This work further elucidates processes involved in promoting and sustaining evolutionary diversification within the gall-inducing thrips that specialise on Australian Acacia. A phylogenetic approach was taken to determine modes of diversification available to these insects. The extension and revision of the gall-thrips phylogeny is central to the work and primarily focuses on cryptic populations of the Kladothrips rugosus and Kladothrips waterhousei species complexes. Parallel diversification, where the radiation of the K. rugosus and K. waterhousei lineages broadly mirror one another, offered a rare opportunity to test hypotheses of coevolution between gall-thrips and their Acacia hosts. In the absence of a reliable host Acacia phylogeny, indirect inference of insect/plant cospeciation can be arrived at as these two complexes share the same set of host species. The expectation is that if the phylogenies for the gall-thrips complexes show a significant level of concordance, then cospeciation between insect and host-plant can be inferred. Results indicate that the K. rugosus species complex comprise populations at species level. A significant level of phylogenetic concordance between the two species complexes is consistent with gall-thrips lineages tracking the diversification of their Acacia hosts. Given the less than strict form of insect/host cospeciation, factors impacting host diversification become important to gall-thrips diversification. Gall-thrips radiated over a period during the expansion of the Australian arid-zone. Cycles of host range expansion and fragmentation during the Quaternary could have played a major role in gall-thrips diversity. An interesting feature of resourse sharing amongst the K. rugosus and K. waterhousei complex members is the apparent absence of competitive exclusion between them. The persistence of this sympatry over millions of years is an unusual feature and merits further investigation.

DNA barcoding has illuminated genetically distinct lineages within what appears to be one morphological species. For example, a large-scale DNA barcode analysis of Lepidoptera in the Área de Conservación Guanacaste has revealed that 8% of the morphospecies show more than one DNA barcode lineage. To assess the evolutionary significance of five of these lineages I conducted further molecular analyses by sequencing mitochondrial cytochrome b, nuclear Elongation Factor 1α and ribosomal Internal Transcribed Spacer 2, and compare their gene phylogenies with the provisional species tree hypothesized by DNA barcode genetic distances. Both mitochondrial and nuclear markers support the existence of three cryptic species in one of the five cases. Morphological and ecological correlates are still lacking to understand the origin of this divergence. The lack of corroboration between markers in the four remaining species either suggests that the chosen nuclear markers have not diverged since speciation, or there has been recent hybridization between lineages. In one case, hybridization is strongly suggested. / Natural Sciences and Engineering Research Council of Canada, Genome Canada through the Ontario Genomics Institute

The morphological phenotypic characters involving sexual selection but with highly individual variability are likely to challenge the prezygotic isolating mechanism driven by differentiation of mechanical structures. This kind of characters may also puzzle species identification and taxonomy. Therefore clarifying the correlation between the phenotypic variability and biological/non-biological factors becomes necessary in order to understand the role of this phenomenon under natural selection and sexual selection. The Syntomini represents one of the few lepidopterous groups that exhibit highly individual variability in both wing pattern and reproductive structures. The evolutionary and taxonomic significance of this phenomenon, however, has never been studied using modern methods although it has been documented for long. In order to test several hypotheses relevant to phenotypic variability, the present study focuses the phylogenetic relationship of Amata wilemani Rothschild, 1914, a subalpine moth species with extremely high variability in wing coloration and genitalia. The phylogenetic relationship between the three color morphs of A. wilemani and 38 Syntomini species plus 2 Lithosiinae outgroups was reconstructed using fragments of COI, EF1a and 28S. All color morphs of A. wilemani were recovered to form a monophyletic group under all data partitioning strategies with Amata formosensis (Wileman, 1928) or its closely related species in China as the potential sister group. The result of gene network analysis suggests low divergence between haplotypes of A. wilemani. Because no correlation between color morphs, phenology, geographical distribution, altitudinal gradient, and genitalic morphlogy was detected, it is concluded that A. wilemani should be regarded as a single species with high phenotypic variability, and this may suggest existence of intraspecific competition. The present study also found that Amata karapinensis (Strand, 1915), which was synonymized with A. wilemani by previous authors, should be revived. The incongruence between the phylogenetic relationships based on morphological and molecular characters shows a need of a comprehensive phylogenetic study of this highly diverse group.

