The evolution of social traits and biodiversity in the ants.

Cooperation has shaped the evolution of life on Earth. The ants are the most numerically diverse of the eusocial Hymenoptera, and display wide variation in social complexity. This positions the ants as an ideal taxon in which to study social evolution in a comparative framework. Social evolution theory has generated many hypotheses that are testable in ants, however the lack of comprehensive or complete phylogenies, and the decentralised and scattered nature of trait data, has been an obstacle to these types of study. In this thesis I construct a large species-level, and a complete genus-level, phylogeny of the ants, and draw together a large dataset of social traits from the literature in order to test hypotheses concerning the evolution of social traits in the ants. I find evidence that the earliest ant was large bodied, and lived in small highly related colonies. I show that group size is a significant trait in the evolution of sociality in ants, predicting the probability of a species having polymorphic workers, or of being polyandrous. I also show that the change in these traits is correlated between ancestral nodes on the phylogeny. Furthermore, in the Attini, colony size correlates closely with non-reproductive and reproductive division of labour. Together these results cement group size as a driving force of social evolution in the ants, and this has interesting implications for social evolution in general. Finally, I report the first evidence that intermediate colony sizes, the presence of discrete worker castes and polygyny are associated with increased diversification rates in ants. This thesis provides a valuable tool for the study of comparative hypotheses in the ants in the form of a complete genus-level phylogeny, and offers significant evidence to support several key hypotheses in social evolution. Furthermore, these results generate hypotheses regarding the evolution of social traits for future research

Eciton army ants - umbrella species for conservation in neotropical forests

Identification of priority areas for conservation is crucial for the maintenance and protection of biodiversity, particularly in tropical forests where biodiversity continues to be lost at alarming rates. Surveys and research on umbrella species can provide efficient and effective approaches to identify potential areas for conservation at small geographical scales. Army ants of the genus Eciton are keystone species in neotropical forests due to their major role as top predators and due to the numerous vertebrate- and invertebrate associated species that depend upon their colonies for survival. These associates range from the iconic army ant-following birds to a wide range of arthropod groups, some of which have evolved intricate morphological, behavioural and/or chemical strategies to conceal their presence and integrate into the colony life. Furthermore, Eciton colonies require large forested areas that support a diverse leaf litter prey community and several field-based and genetic studies have demonstrated the negative consequences of forest fragmentation for the long-term maintenance of these colonies. Therefore, Eciton species will not only act as umbrella for their associates but also for many other species in neotropical forests, in particular for those that require a large extent of forest. This review summarises past and recent accounts of the main taxonomic groups found associated with Eciton colonies, as well research assessing the impact of forest fragmentation on this army ant, to encourage the adoption of Eciton army ants as umbrella species for the identification of priority areas for conservation and assessments of the effect of disturbance in neotropical forests

The genomes of two key bumblebee species with primitive eusocial organization

Background: The shift from solitary to social behavior is one of the major evolutionary transitions. Primitively eusocial bumblebees are uniquely placed to illuminate the evolution of highly eusocial insect societies. Bumblebees are also invaluable natural and agricultural pollinators, and there is widespread concern over recent population declines in some species. High-quality genomic data will inform key aspects of bumblebee biology, including susceptibility to implicated population viability threats. Results: We report the high quality draft genome sequences of Bombus terrestris and Bombus impatiens, two ecologically dominant bumblebees and widely utilized study species. Comparing these new genomes to those of the highly eusocial honeybee Apis mellifera and other Hymenoptera, we identify deeply conserved similarities, as well as novelties key to the biology of these organisms. Some honeybee genome features thought to underpin advanced eusociality are also present in bumblebees, indicating an earlier evolution in the bee lineage. Xenobiotic detoxification and immune genes are similarly depauperate in bumblebees and honeybees, and multiple categories of genes linked to social organization, including development and behavior, show high conservation. Key differences identified include a bias in bumblebee chemoreception towards gustation from olfaction, and striking differences in microRNAs, potentially responsible for gene regulation underlying social and other traits. Conclusions: These two bumblebee genomes provide a foundation for post-genomic research on these key pollinators and insect societies. Overall, gene repertoires suggest that the route to advanced eusociality in bees was mediated by many small changes in many genes and processes, and not by notable expansion or depauperation

Chromosomal dynamics in space and time : evolutionary history of Mycetophylax ants across past climatic changes in the Brazilian Atlantic coast.

