The role of workers in transferring queen substances and the differences between worker castes in the leaf-cutting ant, Acromyrmex subterraneus brunneus (Hymenoptera: Formicidae)

The present study focused on the relationship between the queen and workers in Acromyrmex subterraneus brunneus colonies mediated by the possible transfer of substances from the fertile to the sterile caste. The queens were submitted to different situations of physical limitation, i.e., they were kept isolated in cages with holes that only permitted the entry of workers but left the queen isolated. A tracer (water-soluble dye) was applied to the pronotum and gaster of the queen and its dispersal among workers was analyzed. The results demonstrated that the minor sub caste (0.7-0.9 mm) passed on the dye through allogrooming and self-grooming, or transferred the dye through trophallaxis to the major sub caste (1.2-2.0 mm) when they were not in direct contact with the queen. These findings indicate the communication and probable transfer of substances from the queen to the workers, as well as a substance transfer between workers.

Ghost-ant: Post-embryonic development of the worker caste of Tapinoma melanocephalum (hymenoptera: Formicidae)

This study examines some aspects of the basic biology of the worldwide distributed ant Tapinoma melanocepbalum Fabricius. The number of larval instars and the growth ratio between each instar are given. We used colonies containing only queens and workers, and later removed these queens in order to estimate the production of eggs and the duration of immature development of the worker caste. Measurements of larvae cephalic capsule widths revealed that workers of the ghost-ant have four larval instars from egg-hatching to adult. The mean growth rate for the species is 1.38, in accordance with Dyar's rule. The highest egg production was 5.3±2.2 eggs/day/queen and the analysis of these brood suggested the presence of two kinds of eggs inside the colony. The development of workers from egg to adult lasted 16-52 days with the embryonic development longer than larval, prepupal or pupal stages. Despite the slow egg-laying by queens, our findings also showed that colonies of T. melanocephalum grow faster than colonies of other tramp ants.

Immature development in the tramp species ant Paratrechina longicornis (Hymenoptera, formicidae)

The genus Paratrechina (Motschulsky) is composed of about 147 cosmopolitan ant species and subspecies, of which some were accidentally spread by commerce around the world, infesting houses and hospitals. In Brazil, two species are of remarkable economic importance: Paratrechina fulva (Mayr) and Paratrechina longicornis (Latreille). The present paper aims to report the following aspects on the biology of P. longicornis: number of larval instars, growth rates, duration and viability of the developmental stages and some observations on brood-care behaviour. Artificial colonies were kept under controlled temperature (25±2°C) and relative humidity (60±10%) conditions. To access the number of larval instars, we analyzed 1531 larvae fixed in Dietrich by measuring their maximum head capsule widths. We analysed ten artificial colonies daily to measure the developmental time and viability of immatures. We found P. longicornis presented three larval instars and a mean growth ratio of 1.2988. Egg stage was found to last 16.1 ±0.1 days, with a viability of 24.6%; larval stages were found to last 18.3±0.1 days, with a viability of 33.3% and the pupal stage was found to last 12.3±0.1 days, with a viability of 65.3%.

Nest defense behavior against the attack of ants in colonies of pre-emergent Mischocyttarus cerberus (hymenoptera, vespidae)

The study was carried out at the UNESP Rio Claro campus (SP), where biotests consisting of simulated ant attacks were performed in colonies of Mischocyttarus cerberus. The behaviors of the wasps were recorded with a camcorder, for further analysis. This analysis was done using the Mann-Whitney U test and the Principal Component Analysis. In the pre-emergence development stage, colonies with a single foundress defend the nest only after the first larvae appear. When there are only eggs in the nest, the wasp abandons the nest. Before leaving, the wasp rubs its gaster against the nest, releasing the ant repellent secretion. When the nest contains larvae or larvae and pupae, the foundress defends the colony, vibrating its wings, pumping her abdomen and biting the ant.