3 ABSTRACT Freshwateramphipodsarepoordispersersduetothefragmentarynatureoftheir habitat,benthiclife-styleandlackofresistantpropagules.Consequently,mostspeciesare narrowlyendemic,andwidespreadmorphospeciesconsistofnumerousdivergentlineages. Assuch,thesepatternsareconsideredrelict, mainlyreflectinghistoricalratherthan contemporaryfactors.TheCarpathian Mountainsunderwentadramaticgeomorphological evolutionduringthelast20millionyears,risingfromtheParatethysSeathroughaseriesof islandsthateventuallycoalescedintooneofthelongest mountainchainsinEurope. Moreover,these mountainsandsurroundingbasinsareinhabitedbyaubiquitousand diverseamphipodfauna,providinganidealsettingfortestinghistoricalbiogeographical hypotheses.OurresultshaveshownthatinthesouthernhalfoftheCarpathians,montane Gammarus taxadisplayfragmentedandallopatricdistributionpatterns whilelowland speciesbelongingtothegeneraNiphargus andSynurella are mostlysympatricand widespreadoverlargedistances.Subsequent molecularinvestigationsrevealedthatthe patchydistributionofG.fossaruminthewesternandsouthwesternCarpathianshaslikely resultedfromthebreakupofaformerlycontiguouslandmass16 millionyearsago, suggestingthatgeologicalsubsidencecanplayanimportantroleinshapingbiogeographical patterns. Moreover,the WesternCarpathianlineagesineasternCzechiaandSlovakiaare...

Stylaria is a genus of globally distributed clitellates consisting of two recognized species with a difficult taxonomic history: Stylaria lacustris and Stylaria fossularis. The current species-level taxonomy of this genus is unclear due to variation in several morphological characters. To assess the amount of species-level diversity in Stylaria, an integrative approach using morphological investigations and genetic information was employed. I investigated the phylogenetics of this genus using three loci, two mitochondrial (cox1, and rrnL), and one nuclear (ITS) gene regions. Using several species delimitation tools, I was able to estimate the number of species in this group. Automatic barcode gap discovery, assemble species by automatic partitioning (ASAP), and the Geneious Prime Species delimitation package all delimited at least five species within Stylaria, Bayesian Poisson Tree Processes(bPTP) recovered at least six, and DELINEATE proposed only three species, all within S. fossularis. Morphologically, there seems to be no clear, consistent distinction between members of S. lacustris from Europe and S. lacustris from North America. Based on my morphological investigations and disagreement among species delimitation methods, I conservatively propose that the subclades discovered via phylogenetic analysis of mitochondrial and nuclear sequence data within S. lacustris represent population-level rather than species-level variation. However, there appear to be at least two previously unknown species in need of proper descriptions within the S. fossularis species complex. Within Stylaria, morphological criteria might be useless at the species level as there is a high level of intraspecific morphological variation. Analyses of sexual characteristics of this group could help further document any differences between populations of S. lacustris.

