Convergent genetic architecture underlies social organization in ants.

Complex adaptive polymorphisms are common in nature, but what mechanisms maintain the underlying favorable allelic combinations [1-4]? The convergent evolution of polymorphic social organization in two independent ant species provides a great opportunity to investigate how genomes evolved under parallel selection. Here, we demonstrate that a large, nonrecombining "social chromosome" is associated with social organization in the Alpine silver ant, Formica selysi. This social chromosome shares architectural characteristics with that of the fire ant Solenopsis invicta [2], but the two show no detectable similarity in gene content. The discovery of convergence at two levels-the phenotype and the genetic architecture associated with alternative social forms-points at general genetic mechanisms underlying transitions in social organization. More broadly, our findings are consistent with recent theoretical studies suggesting that suppression of recombination plays a key role in facilitating coordinated shifts in coadapted traits [5, 6].

Relaxed selection is a precursor to the evolution of phenotypic plasticity.

Phenotypic plasticity allows organisms to produce alternative phenotypes under different conditions and represents one of the most important ways by which organisms adaptively respond to the environment. However, the relationship between phenotypic plasticity and molecular evolution remains poorly understood. We addressed this issue by investigating the evolution of genes associated with phenotypically plastic castes, sexes, and developmental stages of the fire ant Solenopsis invicta. We first determined if genes associated with phenotypic plasticity in S. invicta evolved at a rapid rate, as predicted under theoretical models. We found that genes differentially expressed between S. invicta castes, sexes, and developmental stages all exhibited elevated rates of evolution compared with ubiquitously expressed genes. We next investigated the evolutionary history of genes associated with the production of castes. Surprisingly, we found that orthologs of caste-biased genes in S. invicta and the social bee Apis mellifera evolved rapidly in lineages without castes. Thus, in contrast to some theoretical predictions, our results suggest that rapid rates of molecular evolution may not arise primarily as a consequence of phenotypic plasticity. Instead, genes evolving under relaxed purifying selection may more readily adopt new forms of biased expression during the evolution of alternate phenotypes. These results suggest that relaxed selective constraint on protein-coding genes is an important and underappreciated element in the evolutionary origin of phenotypic plasticity.

The assessment of reproductive success of queens in ants and other social insects

Recently considerable research has focused on the causes of evolution of multiple-queen (polygynous) colonies. In order to better understand the factors which may have led to these polygynous associations it is vital to compare the reproductive success of queens in monogynous (one queen per colony) and polygynous colonies as well as the relative fitness of queens in polygynous colonies. This paper addresses the difficulties arising from such comparisons and their implications with regard to the methods commonly used to assess reproductive success in queens. The relative reproductive success of queens in monogynous and polygynous colonies is commonly assessed by comparing the relative number of reproductives they produce during a single reproductive season. However, shift in queen number seems to be only one aspect of a profound shift in social structure and reproductive strategy that constitutes, in effect, a ''polygyny syndrome''. For example, female reproductives produced in polygynous colonies frequently use a different mode of colony founding, which in turn affects the probability of their survival. Furthermore, queens from monogynous and polygynous colonies frequently differ in their life-span and the number of sexual broods they produce. As a result, the reproductive success of queens in monogynous and polygynous colonies may not be directly related to the relative number of sexuals they produce during a single reproductive season.

