30 resultados para Gynes
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The increase in weight, fat and energy content of queens was studied in Iridomyrmex humilis (Mayr) in relation to the mode of colony founding in ants. The increase in energy content of gynes during the time between emergence and mating reaches only 80% in this species in which queens found colonies with the help of workers (dependent mode), whereas it can reach 470% in species in which queens found colonies without the help of workers (independent mode). These results are discussed with regard to the investment in energy required by each mode of colony founding.
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In ants, there are two main processes of colony founding, the independent and the dependent modes. In the first case young queens start colony founding without the help of workers, whereas in the second case they are accompanied by workers. To determine the relation between the mode of colony founding and the physiology of queens, we collected mature gynes of 24 ant species. Mature gynes of species utilizing independent colony founding had a far higher relative fat content than gynes of species employing dependent colony founding. These fat reserves are stored during the period of maturation, i.e. between the time of emergence and mating, and serve as fuel during the time of colony founding to nurture the queen and the brood. Gynes of species founding independently but non claustrally were found to have a relative fat content intermediate between the values found for gynes founding independently and those founding dependently. This suggests that such gynes rely partially on their fat reserves and partially on the energy provided by prey they collect to nurture themselves and the first brood during the time of colony founding. Study of the fat content of mature gynes of all species has shown that it gives a good indication of the mode of colony founding.
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Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)
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The behaviour and morphology of dwarf gynes produced in worker-sized cells of normal colonies in Nannotrigona testaceicornis (Meliponinae, Trigonini) were studied. The behaviour of these dwarf virgin queens was the same as observed for normal Trigonine gynes. The glandular equipment is also the same: Dufour glands, fat bodies and spermathecae are present. Despite these similarities, their ovaries are different. The functional significance of dwarf gynes is unknown, but may be a basis for an alternative reproductive strategy.
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Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)
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Ectatomma parasiticum Feitosa & Fresneau, a new ant species socially parasitic on Ectatomma tuberculatum (Olivier), is described from gynes discovered in Apazapan, state of Veracruz, Mexico, and reared in the laboratory. Ectatomma parasiticum is the first social parasite described in the Ectatomminae. This species can be distinguished from its host by morphological and behavioral features characteristic of the inquilines known in other ant subfamilies including reduced size, thickened petiole, and agonistic interactions with host species.
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The evolution of eusociality is one of the major evolutionary transitions of life on earth. For investigating the conditions and processes that are central to the origin of such integrated social organization, it is best to study organisms in which individuals have retained some flexibility in their reproductive strategies. Halictid bees are especially well suited as model organisms, because they show huge variation in social systems, both within and between species. In this thesis, I investigated female reproductive strategies in the primitively eusocial bee Halictus scabiosae, with a focus on the role of helpers, in order to get insight into the mechanisms governing the evolution and maintenance of eusociality. This species produces two broods per year. The females from the first brood can stay in the natal nest to help raise a second brood of males and gynes that become the next-generation foundresses in spring. We first compared the morphology of females from the two broods, as well as the nutrition they receive as larvae. Then we conducted a helper- removal experiment in the field to quantify the effects of the presence of helpers on colony survival and productivity. Finally, we reconstructed pedigree relationships of individuals using microsatellite markers in order to detect who reproduces in the nest and how much individuals drift between nests. We found that first brood females had a uniformly small size and low fat reserves, which may be caused by the restricted pollen and nectar provisions on which they