In the marine environment, relatively little is known about how the effects of climatic change will manifest into future patterns of biodiversity and community stability, with the exception of recent work on corals in tropical reef ecosystems. In this thesis, I examine quantitatively how patterns of coral reef biodiversity will respond to the impacts of climatic change and provide essential information (i) to address a critical knowledge gap in the understanding of coral reef biodiversity and (ii) predict how a wide range of sessile invertebrates from coral reef ecosystems will respond to a range of potential impacts of climatic change such as increased physical disturbance and rising sea level. An initial descriptive component of my thesis was required to determine the distribution and abundance of a diverse assemblages of sessile non-scleractinian invertebrates, found along gradients of increasing physical disturbance from wave action and increasing intertidal shore height. I also examined the source, intensity and frequency of disturbance along these environmental gradients. This information was used to derive testable hypotheses about the potential impacts of increasing physical disturbance and rising sea level associated with climatic change. I found that physical disturbance had a substantial influence on the types of species that are able to survive in these intertidal rubble habitats, with diversity decreasing at more exposed shores and further down the shore. Physical disturbance was more important for determining the composition of sessile assemblages than other biotic factors such as predation, which only had weak effects on these sessile assemblages. Increased frequency and intensity of waves and storms will increase rates of physical disturbances such as scraping and overturning of rubble plates, which will have a substantial negative impact on biodiversity in these tropical intertidal habitats. These effects may be more complicated that first thought when combined with the effects of rising sea level which will not only alter the extent of inundation, but may also allow wave energy to propagate further up the shore, which has the potential to modify interactions among species through changes to the supply and recruitment of larvae, predator-prey interactions, competition and survival in harsher environments. However, the magnitude of these impacts may depend on how increased physical disturbance and rising sea level affect established species, and whether they will have a substantial effect on larval mortality rates, which currently appears to be limited by physical disturbance. Declines in biodiversity as a result of climatic change over the next 100 years could have important implications for the future health and productivity of coral reef ecosystems, especially given the ecosystem services these organisms provide. A greater understanding of the processes that drive the distribution and abundance of many different types of organisms on coral reefs, and indeed in other ecosystems, will provide essential information that managers can use to better understand and maintain these important ecosystems for future generations.

Dictyostelium discoideum serves as an ideal system to study social evolution because of the social stage of its lifecycle, where individuals aggregate to build a multicellular structure. However, much of its basic biology remains unknown and this limits its utility. I used three separate projects to fill these gaps. In my first project, I examined how speciation and genetic diversity affects kin discrimination using a related dictyostelid, Polysphondylium violaceum . I sequenced the ribosomal DNA of 90 clones of P. violaceum and found that P. violaceum is split into several morphologically identical groups. When allowed to cooperate in pairwise mixes, I found that some clones cooperated with others in their group, but in mixes between groups, clones did not cooperate. For my second project, I looked at whether D. discoideum has sex in natural populations. While sex has been observed in laboratory clones of D. discoideum , it is unclear whether sex occurs in natural populations, and sex can influence the evolution of traits. I used a dataset of microsatellites in 24 clones of D. discoideum to look for a decrease in linkage disequilibrium as a molecular sign of sex. Linkage disequilibrium is higher between physically close loci than between loci on different chromosomes. From this, I conclude that D. discoideum undergoes recombination in nature. Lastly, I used the genome sequence of D. discoideum to look at large scale patterns of evolution. Mutations tend to be biased towards A/T from G/C so, on average, mutations should lower the nucleotide content of sequences. The removal of these mutations, purifying selection, should preserve nucleotide content. I used the genomes of D. discoideum and Plasmodium falciparum identify classes of sequences that should be under different amounts of purifying selection and compared their nucleotide contents. In all cases, those sequences under more purifying selection had higher GC contents than sequences under less purifying selection. Looking at relative nucleotide content may thus serve as an indicator purifying selection. These three studies add insight on how cooperation works in dictyostelids as well as adding an understanding of how traits, social and otherwise, would evolve in this system.