Fungus-farming ants of the genus Mycetophylax exhibit intra and interspecific chromosome variability, which makes them suitable for testing hypotheses about possible chromosomal rearrangements that endure lineage diversification. We combined cytogenetic and molecular data from Mycetophylax populations from coastal environments to trace the evolutionary history of the clade in light of chromosomal changes under a historical and geographic context. Our cytogenetic analyses revealed chromosomal differences within and among species. M. morschi exhibited three distinct karyotypes and considerable variability in the localization of 45S rDNA clusters. The molecular phylogeny was congruent with our cytogenetic findings. Biogeographical and divergence time dating analyses estimated that the most recent common ancestor of Mycetophylax would have originated at about 30?Ma in an area including the Amazon and Southern Grasslands, and several dispersion and vicariance events may have occurred before the colonization of the Brazilian Atlantic coast. Diversification of the psammophilous Mycetophylax first took place in the Middle Miocene (ca. 18?10?Ma) in the South Atlantic coast, while ?M. morschi? lineages diversified during the Pliocene-Pleistocene transition (ca. 3?2?Ma) through founder-event dispersal for the Northern coastal regions. Psammophilous Mycetophylax diversification fits into the major global climatic events that have had a direct impact on the changes in sea level as well as deep ecological impact throughout South America. We assume therefore that putative chromosomal rearrangements correlated with increased ecological stress during the past climatic transitions could have intensified and/or accompanied the divergence of the psammophilous Mycetophylax. We further reiterate that ?M. morschi? comprises a complex of at least three well-defined lineages, and we emphasize the role of this integrative approach for the identification and delimitation of evolutionary lineages

Gene content evolution in the arthropods

Arthropods comprise the largest and most diverse phylum on Earth and play vital roles in nearly every ecosystem. Their diversity stems in part from variations on a conserved body plan, resulting from and recorded in adaptive changes in the genome. Dissection of the genomic record of sequence change enables broad questions regarding genome evolution to be addressed, even across hyper-diverse taxa within arthropods. Using 76 whole genome sequences representing 21 orders spanning more than 500 million years of arthropod evolution, we document changes in gene and protein domain content and provide temporal and phylogenetic context for interpreting these innovations. We identify many novel gene families that arose early in the evolution of arthropods and during the diversification of insects into modern orders. We reveal unexpected variation in patterns of DNA methylation across arthropods and examples of gene family and protein domain evolution coincident with the appearance of notable phenotypic and physiological adaptations such as flight, metamorphosis, sociality, and chemoperception. These analyses demonstrate how large-scale comparative genomics can provide broad new insights into the genotype to phenotype map and generate testable hypotheses about the evolution of animal diversity

Diversification with Recurrent Evolution of Flightless Queens of the Ant Genus Acanthomyrmex in the Eastern Indochina and Adjacent Areas (Hymenoptera: Formicidae, Myrmicinae)

東京都立大学Tokyo Metropolitan University博士（理学）doctoral thesi

Virus discovery in monogynous and polygynous ant societies

Viruslöytö yksi- ja polygyniinimuurahaisissa yhteisöissä. Tiivistelmä. The interaction between host and pathogen is an intricate dance of survival, a constant tug-of-war in the evolutionary race. This relationship is not merely an act of invasion and defense but a complex dynamic process that shapes the natural world. The research focused on ant colonies, illuminating the complex relationship between ants and viruses. Theoretically, polygynous colonies, with their greater size and genetic diversity due to multiple queens, are better equipped to handle environmental fluctuations and disease. This study hypothesized that mature monogynous colonies, owing to their smaller size and the rigorous selection processes during colony establishment, which they have successfully endured, are likely exhibit a significantly lower viral diversity compared to polygynous colonies. To test this, RNA and small RNA were extracted from nine ant species — five monogynous and four polygynous. Data analysis unveiled a spectrum of 100 viruses, including 33 novel and 57 unknown entities. The results challenged initial expectations, with no statistically significant difference in virus diversity, viral load, and active infection between monogynous and polygynous colonies. Interestingly, monogynous colonies demonstrated a trend towards higher virus abundance and active infections. Furthermore, the research unveiled a geographical clustering (measured by total number of viruses per population). Moreover, a statistically significance relationship was identified between the virus abundance and active infections, as determined by RNAseq and small-RNAseq RPKM values. The findings of this study may suggest that polygynous species are better equipped for preventing the widespread transmission of infections within their colonies, in comparison to their monogynous counterparts. This research illuminate’s virus-host dynamics within social insects, enhancing knowledge on viral diversity in ants, and implying potential impacts on evolutionary biology, ecology, disease control, and biosecurity