develop. Colony survival and productivity was increased by the presence of a single helper, but the effect was small and mostly limited to small colonies. By inferring parentage within and across colonies, we could determine that females from the first brood rarely reproduce in their natal nests. However, foundresses are frequently replaced, and foundresses and females from the first brood occasionally move to and reproduce in foreign colonies. As a result, colonies often contain offspring from unrelated individuals, and the relatedness of females to the brood they rear is low. Overall, this thesis shows that the reproductive system of H. scabiosae is highly flexible. The production of helpers in the first brood is important for colony success and productivity, but there is a high colony failure rate and part of the first brood females drift and reproduce in foreign nests. Both foundresses and helpers appear to be constrained by harsh environmental conditions or social factors limiting reproduction and independent colony founding. - L'origine des insectes sociaux est un domaine fascinant pour la recherche. Pour comprendre les mécanismes et les conditions qui sont nécessaires pour l'évolution et le maintien de la vie en société, il est judicieux d'étudier des sociétés primitives d'insectes, où toutes les femelles ont conservé la capacité de se reproduire, même si leur rôle comportemental dans la colonie est d'aider sans se reproduire. Une des familles d'abeilles, les halictes, est idéale pour cette sorte de recherche, en raison de la grande variabilité dans leur comportement social. Dans cette thèse, j'ai étudié les stratégies reproductives des femelles de Halictus scabiosae pour mieux comprendre les mécanismes qui influencent l'évolution de la vie en société. Cette espèce produit deux cohortes de couvain par année. Les femelles du premier couvain restent souvent dans leur nid natal pour aider à élever le deuxième couvain, tandis que les femelles du deuxième couvain s'accouplent et hibernent pour devenir les nouvelles fondatrices au printemps suivant. Nous avons d'abord comparé la morphologie des femelles issues des deux couvains ainsi que leur nutrition au stade de larve. Puis, dans une expérience sur le terrain, nous avons quantifié l'apport d'une ouvrière pour la survie et la productivité de la colonie. Finalement, nous avons reconstruit des pedigrees en utilisant des marqueurs génétiques, pour savoir qui se reproduit dans la colonie et combien d'individus migrent entre colonies. Les résultats montrent que les femelles du premier couvain sont uniformément plus petites et plus maigres, ce qui indique que les fondatrices réduisent les provisions de nourriture pour leur premier couvain afin de les inciter à aider dans le nid au lieu de se reproduire indépendamment. Dans l'expérience sur le terrain, la survie et la productivité de la colonie augmentaient avec la présence d'une ouvrière additionnelle, mais l'effet était petit et limité aux petites colonies. Par la reconstruction de pedigrees, nous pouvions constater que les femelles du premier couvain pondent rarement dans leurs nids natals. Les fondatrices cependant sont souvent remplacées en cours de saison, et migrent fréquemment entre nids, tandis que les femelles du premier couvain pondent parfois des oeufs dans des nids étrangers. De ce fait, les colonies contiennent souvent des descendants d'individus étrangers, et la parenté génétique entre les femelles et le deuxième couvain est basse. Cette thèse démontre que le système reproductif de H. scabiosae est très flexible. La production d'ouvrières est importante pour la survie de la colonie et sa productivité, mais le taux d'échec est élevé et une partie des femelles du premier couvain migrent et pondent dans une colonie étrangère. Autant les fondatrices que les ouvrières semblent être contraintes par des conditions environnementales ou sociales qui limitent la reproduction et les nouvelles fondations de colonie. - Die Entstehung von sozialen Lebensformen ist eines der wichtigsten Entwicklungen in der Geschichte des Lebens. Um die Bedingungen oder Prozesse zu verstehen, welche bei der Entstehung und dem Erhalt von sozialen Merkmalen wichtig sind, sollte man Lebewesen untersuchen, welche je nach Umwelteinflüßen ihr soziales Verhalten flexibel ändern können. Furchenbienen (Halictidae) gehören dazu. Diese weisen nämlich ein breites Spektrum verschiedener sozialer Organisationsformen auf, oftmals sogar innerhalb der einzelnen Arten. In meiner Doktorarbeit befasste ich mich mit den Fortpflanzungsstrategien der Weibchen der Skabiosen-Furchenbiene Halictus scabiosae. Diese Art produziert zwei Brüten pro Jahr. Die Weibchen der ersten Brut bleiben dabei meist als Arbeiterinnen in ihrem Geburtsnest, wohingegen die Weibchen der zweiten Brut nach der Paarung überwintern, um im nächsten Frühling neue Kolonien zu gründen. In einem ersten Schritt verglichen wir die beiden Brüten bezüglich