Studying the molecular causes of ageing in ants

RÉSUMÉ DE THÈSE Au cours de ma thèse, je me suis intéressée aux causes physiologiques du vieillissement en utilisant les fourmis comme modèle. Les trois castes de fourmis - les mâles, les ouvrières et les reines - présentent des longévités très différentes, tout en étant génétiquement identiques. Ceci implique que les différences de longévité sont dues à des variations entre castes dans le pattern d'expression de gènes. Mon travail chez la fourmi a consisté d'une part à mettre en place les outils pour identifier de tels gènes à grande échelle, de l'autre à étudier le rôle de gènes et de mécanismes qui affectent la longévité chez d'autres espèces. Pour identifier de nouveaux gènes potentiellement impliqués dans le vieillissement, nous avons développé des puces à ADN. Cette technique permet la comparaison du niveau d'expression de milliers de gènes entre deux échantillons. L'application de cette méthode aux reines et ouvrières adultes nous a jusqu'à présent permis d'identifier neuf gènes surexprimés chez les reines. Trois d'entre eux sont potentiellement impliqués dans le maintien et la réparation du soma, deux processus qui sont supposés avoir un impact crucial sur la longévité. Parmi les mécanismes impliqués dans le vieillissement chez d'autres espèces, nous nous sommes principalement intéressés aux télomères, qui sont les extrémités des chromosomes. Chez les vertébrés, les télomères se raccourcissent à chaque division cellulaire, entre autres parce que l'ADN polymérase ne peut répliquer cette partie des chromosomes en entier. Or des télomères courts entravent la prolifération des cellules et peuvent même induire l'apoptose, ce qui pourrait se répercuter sur la capacité des organismes à régénérer des tissus. J'ai pu montrer que chez les fourmis mâles (la caste qui vit le moins longtemps) les télomères se raccourcissent beaucoup plus vite que chez les reines et les ouvrières. L'explication la plus plausible pour cette différence est que les mâles, étant adapté à une vie très éphémère, n'investissent qu'un minimum d'énergie dans la machinerie de maintenance qui assure le bon fonctionnement des cellules. Ces résultats sont intéressants car ils permettent pour la première fois de faire le lien entre les théories évolutives du vieillissement et la biologie des télomères. THESIS ABSTRACT During my thesis I used ants as a model to study the proximate (i.e., molecular) causes of ageing and lifespan determination. Ant queens, workers and males differ tremendously in lifespan, although all three castes are genetically identical. Importantly, this implies that genes and molecular pathways responsible for modulating lifespan are regulated in a caste-specific manner. To find new genes potentially involved in ageing, we first constructed 371-gene-cDNA microarrays for the ant L. niger. This molecular tool can be used to survey the relative gene expression levels of two samples for thousands of genes simultaneously. By applying this method to adult queens and workers we identified nine genes that are overexpressed in queens. Three of them are putatively involved in somatic maintenance and repair, two processes that have been previously suggested as important for ageing and lifespan determination. We expect to identify many more candidate genes in the near future by using the 9000-gene fire ant microarrays we have recently developed. We also investigated whether factors linked to ageing in other organisms could affect lifespan determination in ants. One project was on telomeres, the ends of linear chromosomes. For various reasons telomeres shorten with every cell division. Since short telomeres can lead to cellular defects such as impaired cell division, telomeres have been hypothesized as playing a role in ageing. We tested whether telomere length in ant somatic tissues correlates with caste-specific lifespan in young adults. The short-lived L. niger mates did indeed have significantly shorter telomeres than the longer-lived queens and workers, probably because telomere attrition is faster in males than in queens and workers. Queens did not, however, have longer telomeres than the shorter-lived workers. These findings are consistent with the idea that telomere length may play a role in ageing under some circumstances, but they also clearly demonstrate that other factors must be involved. We argue that sex-specific telomere length patterns in ants ultimately reflect adaptive differences in the level of somatic maintenance between males and females, and thus create a link between telomere biology and the evolutionary theory of ageing.

Female reproductive system of the decapitating fly Pseudacteon wasmanni Schmitz (Diptera : Phoridae)

Pseudacteon wasmanni is a South American decapitating fly that parasitizes workers of Solenopsis fire ants. We used light microscopy (historesin serial-sectioning stained with Haematoxylin/Eosin) and scanning electron microscopy to show and analyze internal and whole external views of the female reproductive system. All specimens analyzed (n = 9) by light microscopy showed post-vitellogenic oocytes inside the ovaries. The lack of typical follicles (oocyte-nurse cell complexes) in all specimens suggests that oogenesis occurs during the pupal stage. The total number of eggs found ranged from 31 to 280 (X = 142 +/- 73, SD). The egg has a slugform or torpedo shape (about 130 by 20 mum) with a pointed apex at the posterior pole as defined by the fly; the micropyle appears to be in a depression or invagination at the anterior pole. An acute hypodermic-like ovipositor is evaginated from the hard sclerotized external genitalia during egg laying. The existence of a muscular bulb associated with the end of the common oviduct suggests that the egg is injected into the ant's body by a strong contraction of the bulb which probably is stimulated by bending of several ventral sensilla. During contraction, the abdomen extends out along a large fold between the sixth and seventh tergites in such a way that the sclerotized genitalia is rotated ventrally into a slightly anterior orientation in preparation for oviposition. (C) 2003 Elsevier B.V. Ltd. All rights reserved.

