Abstract In social insects a major cost of social life is the high number of pathogens found in large societies and the greater likelihood of transmission of pathogens among closely related individuals. The aim of this thesis was to investigate the effect of high pathogen pressure on the molecular evolution of genes involved in the innate immune system in social insects. In addition, the transmission dynamics of the intracellular bacteria Wolbachia in wood ants was examined. By comparing DNA sequences from diverse species of ants and honeybees it was shown that the immune genes in hymenopteran social insects have evolved rapidly. However, by using codon-based likelihood models of evolution positive selection was detected in only two ant genes. This may reflect behaviourally based colony-level defences that can reduce selective pressure on the immune genes. The transmission modes of Wolbachia were studied by comparing DNA sequence variation of the bacteria with that of the host ants. First, it was found that all the studied ants carry Wolbachia. Second, Wolbachia have been transmitted both vertically from mother to offspring and horizontally between individuals of the same as well as of different species. / Tiivistelmä Yhteiskuntahyönteisten (muurahaiset, ampiaiset, mehiläiset ja termiitit) ekologisen menestyksen kääntöpuolena on ollut jatkuva riesa taudinaiheuttajista, joita suurissa yhteisöissä tavataan runsaammin kuin yksittäin elävissä eliöissä. Taudinaiheuttajien tuoman paineen myötä yhteiskuntahyönteisille on kehittynyt käyttäytymiseen perustuvia puolustusmekanismeja täydentämään kaikille monisoluisille eliöille yhteistä synnynnäistä, fysiologista immuniteettia. Nämä puolustusmekanismit ovat todiste siitä, että taudeilla on ollut suuri merkitys yhteiskuntahyönteisten käyttäytymisen evoluutiossa. Toisaalta taudinaiheuttajien vaikutuksista synnynnäiseen immuunipuolustukseen tiedetään hyvin vähän. Väitöstutkimuksen ensisijainen kohde oli taudinaiheuttajien merkitys yhteiskuntahyönteisten synnynnäisen immuunipuolustuksen evoluutiossa. Tutkimuksessa tarkasteltiin, miten immuunijärjestelmän geenit ovat ajan mittaan muuttuneet. Tulokset osoittivat että muutoksia, jotka johtavat proteiinien aminohappojen vaihtumiseen on tapahtunut tiuhempaan tahtiin muurahaisilla ja mehiläisillä kuin yksittäin elävällä banaanikärpäsellä. Merkkejä erityisen voimakkaasta luonnonvalinnasta löydettiin kuitenkin yllättävän pienestä määrästä geenejä. Tämä voi johtua siitä, että käyttäytymiseen perustuvat puolustusmekanismit lieventävät taudinaiheuttajien vaikutusta synnynnäiseen immuunipuolustukseen. Väitöstutkimukseen sisältyi myös hyönteisten solunsisäisen bakteerin, Wolbachian, siirtymismekanismien kartoitus kekomuurahaisilla. Wolbachia on loinen, joka siirtyy yleensä äidiltä jälkeläisille munasolussa. Leviäminen voi tapahtua myös horisontaalisesti lajitoverien ja jopa eri lajien edustajien kesken. Geenisekvensseihin perustuvassa tutkimuksessa kaikista muurahaisista löytyi Wolbachia-bakteereja, ja samasta yksilöstä saattoi löytyä useaa eri bakteerikantaa. Koska muurahaislajien väliset geneettiset erot olivat paljon suurempia kuin erot niissä elävien bakteerien välillä, voitiin päätellä että bakteerien pääasiallinen leviämistapa tutkituilla muurahaisilla on ollut horisontaalinen.