Thelytokous parthenogenesis in the fungus-gardening ant Mycocepurus smithii (Hymenoptera: Formicidae)

The general prevalence of sexual reproduction over asexual reproduction among organisms testifies to the evolutionary benefits of recombination, such as accelerated adaptation to changing environments and elimination of deleterious mutations. Documented instances of asexual reproduction in groups otherwise dominated by sexual reproduction challenge evolutionary biologists to understand the special circumstances that might confer an advantage to asexual reproductive strategies. Here we report one such instance of asexual reproduction in the ants. We present evidence for obligate thelytoky in the asexual fungus-gardening ant, Mycocepurus smithii, in which queens produce female offspring from unfertilized eggs, workers are sterile, and males appear to be completely absent. Obligate thelytoky is implicated by reproductive physiology of queens, lack of males, absence of mating behavior, and natural history observations. An obligate thelytoky hypothesis is further supported by the absence of evidence indicating sexual reproduction or genetic recombination across the species' extensive distribution range (Mexico-Argentina). Potential conflicting evidence for sexual reproduction in this species derives from three Mycocepurus males reported in the literature, previously regarded as possible males of M. smithii. However, we show here that these specimens represent males of the congeneric species M. obsoletus, and not males of M. smithii. Mycocepurus smithii is unique among ants and among eusocial Hymenoptera, in that males seem to be completely absent and only queens (and not workers) produce diploid offspring via thelytoky. Because colonies consisting only of females can be propagated consecutively in the laboratory, M. smithii could be an adequate study organism a) to test hypotheses of the population-genetic advantages and disadvantages of asexual reproduction in a social organism and b) inform kin conflict theory. For a Portuguese translation of the abstract, please see Abstract S1. © 2009 Rabeling et al.

Polydomy in the ant Ectatomma opaciventre

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Protein content of leaf-cutting ant queens before the nuptial flight and during the post-claustral phase

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Development of twenty-one polymorphic microsatellite markers for the fungus-growing ant, Mycocepurus goeldii (Formicidae: Attini), using Illumina paired-end genomic sequencing

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

A Social Parasite Evolved Reproductive Isolation from Its Fungus-Growing Ant Host in Sympatry

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Toxic effects of Canavalia ensiformis L. (Leguminosae) on laboratory colonies of Atta sexdens L. (Hym., Formicidae)

Laboratory colonies of the leaf-cutting ants Atta sexdens that were fed daily with leaves of Canavalia ensiformis showed a high ant mortality, and a significant decrease in fungal garden volume, with complete depletion of nests after 11 weeks of treatment.

A taxonomic revision of the Neotropical myrmicine ant genus Lachnomyrmex wheeler (Hymenoptera : Formicidae)

The Neotropical ant genus Lachnomyrmex Wheeler, 1910 is revised for the first time. A revised generic description is provided for workers and gynes; males remain unknown. Morphological patterns combined with geographical data led to the recognition of 16 species, of which 10 are here described as new: L. amazonicus sp. n., L. fernandezi sp. n., L. grandis Fernandez & Baena, L. haskinsi Smith, L. lattkei sp. n., L. longinodus Fernandez & Baena, L. longinoi sp. n., L. mackayi sp. n., L. nordestinus sp. n., L. pilosus Weber, L. laticeps sp. n., L. platynodus sp. n., L. plaumanni Borgmeier, L. regularis sp. n., L. scrobiculatus Wheeler, and L. victori sp. n. The gynes of L. grandis, L. haskinsi, and L. pilosus are described for the first time. Illustrated identification keys for workers, distribution maps, and high resolution illustrations are supplied for all species. Some Lachnomyrmex species are relatively common in the leaf-litter of submontane wet forests in Central and South America; others remain known by very few individuals. The relatively small and apparently monogynic Lachnomyrmex colonies inhabit small nests in the ground, among rotten leaves and inside fallen logs. Workers generally forage alone on the ground or within the leaf litter, but have been recorded also on tree trunks, and apparently do not recruit nestmates.