Sobre um novo Nicoletiidae (Zygentoma: Insecta) do Brasil vivendo com formigas lava-pés (Hymenoptera: Formicidae)

Descreve-se um novo Zygentoma (Nicoletiidae: Subnicoletiinae), mirmecófilo da formiga lava-pés Solenopsis saevissima (Formicidae: Myrmicinae) no Estado do Rio de Janeiro, Brasil: Allotrichotriura saevissima gen. nov. sp. nov. que é comparado com os géneros e subgéneros conhecidos na subfamília. As principais características diagnósticas respeitam a combinação da forma do corpo, quetotaxia cefálica e do corpo, morfologia do prétarso e número de estilos e vesículas abdominais. Embora prospecções recentes tenham sido levadas a cabo na localidade típica, apenas se conhece a amostra original, que integra exclusivamente fêmeas.

EXPERIMENTAL INCIPIENT NEST TRANSPLANTS AND SURVIVORSHIP OF SOLENOPSIS-INVICTA BUREN (HYM, FORMICIDAE) IN BRAZIL

Transplants of incipient colonies of the fire ant, Solenopsis invicta into areas in which mature colonies had been eliminated did not result in higher rates of success than in areas in which mature colonies were present. However for both types of sites, Pogonomyrmex (Ephebomyrmex) nagelli colonies were found in cleared sites used for transplants, with the number of translocated P. nagelli nests higher in areas which had not been treated with Mirex for S. invicta colony reduction. These results suggest that small colonies may be displaced by larger colonies of other species.

Análises moleculares das formigas lava-pés (Solenopsis spp.) (Hymenoptera : Formicidae) e da presença da endobactéria Wolbachia

Pós-graduação em Ciências Biológicas (Biologia Celular e Molecular) - IBRC

Biologia, morfologia e bioquímica de veneno da formiga lava-pés Solenopsis saevissima Smith (Insecta, Hymenoptera, Formicidae)

Pós-graduação em Ciências Biológicas (Zoologia) - IBRC

A 15-year post evaluation of the fire effects on ant community in an area of Amazonian forest

Fire represents an important disturbance to ant communities in areas of fire regime. Otherwise, little is known about the effects of fire on ant communities in areas of non-fire regimes, such as in the Amazonian region. We evaluated the long-term effect of fire on ant species richness in a rain forest (Bacaba Plateau) burned 15-years ago and compare our data with the data of primary unburned forest. A total of 85 ant species distributed in 21 genera and 14 tribes were collected; among them, 72 and 44 species were found on the litter and vegetation, respectively. The fire damaged forest studied supports an intermediate richness of ants when compared to a primary unburned rain forest in the same region. A comparative analysis of ant species richness showed that the Bacaba Plateau presented a different ant fauna when compared with the primary unburned forests, suggesting that fire can alter ant species composition. Although, our results cannot be conclusive on the effects of fire on ant community, they represent a pioneer data on human induced fire in tropical rain forests.