der Grösse und der Fettreserven der Weibchen sowie der Pollen-Nektar-Vorräte für die Larven. Dann bestimmten wir in einem Feldexperiment, wieviel eine zusätzliche Arbeiterin zum Überleben und zur Produktiviät der Kolonie beiträgt. Schliesslich ermittelten wir durch genetische Tests die Verwandtschaftsbeziehungen zwischen den Bienen, um herauszufinden, wer in den Kolonien tatsächlich die Eier legt und ob und wieviel die Bienen zwischen verschiedenen Nestern wandern. Wir stellten fest, dass die Weibchen von der ersten Brut einheitlich kleiner sind und weniger Fettreserven besitzen. Das weist daraufhin, dass die Nestgründerin die erste Brut unterernährt, um die Wahrscheinlichkeit zu erhöhen, dass diese Weibchen als Arbeiterinnen im Nest bleiben anstatt sich unabhängig fortzupflanzen. Schon eine einzelne zusätzliche Arbeiterin verbesserte die Überlebenschancen und Produktivität der Kolonie, der Effekt war allerdings klein und auf kleine Kolonien beschränkt. Die Verwandtschaftsanalysen zeigten, dass die Arbeiterinnen nur sehr selten ein Ei in ihr Geburtsnest legen. Erstaunlicherweise wanderten die Nestgründerinnen oft zwischen verschiedenen Nestern. Einige Weibchen der ersten Brut wanderten auch in ein fremdes Nest und produzierten dort Nachkommen. Diese Doktorarbeit zeigt, dass die Fortpflanzungsstrategien der Skabiosen-Furchenbiene tatsächlich sehr flexibel sind. Die Anwesenheit von Arbeiterinnen ist wichtig für das Überleben und die Produktivität der Kolonie. Die Misserfolgsraten bleiben jedoch hoch, und ein Teil der Weibchen der ersten Brut pflanzt sich in fremden Nestern fort. Sowohl die Nestgründerinnen als auch die Weibchen der ersten Brut scheinen durch Umweltsbedingungen oder durch soziale Faktoren in der Wahl ihrer Fortpflanzungs¬strategie eingeschränkt zu sein.
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ABSTRACT: INTRODUCTION: Primitively eusocial halictid bees are excellent systems to study the origin of eusociality, because all individuals have retained the ancestral ability to breed independently. In the sweat bee Halictus scabiosae, foundresses overwinter, establish nests and rear a first brood by mass-provisioning each offspring with pollen and nectar. The mothers may thus manipulate the phenotype of their offspring by restricting their food provisions. The first brood females generally help their mother to rear a second brood of males and gynes that become foundresses. However, the first brood females may also reproduce in their maternal or in other nests, or possibly enter early diapause. Here, we examined if the behavioural specialization of the first and second brood females was associated with between-brood differences in body size, energetic reserves and pollen provisions. RESULTS: The patterns of variation in adult body size, weight, fat content and food provisioned to the first and second brood indicate that H. scabiosae has dimorphic females. The first-brood females were significantly smaller, lighter and had lower fat reserves than the second-brood females and foundresses. The first-brood females were also less variable in size and fat content, and developed on homogeneously smaller pollen provisions. Foundresses were larger than gynes of the previous year, suggesting that small females were less likely to survive the winter. CONCLUSIONS: The marked size dimorphism between females produced in the first and second brood and the consistently smaller pollen provisions provided to the first brood suggest that the first brood females are channelled into a helper role during their pre-imaginal development. As a large body size is needed for successful hibernation, the mother may promote helping in her first brood offspring by restricting their food provisions. This pattern supports the hypothesis that parental manipulation may contribute to promote worker behaviour in primitively eusocial halictids.
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A study was undertaken to determine if there was a relation between the mode of colony founding in ants and the physiology of the new queens produced, in which mature gynes of 24 ant species were examined. Gynes of species utilizing independent colony founding had a far higher relative fat content (X±SD; 54±6%)(g fat/g dry weight) than gynes of species employing dependent colony founding(19±8%). Dimorphism between queens and workers was significantly higher in species employing independent colony founding. Thus independent colony founding not only results in production of queens with a relatively higher fat content and therefore with a higher energy content per g, but also results in the production of larger queens (in comparison with worker size). Of species employing independent colony founding, 80% were monogynous, whereas only 11% of the species employing dependent colony founding were monogynous. These results are discussed with regard to the social structure and life-history of ant species.