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La sélection de la parentèle est le concept actuellement le plus fréquemment avancé pour justifier l’évolution et le maintien d'une caste ouvrière stérile chez les Hyménoptères sociaux. La fourmi méditerranéenne Cataglyphis cursor possède plusieurs traits biologiques qui font de cette espèce un modèle particulièrement intéressant pour tester les prédictions de la théorie de la sélection de la parentèle, le plus important d'entre eux étant la capacité des ouvrières, qui ne s'accouplent jamais, à produire une descendance mâle (haploïde) par parthénogenèse arrhénotoque, ou femelle (diploïde) par parthénogenèse thélytoque. Nos analyses génétiques, basées sur des marqueurs microsatellites développés au préalable pour cette espèce, ont révélé que les reines utilisent la reproduction sexuée et asexuée respectivement pour la production de la caste ouvrière et reproductrice. L'analyse du pedigree des reproductrices issues de la reproduction asexuée nous a permis d'identifier le mécanisme cytologique de la parthénogenèse thélytoque et d'estimer la proportion de reines issues de la reproduction des ouvrières au sein de la population. De plus, bien que les reines soient capables de produire une descendance diploïde (femelle) par parthénogenèse thélytoque, elles ont conservé la reproduction sexuée pour la caste ouvrière et s'accouplent avec plusieurs mâles. Ceci indique que la reproduction sexuée a une fonction importante au niveau de la colonie, et nous avons testé certaines des hypothèses avancées pour justifier l'évolution de la polyandrie. Finalement, nous avons étudié l'impact des stratégies de dispersion de cette espèce sur le sex-ratio de la descendance sexuée. Ces résultats confirment l'intérêt que représente l'étude des stratégies reproductrices chez les Hyménoptères sociaux pour tester les prédiction de diverses théories en biologie évolutives et ouvrent également de nouvelles perspectives de recherche, tant chez C. cursor que chez d'autres espèces appartenant au genre Cataglyphis. / Kin selection is, to date, the most widely accepted theory to justify the evolution of a sterile worker caste among social Hymenoptera. The Mediterranean ant Cataglyphis cursor represents an interesting biological model for several reasons, the most important of them being the ability for unmated workers to produce haploid (male) offspring, through arrhenotokous parthenogenesis, and diploid (female) offspring, through thelytokous parthenogenesis. Our genetic analyses, based on microsatellite loci developed for this purpose, revealed that queens selectively use sexual and asexual reproduction to produce workers and sexuals, respectively. Pedigree analyses allowed us to identify the cytological mechanism involved in thelytokous parthenogenesis and to estimate the proportion of worker-produced queens in the study population. Although C. cursor queens do not require mating to produce diploid offspring, they have retained sexual reproduction and mate multiply with up to 8 males. This suggests that sexual reproduction has important benefits for colony function, and we tested several hypotheses accounting for the evolution of polyandry. Eventually, we studied the effect of dispersal strategies on sex-ratio of the sexual brood. These results confirm the interest of investigating the reproductive strategies of social Hymenoptera to test the predictions of diverse theories in the field of evolutionary biology, and open new research perspectives in C. cursor and other ants of the Cataglyphis genera. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

Doctorat en Sciences / info:eu-repo/semantics/nonPublished

Les savanes tropicales sont très importantes pour la production agricole et de nombreux services écosystémiques. En Colombie, ces savanes ont été traditionnellement gérées par l'élevage extensif et l'agriculture à faible intrants. La conversion actuelle de ces systèmes naturels en agriculture intensive peut avoir des effets dévastateurs sur la biodiversité du sol et au-dessus du sol. La macrofaune du sol représente une part importante de la biodiversité des agroécosystèmes et certains groupes ont reçu une attention considérable en tant qu'ingénieurs des écosystèmes. L'objectif général de la thèse est d'évaluer et d'analyser les impacts des paysages agricoles sur les communautés d'ingénieurs des sols et les services écosystémiques fournis par le sol dans les Llanos colombiens. Trois questions principales ont été abordées: (1) Quel est l'impact de la gestion agricole sur les communautés de fourmis et est-il possible d'identifier des espèces de fourmis qui pourraient servir d'indicateurs des services écosystémiques basés sur le sol? (2) Quel est l'impact de l’usage des sols sur les communautés de fourmis et de termites et cet impact est-il lié aux modifications des propriétés physiques et chimiques des sols? (3) Les traits écologiques et morphologiques des fourmis répondent-ils aux usages du sol et aux propriétés du sol?Les résultats de cette thèse confirment que les communautés de fourmis sont très sensibles aux changements d'usage de sols et constituent de bons indicateurs précoces des services écosystémiques basés sur les sols. Plus précisément, les cultures annuelles se sont révélées le plus préjudiciables aux communautés de fourmis et de termites, et cela est probablement dû à l'application d'intrants chimiques, ainsi qu'au travail du sol et aux pesticides. Tous les usages du sol (savanes, plantation de caoutchouc ou palmiers à huile, pâturages améliorés et cultures annuelles) ont une faune particulière, mais certaines espèces ont des besoins particuliers en termes d’habitat pour leurs nids, leur nourriture, ou des refuges. Certaines espèces sont plus généralistes et particulièrement adaptées aux environnements perturbés. En ce sens, la présence de toutes les utilisations du sol dans un paysage contribue dans une certaine mesure à la diversité totale. On a constaté que les usages des sols avec une couverture arborée permettent le maintien d’espèces rares, discrètes et souvent avec exigences de habitat particulaires. / Tropical savannas are highly important for agricultural production and many other ecosystems services. In Colombia, these savannas have been traditionally managed through extensive livestock production and low-input agriculture. The current conversion of these natural systems to intensified agriculture can have devastating impacts on belowground and aboveground biodiversity. Soil macrofauna represents an important part of agroecosystem biodiversity and some groups have received considerable attention as ecosystem engineers. The general goal of the thesis is to evaluate and analyze the impacts of agricultural landscapes on soil engineer communities and soil ecosystem services in the Colombian Llanos. Three main questions were addressed: (1) What is the impact of agricultural management on ant communities and is it possible to identify ant species that could be used as indicators of soil-based ecosystem services? (2) What is the impact of land uses on ant and termite communities and is this impact associated with modifications of soil physical and chemical properties? (3) Do the ecological and morphological traits of ants respond to land uses and soil properties?The results of this thesis confirm that ant communities are highly sensitive to land use changes and constitute good early indicators of soil-based ecosystem services. More accurately, annual crops have shown to be detrimental to ant and termites communities and this is probably due to the application of chemical inputs, as well as to tillage and pesticides. All land uses (Savannas, rubber or oil palm plantations, improved pastures and annual crops) have a particular soil fauna, but some species have particular habitat requirements for nest sites, food, refugees, etc. Some species are more generalist and are particularly adapted to disturbed environment. The presence of all land uses within a landscape contributes to a certain extent to the total diversity. It was found that sites with a tree cover protect rare, sometimes inconspicuous and perhaps fragile species.