Sodium-specific foraging by leafcutter ant workers (Atta cephalotes, Hymenoptera: Formicidae)

1. Sodium is often a limiting nutrient for terrestrial animals, and may be especially sought by herbivores. Leafcutter ants are dominant herbivores in the Neotropics, and leafcutter foraging may be affected by nutritional demands of the colony and/or the demands of their symbiotic fungal mutualists. We hypothesized that leafcutter colonies are sodium limited, and that leafcutter ants will therefore forage specifically for sodium. 2. Previous studies demonstrated that leafcutter Atta cephalotes Linnaeus workers preferentially cut and remove paper baits treated with NaCl relative to water control baits. Atta cephalotes colonies in this study were presented with baits offering NaCl, Na2SO4, and KCl to test whether leafcutters forage specifically for sodium. Sucrose and water were used as positive and negative controls, respectively. 3. Atta foragers removed significantly more of the baits treated with NaCl and Na2SO4 than the KCl treatment, which did not differ from water. The NaCl and Na2SO4 treatments were collected at similar rates. We conclude A. cephalotes forage specifically for sodium rather than for anions (chloride) or solutes in general. This study supports the hypothesis that leafcutter ants are limited by, and preferentially forage for, sodium.

Aggression and its function as a hub mediating host defense against the slavemaking ant Protomognathus americanus

In meiner Dissertation beschäftigte ich mich mit unterschiedlichen Verteidungsstrategien, derenrnEffektivität und Evolution, der Ameisenart Temnothorax longispinosus (“Sklaven”), gegenüberrneinem sozialen Parasiten - der nahverwandten, sklavenhaltenden Art Protomognathusrnamericanus (“Sklavenhalter”). Wir entdeckten eine neue Kategorie der Verteidigungsstrategie,rnwelche es dem Wirten ermöglicht, flexibel auf die nicht vorhersagbaren Angriffe des Parasitenrnzu reagieren. Darüber hinaus erforschten wir, wie die Wirte ihre kollektive Verteidigung an einernVielzahl unterschiedlicher Angreifer anpassen können. Wir konnten feststellen, dass Wirte in derrnLage sind ihre kollektive Verteidigung dem Grad der Bedrohung anzupassen. Dies weist daraufrnhin, dass Selektion die Verteidigung gegen unterschiedliche Typen von Angreifern voneinanderrnunabhängig beeinflussen könnte. In einer dritten Studie belegten wir experimentell, dass diernParasiten die Evolution der Kolonieaggressivität der Wirtsart direkt beeinflussen. Die letztenrnbeiden Publikationen beschäftigten sich mit Sklavenrebellion, einer rätselhaftenrnVerteidigungsstrategie, da noch unklar ist, wie eine Eigenschaft von nicht reproduzierendenrnIndividuen vererbt werden kann. In einer Metaanalyse konnten wir die weite Verbreitung undrnhohe Variabilität dieser Eigenschaft dokumentieren, und fanden Hinweise, dassrnVerwandtenselektion eine mögliche Erklärung für die Evolution dieses Merkmals darstellenrnkönnte.

Alternative reproductive tactics, sexual selection and the effects of inbreeding in the ant Hypoponera opacior