The Rhodophyta (red algae) are an ancient crown group of the Eukarya (ca. 1400-1500 million years), comprised of 5000 - 6000 species. Gametophytes of taxa excluding the speciose Class Florideophyceae are typically of very simple unicellular, filamentous or foliose morphologies. These simple morphologies are often homoplasious (resulting from convergent or parallel evolution) and can be indistinguishable among distinct taxa, leading to cryptic species. As a result, historical morphology-based taxonomy is often not congruent with evolutionary history. Intraspecific genetic variation is not yet characterized for non-Florideophyceae taxa. Here the intraspecific genetic variation was characterized for a locally endemic, morphologically distinct bangiophyte red alga, Bangia maxima Gardner using inter simple sequence repeat (ISSR) patterns from 91 individual filaments across seven local populations. A high degree of genetic variation was observed over very small distances (< 25 cm) and very little genetic exchange was observed between populations. It is possible that B. maxima is a true endemic species and its population dynamics may differ from other Bangia species. Metrics of sequence-based identification rely on genetic divergence among isolates to distinguish taxonomic units independent of morphology. Such metrics are especially useful for morphologically simple or cryptic species. The mitochondrial cytochrome oxidase c subunit 1 gene has been proposed for the Florideophyceae. An evaluation of this gene as a metric for non-Florideophyceae taxa was undertaken and limited utility was demonstrated in most lineages of Rhodophyta due to poor or inconsistent amplification and conflicts with nuclear and plastid phylogenies. Patterns of genetic divergence among taxa are used to infer evolutionary relationships. The nuclear ribosomal small subunit (nSSU rRNA) is the taxonomically broadest pool of gene sequence data for the Rhodophyta. The use of stochastic models of nucleotide evolution is the most common approach to inferring phylogenies using this gene, ignoring much of its evolutionary information as different characters that contribute to secondary structure (e.g. paired nucleotides) are treated independently. The incorporation of structural information leads to more biologically realistic evolutionary models increasing phylogenetic resolution. Parametric models incorporating structural information were used here to more fully resolve phylogenies for all known Rhodophyta lineages. Novel phylogenetic topologies were observed and well supported for each Class within the Rhodophyta resulting in a number of formally proposed or suggested taxonomic revisions. These include phylogenetic resolution of Rhodophyta Classes, support for the introduction of 11 genera within the Bangiales and support for various taxonomic revisions within the Florideophyceae previously proposed but not yet fully adopted. As structure evolves more slowly than its constituent sequence, secondary structure elements can further resolve evolutionary relationships, especially in lineages as old as the Rhodophyta. A novel encoding of secondary structure elements and subsequent multivariate analysis was performed for all known Rhodophyta nSSU rRNA gene sequences, reinforcing phylogenetic results. Computer programs developed for these analyses are publicly available. The analyses presented here significantly advanced understanding of the evolutionary distribution of cryptic species within the Rhodophyta. Furthermore, useful methods for the characterization of such species are presented, as is a demonstration of the utility of biologically realistic sequence models parameterizing nSSU rRNA structure in resolving ambiguous phylogenetic relationships. Most importantly, this work also represents a significant improvement toward taxonomy congruent with evolutionary history for the Rhodophyta.

Gastrotricha are small aquatic invertebrates with approximately 770 known species. The group has a cosmopolitan distribution and is currently classified into two orders, Chaetonotida and Macrodasyida. The gastrotrich fauna of Sweden is poorly known: a couple of years ago only 29 species had been reported. In Paper I, III, and IV, 5 freshwater species new to science are described. In total 56 species have been recorded for the first time in Sweden during the course of this thesis. Common species with a cosmopolitan distribution, e. g. Chaetonotus hystrix and Lepidodermella squamata, as well as rarer species, e. g. Haltidytes crassus, Ichthydium diacanthum and Stylochaeta scirtetica, are reported. In Paper II molecular data is used to infer phylogenetic relationships within the morphologically very diverse marine family Thaumastodermatidae (Macrodasyida). Results give high support for monophyly of Thaumastodermatidae and also the subfamilies Diplodasyinae and Thaumastodermatinae. In Paper III the hypothesis of cryptic speciation is tested in widely distributed freshwater gastrotrichs. Heterolepidoderma ocellatum f. sphagnophilum is raised to species under the name H. acidophilum n. sp. The results indicate that L. squamata may be a complex of at least two species. In Paper III and V the phylogeny of Chaetonotidae (Chaetonotida), the largest family within Gastrotricha, is inferred. The group suffers from a troubled taxonomy and is hypothesized to be nonmonophyletic. Results show that members of Dasydytidae are nested within the group. Since only 3 of 17 sampled genera are monophyletic, it is hypothesized that the cuticular structures used in current classification do not reflect phylogenetic relationships. The phylogenetic hypothesis generated in Paper V indicates a marine origin of the predominantly limnic Chaetonotidae with a subsequent secondary invasion to marine environments of some taxa. / Gastrotricha of Sweden - Biodiversity and Phylogeny