Evolution of dispersal in ants (Hymenoptera: Formicidae) : a review on the dispersal strategies of sessile superorganisms

The extreme diversity of dispersal strategies in ants is unique among terrestrial animals. The nature of ant colonies as social, perennial, and sessile superorganisms is the basis for understanding this diversity, together with the inclusive-fitness framework for social evolution. We review ant dispersal strategies, with the aim of identifying future research directions on ant dispersal and its evolution. We list ultimate and proximate determinants of dispersal traits and the ecological and evolutionary consequences of dispersal for population structures and dynamics, as well as species communities. We outline the eco-evolutionary feedbacks between the multitude of traits affecting dispersal evolution and the likely evolutionary routes and ecological drivers in transitions among the diverse ant dispersal strategies. We conclude by presenting a research framework to fill the gaps in current knowledge, including comparative studies of colony life histories and population structures and theoretical models of the eco-evolutionary dynamics affecting dispersal, in an inclusive-fitness framework.Peer reviewe

Diversity, Biogeography and Community Ecology of Ants

Ants are a ubiquitous, highly diverse, and ecologically dominant faunal group. They represent a large proportion of global terrestrial faunal biomass and play key ecological roles as soil engineers, predators, and re-cyclers of nutrients. They have particularly important interactions with plants as defenders against herbivores, as seed dispersers, and as seed predators. One downside to the ecological importance of ants is that they feature on the list of the world’s worst invasive species. Ants have also been important for science as model organisms for studies of diversity, biogeography, and community ecology. Despite such importance, ants remain remarkably understudied. A large proportion of species are undescribed, the biogeographic histories of most taxa remain poorly known, and we have a limited understanding of spatial patterns of diversity and composition, along with the processes driving them. The papers in this Special Issue collectively address many of the most pressing questions relating to ant diversity. What is the level of ant diversity? What is the origin of this diversity, and how is it distributed at different spatial scales? What are the roles of niche partitioning and competition as regulators of local diversity? How do ants affect the ecosystems within which they occur? The answers to these questions provide valuable insights not just for ants, but for biodiversity more generally

The role of visual adaptation in cichlid fish speciation

D. Shane Wright (1) , Ole Seehausen (2), Ton G.G. Groothuis (1), Martine E. Maan (1) (1) University of Groningen; GELIFES; EGDB(2) Department of Fish Ecology & Evolution, EAWAG Centre for Ecology, Evolution and Biogeochemistry, Kastanienbaum AND Institute of Ecology and Evolution, Aquatic Ecology, University of Bern.In less than 15,000 years, Lake Victoria cichlid fishes have radiated into as many as 500 different species. Ecological and sexual sel ection are thought to contribute to this ongoing speciation process, but genetic differentiation remains low. However, recent work in visual pigment genes, opsins, has shown more diversity. Unlike neighboring Lakes Malawi and Tanganyika, Lake Victoria is highly turbid, resulting in a long wavelength shift in the light spectrum with increasing depth, providing an environmental gradient for exploring divergent coevolution in sensory systems and colour signals via sensory drive. Pundamilia pundamila and Pundamilia nyererei are two sympatric species found at rocky islands across southern portions of Lake Victoria, differing in male colouration and the depth they reside. Previous work has shown species differentiation in colour discrimination, corresponding to divergent female preferences for conspecific male colouration. A mechanistic link between colour vision and preference would provide a rapid route to reproductive isolation between divergently adapting populations. This link is tested by experimental manip ulation of colour vision - raising both species and their hybrids under light conditions mimicking shallow and deep habitats. We quantify the expression of retinal opsins and test behaviours important for speciation: mate choice, habitat preference, and fo raging performance