger is sensitive to red wavelengths. Moreover, we show that workers behave differently depending on their ethological caste: foragers aggregate well in total darkness but show low assembly under red light, whereas brood-tenders aggregate well in both conditions. For the first time, a link between vision, social organization and spatial patterns is revealed. The results are discussed relative to their adaptive value and relative to the physiology of the workers. Hypotheses are formulated concerning the acquisition of this change of behavior between castes.