The inadequacy of arbitrarily classifying interactions between species as antagonistic, neutral or mutualistic has become clear in recent years. Both direct and indirect interactions between species can vary between mutualism and antagonism, depending on the intrinsic and extrinsic contexts of the interaction. This study investigated the characteristics of an ant-plant-homopteran interaction in southern Africa. The polyphagous homopteran Hilda patruelis (Tettigometridae) feeds primarily on the trunk-borne fruiting branches and figs of the Cape fig tree, Ficus sur, and produces honeydew which attracts tending ants. Ten of the sixteen ant species/species groups present on F. sur tended H. patruelis, with Pheidole megacephala the most frequent attendant. Ants attracted to F. sur by H. patruelis honeydew or other liquid food sources also preyed on insects on the tree, including adults of the small agaonid fig wasps whose larvae feed on the ovules in the developing figs. One fig wasp species (Ceratosolen capensis) is also the tree's only pollinator. No benefits to H. patruelis from being tended by ants were detected, either in terms of reduced parasitism, or predation by a lycaenid caterpillar. A P. megacephala colony foraging on a F. sur tree was found to receive a high proportion of its likely energy requirements from the tree, mainly in the form of H. patruelis honeydew, during periods when it was bearing fruit. It is probable that the H. patruelis-P. megacephala interaction constitutes a direct mutualism at times, but that benefits to the homopteran are intermittent or weak. Both H. patruelis and ants benefitted from F. sur, directly or indirectly, through the provision of food (and for some ants, nesting sites). The removal of phloem sap by H. patruelis did not detectably reduce the trees' reproductive output, either in terms of pollinator or viable seed production. The indirect effects of ant and H. patruelis presence on the F. sur trees were on average positive, as ants preyed disproportionately heavily on fig wasp species parasitic on or competing with the pollinator, thus increasing pollinator production. Effects of ant presence on seed production were not investigated, but have been demonstrated as beneficial elsewhere. However, there is great varatlon both in the composition of the wasp fauna arriving to oviposit at different crops, and in ant densities per fig, on several temporal and spatial scales. This results in high variability in the effects of ants on the pollinator and, through it, the tree, from positive to zero and potentially even negative. Despite this conditionality of beneficial outcomes for the tree, the mean effect of ants on the F. sur population studied was to increase pollinator production by up to nearly 20%. This study is among the few to have demonstrated an overall benefit to a plant of having homopteran-tending ants present on it.