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Summary Among ants, wood ants are probably the most fascinating and studied species in temperate European forests. Unfortunately, due to several threats they are nowadays registered in red lists. Recent studies made in the Swiss Jura Mountains ended up in the description of a new sympatric sibling species of Formica lugubris (i.e. Formica paralugubris Seifert 1996). Because of this confusion the biology of F. lugubris is incomplete. Due to the extreme difficulties to distinguish morphologically F. lugubris from F. paralugubris we studied their cuticular hydrocarbons profiles. Irrespective of their geographic origin, we observed quantitative discrimination between species within each caste (workers, males and gynes =young alate female). Moreover, using a behavioural taxonomic approach (i.e. the pupa-carrying test) we showed that ants preferred conspecific worker pupae to those of the sibling species. These first results allowed us to consider the two species as two separate taxonomic units. To understand their coexistence, habitat distribution models were fitted with GIS predictors and factors known to influence wood ant distribution. In the Jura Mountains, although the two species share very similar habitats, they are spatially segregated. F. lugubris occurs more frequently at woodland borders than in forest interiors. We demonstrated with genetic and field data that Formica lugubris displays two different social forms in close proximity in alpine zone (e.g. unmanaged forests of the Swiss National Park). We discovered populations mostly monogynous to weakly polygynous (i.e. one to a few egg laying queens per colony) and monodomous (i.e. one nest per colony), and polygynous/polydomous populations (new nests being founded by colony budding). It is generally admitted that monogyne species disperse well in order to find suitable habitat to found new colonies whereas polygyne species have restricted dispersal and local mating within the nest. In order to compare reproductive strategies of F. lugubris and F. paralugubris (i.e. matings and dealation process) we conducted experiments with sexuals. F, lugubris gynes from monogynous/monodomous populations do not show a local strategy like the obligately polygynous F. paralugubris (i.e. early dealation even without mating, insemination without flight activity and low fat reserve). They always keep their wings, do not mate when not able to fly and have high amount of fat content revealing high survival capacities. On the other side, F, lugubris gynes from polygynous/polydomous populations have lower lipid reserves and displayed a reproductive behaviour close to the F. para lugubris one. After dispersal, wood ant gynes can either start new societies by temporary social parasitism of another species (i.e. subgenus Serviformica) or be adopted intraspecifically in an existing nest. In F. lugubris, we demonstrated that gynes from monogynous/monodomous colonies showed a high success for temporary social parasitism compare to the lower success of gynes from polygynous/polydomous colonies. However, physiological analyses suggested that only gynes from monogynous/ monodomous populations can efficiently disperse and found new nest by temporary social parasitism. Intraspecifically, gynes were accepted to a high degree in polygynous nest and in monogynous nests as long as these nests contained sexuals. In conclusion, Formica lugubris displays a social and dispersal polymorphism (mixed mating and founding system) representing a behavioural plasticity in relation to environmental and ecological conditions. Therefore, conservation measures directed toward this species should try to maintain a maximum of diversity at the habitat level. Résumé Les fourmis des bois sont probablement parmi les espèces de fourmis les plus fascinantes et les plus étudiées des forêts tempérées Européennes. Actuellement, du fait de différentes menaces, elles figurent malheureusement sur listes rouges. Plusieurs études menées au sein du Jura Suisse ont abouti à la description d'une nouvelle espèce jumelle et sympatrique de Formica lugubris (F. para- lugubris Seifert 1996). A cause de cette confusion la biologie de F lugubris est lacunaire. La distinction morphologique de F. lugubris et de F. para lugubris est si difficile que nous avons étudié leurs hydrocarbures cuticulaires. Indépendamment de l'origine géographique, nous avons observé une discrimination quantitative entre les espèces au sein de chaque caste (ouvrières, mâles et jeunes femelles ailées). De plus, à l'aide d'une approche taxonomique comportementale (le test de transport de cocons) nous avons montré que les fourmis préfèrent des cocons d'ouvrières conspécifiques à ceux de l'espèce jumelle. Ces premiers résultats nous permettent de considérer ces deux espèces comme deux unités taxonomiques distinctes et valables. Afin de comprendre leur coexistence, des modèles mathématiques ont