Inbreeding can lead to a fitness reduction due to the unmasking of deleterious recessive alleles and the loss of heterosis. Therefore, most sexually reproducing organisms avoid inbreeding, often by disperal. Besides the avoidance of inbreeding, dispersal lowers intraspecific competition on a local scale and leads to a spreading of genotypes into new habitats. In social insects, winged reproductives disperse and mate during nuptial flights. Therafter, queens independently found a new colony. However, some species also produce wingless sexuals as an alternative reproductive tactic. Wingless sexuals mate within or close to their colony and queens either stay in the nest or they found a new colony by budding. During this dependent colony foundation, wingless queens are accompanied by a fraction of nestmate workers. The production of wingless reproductives therefore circumvents the risks associated with dispersal and independent colony foundation. However, the absence of dispersal can lead to inbreeding and local competition.rnIn my PhD-project, I investigated the mating biology of Hypoponera opacior, an ant that produces winged and wingless reproductives in a population in Arizona. Besides the investigation of the annual reproductive cycle, I particularly focused on the consequences of wingless reproduction. An analysis of sex ratios in wingless sexuals should reveal the relative importance of local resource competition among queens (that mainly compete for the help of workers) and local mate competition among males. Further, sexual selection was expected to act on wingless males that were previously found to mate with and mate-guard pupal queens in response to local mate competition. We studied whether males are able to adapt their mating behaviour to the current competitive situation in the nest and which traits are under selection in this mating situation. Last, we investigated the extent and effects of inbreeding. As the species appeared to produce non-dispersive males and queens quite frequently, we assumed to find no or only weak negative effects of inbreeding and potentially mechanisms that moderate inbreeding levels despite frequent nest-matings.rnWe found that winged and wingless males and queens are produced during two separate seasons of the year. Winged sexuals emerge in early summer and conduct nuptial flights in July, when climate conditions due to frequent rainfalls lower the risks of dispersal and independent colony foundation. In fall, wingless sexuals are produced that reproduce within the colonies leading to an expansion on the local scale. The absence of dispersal during this second reproductive season resulted in a local genetic population viscosity and high levels of inbreeding within the colonies. Male-biased sex ratios in fall indicated a greater importance of local resource competition among queens than local mate competition among males. Males were observed to adjust mate-guarding durations to the competitive situation (i.e. the number of competing males and pupae) in the nest, an adaptation that helps maximising their reproductive success. Further, sexual selection was found to act on the timing of emergence as well as on body size in these males, i.e. earlier emerging and larger males show a higher mating success. Genetic analyses revealed that wingless males do not actively avoid inbreeding by choosing less related queens as mating partners. Further, we detected diploid males, a male type that is produced instead of diploid females if close relatives mate. In contrast to many other Hymenopteran species, diploid males were here viable and able to sire sterile triploid offspring. They did not differ in lifespan, body size and mating success from “normal” haploid males. Hence, diploid male production in H. opacior is less costly than in other social Hymenopteran species. No evidence of inbreeding depression was found on the colony level but more inbred colonies invested more resources into the production of sexuals. This effect was more pronounced in the dispersive summer generation. The increased investment in outbreeding sexuals can be regarded as an active strategy to moderate the extent and effects of inbreeding. rnIn summary, my thesis describes an ant species that has evolved alternative reproductive tactics as an adaptation to seasonal environmental variations. Hereby, the species is able to maintain its adaptive mating system without suffering from negative effects due to the absence of dispersal flights in fall.rn

Classified Adélie penguin colonies from Landsat data

Breeding distribution of the Adelie penguin, Pygoscelis adeliae, was surveyed with Landsat-7 Enhanced Thematic Mapper Plus (ETM+) data along the coastline of Antarctica, an area covering approximately 330° of longitude. An algorithm was designed to minimize the radiometric contribution from exogenous sources and to retrieve Adelie penguin colony location and spatial extent from the ETM+ data. In all, 9143 individual pixels were classified as belonging to an Adelie penguin colony class out of the entire dataset of 195 ETM+ scenes, where the dimension of each pixel is 30 m by 30 m, and each scene is approximately 180 km by 180 km. Pixel clustering identified a total of 187 individual Adelie penguin colonies, ranging in size from a single pixel (900 m**2) to a maximum of 875 pixels (0.788 km**2). Colony retrievals have a very low error of commission, on the order of 1 percent or less, and the error of omission was estimated to be 2.9 percent by population based on comparisons with direct observations from surveys across east Antarctica. Thus, the Landsat retrievals can successfully locate Adelie penguin colonies that account for ~97 percent of a regional population. Geographic coordinates and the spatial extent of each colony retrieved from the Landsat data are available publically. Regional analysis found several areas where the Landsat retrievals suggest populations that are significantly larger than published estimates. Six Adelie penguin colonies were found that are believed to be unreported in the literature.