Fire management and the nesting of Athene cunicularia (Aves, Strigidae) in grasslands in central Cerrado, Brazil

Fire management is a common practice in several reserves in the Cerrado, but its influences on bird reproduction remain unknown. In addition, the nesting biology of the Burrowing Owl (Athene cunicularia) has been studied in numerous environments, but not in tropical grasslands managed by fire. This study examined the effects of fire management on the nesting biology of A. cunicularia in Emas National Park, State of Goias, central Brazilian Cerrado. We compared the number of breeding pairs and their burrows in October and November 2009 at 15 study sites in grasslands managed by fire (firebreaks) and unmanaged grasslands adjacent to and distant from firebreaks. We visited active burrows two-four times and described the burrow entrances and sentinel sites and counted and observed adults and young. A total of 19 burrows were found at firebreaks. One and two burrows were found in grasslands adjacent to and distant from firebreaks, respectively. For all burrows found, one to three young reached the adult size, being able to fly and/or run in early November. The 22 burrows found were in the ground, associated or not with termite and ant nests. Most (86.4%) burrows had only one entrance. Only three burrows had two or three entrances. Structures used as sentinel perches by adults were mounds in front of the burrow entrances, termite nests, shrubs and trees. Most of these sentinel sites were shorter than 2 m high and located less than 10 m from the burrow entrance. At Emas National Park, firebreaks appear to provide more attractive conditions to the nesting of A. cunicularia than unmanaged grasslands, likely because of the short herbaceous stratum due to frequent burning of firebreaks. This study suggests that fire management provides suitable conditions for the successful reproduction of A. cunicularia in firebreaks at Emas National Park.

Mitochondrial genome evolution in fire ants (Hymenoptera: Formicidae).

BACKGROUND: Complete mitochondrial genome sequences have become important tools for the study of genome architecture, phylogeny, and molecular evolution. Despite the rapid increase in available mitogenomes, the taxonomic sampling often poorly reflects phylogenetic diversity and is often also biased to represent deeper (family-level) evolutionary relationships. RESULTS: We present the first fully sequenced ant (Hymenoptera: Formicidae) mitochondrial genomes. We sampled four mitogenomes from three species of fire ants, genus Solenopsis, which represent various evolutionary depths. Overall, ant mitogenomes appear to be typical of hymenopteran mitogenomes, displaying a general A+T-bias. The Solenopsis mitogenomes are slightly more compact than other hymentoperan mitogenomes (~15.5 kb), retaining all protein coding genes, ribosomal, and transfer RNAs. We also present evidence of recombination between the mitogenomes of the two conspecific Solenopsis mitogenomes. Finally, we discuss potential ways to improve the estimation of phylogenies using complete mitochondrial genome sequences. CONCLUSIONS: The ant mitogenome presents an important addition to the continued efforts in studying hymenopteran mitogenome architecture, evolution, and phylogenetics. We provide further evidence that the sampling across many taxonomic levels (including conspecifics and congeners) is useful and important to gain detailed insights into mitogenome evolution. We also discuss ways that may help improve the use of mitogenomes in phylogenetic analyses by accounting for non-stationary and non-homogeneous evolution among branches.

Morphological Notes on the Worker and Queen Larvae of the Thief Ant Solenopsis helena (Hymenoptera, Formicidae, Myrmicinae) from Brazil

Young and mature larvae of queen and workers of the thief ant, Solenopsis helena, are herein described for the first time. Specimens were treated and described by usual methods of light and scanning electron microscopy. Many of the observed characteristics confirmed traits of the thief ant larva, reinforcing the assumption that they are a distinct group from fire ants. The larva of S. helena can be recognized from other close species by details of the mouthparts, but seem extensively similar to Solenopsis molesta. These findings are useful for taxonomic studies and phylogenetic inferences.

Ant predation of larval Diatraea saccharalis Fab. (Lep., Crambidae) in new sugarcane in Brazil

Ant predation of natural field populations of larval sugarcane borers, Diatraea saccharalis, was evaluated by using insecticidal check techniques, and through survivorship studies of artificial high-density infestations, in plantations in Brazil. Areas of ant suppression had higher levels of D. saccharalis than unsuppressed areas, although both types of areas normally had extremely low D. saccharalis populations. Artifical infestations of D. saccharalis yielded significantly lower larval survivorship in unsuppressed areas than in suppressed areas. Fire ants (Solenopsis spp.) were always the most abundant ant. This study has demonstrated that ants are important pest predators in Brazilian sugarcane fields.