été développés avec des données SIG et des facteurs écologiques influençant la répartition des fournis des bois. Dans le Jura, même si elles partagent des habitats fortement similaires, les deux espèces n'occupent pas les mêmes secteurs. F. lugubris est plus fréquente en lisière forestière plutôt qu'en pleine forêt. Nous avons démontré grâce à des données génétiques et de terrain que F. lugubris présente deux formes sociales au sein de la zone alpine (forêts protégées du Parc National Suisse). D'autre part, nous avons découvert des populations monogynes à faiblement polygynes (une à quelques reines pondeuses par colonie) et monodomes (colonies composées d'une seule fourmilière), et des populations polygynes/polydomes (les nouveaux nids étant produit par bourgeonnement). Généralement, les espèces monogynes dispersent sur de grandes distances et peuvent coloniser des habitats favorables à la fondation de nouvelles colonies alors que les espèces polygynes possèdent une dispersion limitée avec des accouplements à l'intérieur des nids. Afin de comparer les stratégies de reproduction de F. lugubris et de F. paralugubris (accouplements et perte des ailes) nous avons mené des expériences avec les sexués. Les jeunes femelles ailées de F. lugubris issues de populations monogynes/monodomes ne présentent pas de stratégie locale comparée à l'espèce obligatoirement polygyne F paralugubris (perte des ailes précoce même si il n'y a pas eu accouplement, insémination possible sans avoir volé activement et faibles réserves de graisse). Elles conservent toujours leurs ailes, ne s'accouplent pas lorsqu'elles sont empêchées de voler et possèdent de grandes quantités de graisse révélant de fortes capacités de survie. D'autre part, les jeunes femelles ailées de F. lugubris provenant de populations polygynes/polydomes ont peu de réserves lipidiques et ont un comportement de reproduction proche de celles de F. paralugubris. Après leur dispersion, les jeunes sexués femelles de fourmis des bois peuvent soit fonder une nouvelle société par parasitisme social temporaire d'un nid d'une autre espèce (sous-genre Serviformica) soit être adoptées dans un nid déjà existant de leur propre espèce. Chez F. lugubris, nous avons pu démontrer que les jeunes sexués femelles de colonies monogynes/monodomes présentent un succès élevé au parasitisme sociale temporaire en comparaison au plus faible succès obtenu avec des sexués provenant de colonies polygynes/polydomes. Cependant, les données physiologiques suggèrent que seules les jeunes sexués femelles de populations mono-gynes/monodomes peuvent disperser efficacement et fonder un nouveau nid par parasitisme social temporaire. Au niveau intraspécifique, les jeunes femelles sont acceptées à un taux élevé dans les nids polygynes mais aussi dans les nids monogynes tant que ces nids possèdent encore de jeunes sexués. En conclusion, F. lugubris est caractérisée par un polymorphisme dans ses structures sociales et ses stratégies de dispersion (système mixte d'accouplement et de fondation) ce qui représente une forte plasticité comportementale en relation avec les conditions environnementales et écologiques. Par conséquent, les mesures de conservation de cette espèce devraient s'attacher à maintenir un maximum de diversité au niveau des habitats.
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Genetic diversity benefits for social insect colonies headed by polyandrous queens have received intense attention, whereas sexual selection remains little explored. Yet mates of the same queen may engage in sperm competition over the siring of offspring, and this could confer benefits on queens if the most successful sire in each colony (the majority sire) produces gynes (daughter queens) of higher quality. These benefits could be increased if high-quality sires make queens increase the percentage of eggs that they fertilize (unfertilized eggs develop into sons in social hymenopterans), or if daughters of better genetic quality are over-represented in the gyne versus worker class. Such effects would lead to female-biased sex ratios in colonies with high-quality majority gynes. I tested these ideas in field colonies of Lasius niger black garden ants, using body mass of gynes as a fitness trait as it is known to correlate with future fecundity. Also, I established the paternity of gynes through microsatellite DNA offspring analyses. Majority sires did not always produce heavier gynes in L. niger, but whenever they did do so colonies produced more females, numerically and in terms of the energetic investment in female versus male production. Better quality sires may be able to induce queens to fertilize more eggs or so-called caste shunting may occur wherever the daughters of better males are preferentially shunted to into the gyne caste. My study supports that integrating sexual selection and social evolutionary studies may bring a deeper understanding of mating system evolution in social insects.