The Rhodophyta (red algae) are an ancient crown group of the Eukarya (ca. 1400-1500 million years), comprised of 5000 - 6000 species. Gametophytes of taxa excluding the speciose Class Florideophyceae are typically of very simple unicellular, filamentous or foliose morphologies. These simple morphologies are often homoplasious (resulting from convergent or parallel evolution) and can be indistinguishable among distinct taxa, leading to cryptic species. As a result, historical morphology-based taxonomy is often not congruent with evolutionary history. Intraspecific genetic variation is not yet characterized for non-Florideophyceae taxa. Here the intraspecific genetic variation was characterized for a locally endemic, morphologically distinct bangiophyte red alga, Bangia maxima Gardner using inter simple sequence repeat (ISSR) patterns from 91 individual filaments across seven local populations. A high degree of genetic variation was observed over very small distances (< 25 cm) and very little genetic exchange was observed between populations. It is possible that B. maxima is a true endemic species and its population dynamics may differ from other Bangia species. Metrics of sequence-based identification rely on genetic divergence among isolates to distinguish taxonomic units independent of morphology. Such metrics are especially useful for morphologically simple or cryptic species. The mitochondrial cytochrome oxidase c subunit 1 gene has been proposed for the Florideophyceae. An evaluation of this gene as a metric for non-Florideophyceae taxa was undertaken and limited utility was demonstrated in most lineages of Rhodophyta due to poor or inconsistent amplification and conflicts with nuclear and plastid phylogenies. Patterns of genetic divergence among taxa are used to infer evolutionary relationships. The nuclear ribosomal small subunit (nSSU rRNA) is the taxonomically broadest pool of gene sequence data for the Rhodophyta. The use of stochastic models of nucleotide evolution is the most common approach to inferring phylogenies using this gene, ignoring much of its evolutionary information as different characters that contribute to secondary structure (e.g. paired nucleotides) are treated independently. The incorporation of structural information leads to more biologically realistic evolutionary models increasing phylogenetic resolution. Parametric models incorporating structural information were used here to more fully resolve phylogenies for all known Rhodophyta lineages. Novel phylogenetic topologies were observed and well supported for each Class within the Rhodophyta resulting in a number of formally proposed or suggested taxonomic revisions. These include phylogenetic resolution of Rhodophyta Classes, support for the introduction of 11 genera within the Bangiales and support for various taxonomic revisions within the Florideophyceae previously proposed but not yet fully adopted. As structure evolves more slowly than its constituent sequence, secondary structure elements can further resolve evolutionary relationships, especially in lineages as old as the Rhodophyta. A novel encoding of secondary structure elements and subsequent multivariate analysis was performed for all known Rhodophyta nSSU rRNA gene sequences, reinforcing phylogenetic results. Computer programs developed for these analyses are publicly available. The analyses presented here significantly advanced understanding of the evolutionary distribution of cryptic species within the Rhodophyta. Furthermore, useful methods for the characterization of such species are presented, as is a demonstration of the utility of biologically realistic sequence models parameterizing nSSU rRNA structure in resolving ambiguous phylogenetic relationships. Most importantly, this work also represents a significant improvement toward taxonomy congruent with evolutionary history for the Rhodophyta.