Human activities are increasing the size, frequency and severity of disturbance across earth’s ecosystems including deserts. Exotic annual grasses have altered fire regimes by increasing the size, frequency, and severity of fires in these systems. Invertebrates make up a large proportion of ecosystem diversity, provide a wide range of ecosystem functions, and are good indicators of ecosystem function and resilience. Ants are particularly good indicators of ecosystem stability. The ability of rodents to modify plant community structure post-fire, could result in rodent communities having important indirect effects on invertebrate communities. In chapter 1 we report changes in ant forager abundance and diversity with fire and rodent treatments over a three year period in the Great Basin. We found that while rodents had significant effects on the plant community in burned plots, this did not affect the ant community. Fire, however played a significant role in determining ant species richness and Shannon’s diversity index. Ant richness and diversity were reduced in burned areas compared to unburned areas. Total ant forager abundance was unaffected by fire, however, the abundance of the most common ant species, Pogonomyrmex occidentalis, increased in burned areas. The overall abundance of the other species was reduced in burned areas. We saw increases in the densities of P. occidentalis mounds in burned areas, but the average size of those discs decreased. The total area occupied by P. occidentalis mounds remained equal between burned and unburned plots. In chapter 2 we compare the abundances of different groups of invertebrates, as well as the abundances and diversity of the ant communities, between fire and rodent treatments. We then compared how those responses differed between sites in the Great Basin and Mojave deserts. In this study, we found that the abundances of most invertebrate groups remained unaffected by fire and rodent treatments. In the Great Basin, however, the abundance of flying-foragers was reduced in burned areas. At both locations, ant species richness and Shannon’s diversity were reduced in burned areas. Species richness and Shannon’s diversity were negatively correlated with invasive plant cover at both sites, and invasive plant cover was positively correlated with fire. The loss of diversity can spell losses in important ecosystem functions, and invasive grass fire cycles threaten to make these losses permanent.

Dans ce travail de thèse, nous présentons une méthode originale s’inspirant des comportements des fourmis réelles pour la résolution de problème de classification non supervisée non hiérarchique. Cette approche créée dynamiquement des groupes de données. Elle est basée sur le concept des fourmis artificielles qui se déplacent en même temps de manière complexe avec les règles de localisation simples. Chaque fourmi représente une donnée dans l’algorithme. Les mouvements des fourmis visent à créer des groupes homogènes de données qui évoluent ensemble dans une structure de graphe. Nous proposons également une méthode de construction incrémentale de graphes de voisinage par des fourmis artificielles. Nous proposons deux méthodes qui se dérivent parmi les algorithmes biomimétiques. Ces méthodes sont hybrides dans le sens où la recherche du nombre de classes, de départ, est effectuée par l’algorithme de classification K-Means, qui est utilisé pour initialiser la première partition et la structure de graphe. / In this work, we present a novel method based on behavior of real ants for solving unsupervised non-hierarchical classification problem. This approach dynamically creates data groups. It is based on the concept of artificial ants moving complexly at the same time with simple location rules. Each ant represents a data in the algorithm. The movements of ants aim to create homogenous data groups that evolve together in a graph structure. We also propose a method of incremental building neighborhood graphs by artificial ants. We propose two approaches that are derived among biomimetic algorithms, they are hybrid in the sense that the search for the number of classes starting, which are performed by the classical algorithm K-Means classification, it is used to initialize the first partition and the graph structure.

abstract: An insect society needs to share information about important resources in order to collectively exploit them. This task poses a dilemma if the colony must consider multiple resource types, such as food and nest sites. How does it allocate workers appropriately to each resource, and how does it adapt its recruitment communication to the specific needs of each resource type? In this dissertation, I investigate these questions in the ant Temnothorax rugatulus. In Chapter 1, I summarize relevant past work on food and nest recruitment. Then I describe T. rugatulus and its recruitment behavior, tandem running, and I explain its suitability for these questions. In Chapter 2, I investigate whether food and nest recruiters behave differently. I report two novel behaviors used by recruiters during their interaction with nestmates. Food recruiters perform these behaviors more often than nest recruiters, suggesting that they convey information about target type. In Chapter 3, I investigate whether colonies respond to a tradeoff between foraging and emigration by allocating their workforce adaptively. I describe how colonies responded when I posed a tradeoff by manipulating colony need for food and shelter and presenting both resources simultaneously. Recruitment and visitation to each target partially matched the predictions of the tradeoff hypothesis. In Chapter 4, I address the tuned error hypothesis, which states that the error rate in recruitment is adaptively tuned to the patch area of the target. Food tandem leaders lost followers at a higher rate than nest tandem leaders. This supports the tuned error hypothesis, because food targets generally have larger patch areas than nest targets with small entrances. This work shows that animal groups face tradeoffs as individual animals do. It also suggests that colonies spatially allocate their workforce according to resource type. Investigating recruitment for multiple resource types gives a better understanding of exploitation of each resource type, how colonies make collective decisions under conflicting goals, as well as how colonies manage the exploitation of multiple types of resources differently. This has implications for managing the health of economically important social insects such as honeybees or invasive fire ants. / Dissertation/Thesis / Doctoral Dissertation Biology 2019

MEnvSC / Department of Zoology