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Sweat bees exhibit a range of social behaviours, from solitary nesting, in which no workers are produced, to strong eusociality, in which workers exhibit a high degree of altruism, behaviour that is measured by the degree of personal reproductive sacrifice. Field studies were carried out for seven weeks during May-June 2000 in southern Greece in order to investigate intraspecific social variation, and test the hypothesis of a north-south cline of decreasing eusociality in the obligately eusocial sweat bee L. (E.) malachurum. A comparative study, using principal components analysis, was performed to determine if patterns of intraspecific social variation in L. malachurum reflect the patterns of social variation within the subgenus, Evylaeus, as a whole. The results of the field study reveal that, in Greece, two worker broods were produced followed by a third brood consisting of gynes, males and some workers, indicating that there was an overlap in worker and gyne production. There was strong caste distinction between queens and workers. Workers actively foraged and participated in nest construction as most workers (58%, n=303) had a high degree of mandibular wear. Workers did not participate in the oviposition of Brood 3 gynes since only 0.7% (n=278) of workers were mated. Furthermore, queen survival until the end of Brood 3 and a substantial size differential of 10.6% between queens and workers suggested that queen domination over worker behaviour during the early to mid-part of the colony cycle was plausible. Male production in Brood 3 by some workers was likely, since the timing of worker ovarian development corresponded with the timing of male production. These findings suggest that workers of the first two broods were primarily altruistic, but some (28%) Brood 1 (9%) and Brood 2 (19%) workers produced males, indicating that the degree of altruistic behaviour declined during the lifetime of the colony. In comparison with other L. malachurum populations in Europe, the Greek population of L. malachurum had a weaker social level as a result of the higher proportion of workers potentially involved in male production, thus 3 supporting the hypothesis of a southerly cline of decreasing eusociality. Furthermore, intraspecific variation in social level across Europe appears to be due to longer breeding seasons in more southerly locations that would promote the production of larger colonies and provide opportunities for workers to evade queen control. The comparative study using principal components analysis on 20 solitary (of the subgenera Evylaeus and Lasioglossum), eusocial and socially polymorphic Evylaeus species and populations reveals that six traits are closely associated with stronger eusociality in Evylaeus. These traits are: (1) a reduction in the proportion of males in the early brood(s); (2) a reduction in the proportion of females that mate; (3) an increase in the mean number of first brood workers; (4) a reduction in the proportion of females with developed ovaries; (5) an increase in size dimorphism between castes, and (6) nest guarding. These are traits that most significantly define principal component one and therefore distinguish social type as indicated by a clear separation of the eusocial and the solitary populations, with a socially polymorphic species falling in between. Furthermore, most of these traits are under foundress control and may suggest that the evolutionary loss or gain of eusociality is based on selection pressures on a founding female. Colony size and female ovarian development are common factors distinguishing social variation in L. malachurum and within the subgenus as a whole. The principal components analysis excluding the solitary species and the socially aberrant L. marginatum populations show the L. malachurum populations separated based on an increasing proportion of workers with developed ovaries as populations are found more south, lending further support to the hypothesis of a north-south cline of decreasing eusociality.
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Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)
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Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)
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The production of sexual forms in laboratory colonies of Monomorium pharaonis was followed over a 19-month period. Cycles of production occurred at intervals of 4 months, with male investment inversely proportional to the number of gynes produced. There was a tendency for colonies to produce only males or females as total sexual production increased. Sperm counts of known-aged males suggest that they mate twice, in contrast to European populations. Sex-ratio was heavily female-biased, contrary to predictions for polygynous ant species.