The ant genus Lasius (Formicinae) arose during the early Tertiary approximately 65 million years ago. Lasius is one of the most abundant and widely distributed ant genera in the Holarctic region, with 95 described species placed in six subgenera: Acanthomyops, Austrolasius, Cautolasius, Chthonolasius, Dendrolasius and Lasius. Many species of Lasius have been central to numerous species-level studies and the focus of many ecological, agricultural, and behavioral investigations. The focus of this study was to use molecular phylogenetic analysis of 781 base pairs of the mitochondrial gene cytochrome oxidase I (COI) and 251 base pairs of an anonymous nuclear gene (ANG) to address questions about the evolutionary relationships of North American Lasius species and subgenera. These relationships were used to better understand the biological and evolutionary complexities associated with these species given their North American distributions. The resulting hypotheses generated in this study from the analyses of these genes produced unexpected patterns of phylogenetic placement of Lasius species and subgenera. A number of biological processes alone or together could explain these patterns, including interspecific hybridization and gene introgression, incomplete lineage sorting, and the presence of multiple cryptic species.

Widespread species that occupy continents and oceanic islands provide an excellent opportunity to study evolutionary forces responsible for population divergence. Here, I use multilocus coalescent based population genetic and phylogenetic methods to infer the evolutionary history of the common chaffinch (Fringilla coelebs), a widespread Palearctic passerine species. My results showed strong population structure between Atlantic islands. However, the two European subspecies can be considered one panmictic population based on gene flow estimates. My investigation of effects of sampling on concatenated and Bayesian estimation of species tree (BEST) methods demonstrated that concatenation is more sensitive to sampling than BEST. Furthermore, concatenation can provide incorrect evolutionary relationships with high confidence when sample size is small. In conclusion, my results suggest European ancestry for the common chaffinch and Atlantic islands appear to have been colonized sequentially from north to south via Azores.

Widespread species that occupy continents and oceanic islands provide an excellent opportunity to study evolutionary forces responsible for population divergence. Here, I use multilocus coalescent based population genetic and phylogenetic methods to infer the evolutionary history of the common chaffinch (Fringilla coelebs), a widespread Palearctic passerine species. My results showed strong population structure between Atlantic islands. However, the two European subspecies can be considered one panmictic population based on gene flow estimates. My investigation of effects of sampling on concatenated and Bayesian estimation of species tree (BEST) methods demonstrated that concatenation is more sensitive to sampling than BEST. Furthermore, concatenation can provide incorrect evolutionary relationships with high confidence when sample size is small. In conclusion, my results suggest European ancestry for the common chaffinch and Atlantic islands appear to have been colonized sequentially from north to south via Azores.

Once thought to be an evolutionary dead-end, hybridization is now being detected in genomes across kingdoms, perhaps even playing an integral role in evolution. In chapter 1, I investigated the potential influences of hybridization on the evolution of a group of vagrant, asexual species in the Rhizoplaca melanophthalma species group. I sequenced the mitochondrial and nuclear genomes of 55 specimens and found well-supported nuclear phylogenies of both datasets. There were, however, multiple instances of discordance between the mitochondrial and nuclear trees, which can be caused by hybridization. PhyloNet and ABBA-BABA also detected widespread hybridization among this group. In chapter 2, I shifted to the Holarctic clade of lichen-forming fungi in Xanthoparmelia to characterize gene tree conflict and investigate the potential for hybrization. Here, I used three different tests for hybridization that account for incomplete lineage sorting "ABBA-BABA, PhyloNet, and SplitsTree" as well as PhyParts to characterize gene tree conflict. Like the Rhizoplaca species group, widespread hybridization was detected in the Holarctic clade despite robust phylogenies. My research underscores the value of investigating hybridization when studying species boundaries and evolutionary history.

This thesis investigates a range of genealogical approaches to making quantitative inferences about the spatial and demographic history of populations with application to two insect systems: A local radiation of high alpine ground beetles (Carabidae) in the genus Trechus and major refugial populations of the oak gall parasitoid Cecidostiba fungosa (Pteromalidae). i) Summary statistics, which make explicit use of genealogical information are developed. Using simulations their power to detect a history of population growth is shown to be higher than that of standard measures such as Tajima’s D for single and multilocus data. The improvement arises from the fact that in contrast to pairwise measures, the new statistics are minimally confounded with the topology. ii) A Bayesian method to reconstructing character states is used to infer the Pleistocene history of populations of high alpine Trechus sampled along a singlemountain range frommitochondrial and nuclear data. Despite evidence for some incomplete lineage sorting, a simple model of a series of extreme founder events out of two refugia during or before the last glacial maximum provides a good fit to the data. iii) A large set of exon-primed, intron-spanning (EPIC) loci is developed for Hymenoptera from EST and genomic data. Amplification success is screened on a range of Hymenopteran species associated with two insect-plant interactions: Oak galls and figs. iv) Borrowing model-based approaches developed to quantify species divergence, the new EPIC loci are used to investigate the relationships between three major European refugia in the oak gall parasitoid C. fungosa. These analyses reveal strong support for an eastern origin, effective ancestral population sizes comparable to insect model species and evidence for recent population divergence during the last interglacial. The results also suggest that there is significant information in minimal samples provided a large number of loci are available. v) Results for the probability of gene tree topologies are derived for a model of divergence with gene flow between three populations. I outline how the asymmetries in the frequency of gene tree topologies may be used to distinguish incomplete lineage sorting from migration and discuss the results in the context of next generation sequence data from D. melanogaster and humans and Neanderthals.

Despite more than 50 years of research on the genetic basis of speciation, we still know very little about the early stages of this process. A general rule of speciation is the Haldane’s Rule, which states that postzygotic isolation is expressed earlier in the heterogametic sex: sterility and inviability in hybrids is much more likely to occur in the heterogametic sex. However, it is still unknown how long hybrid sterility takes to be established in the heterogametic sex, since most studies have focused on highly divergent species that no longer hybridize in nature. The meadow grasshopper (Chorthippus parallelus) is a suitable system to shed light on this matter because hybrid male sterility exists between subspecies that show low divergence and are still able to hybridize readily in nature. C. p. parallelus is distributed throughout Northern, Central and Eastern Europe while C. p. erythropus is restricted to the Iberian Peninsula. Both subspecies meet in the Pyrenees, where they form a narrow hybrid zone. Yet, it is unknown 1) how long ago have these subspecies diverged and 2) how have they expanded to form the hybrid zone where these incompatibilities take place. Here, we applied phylogenomic methods to estimate the time of divergence of the subspecies and to study its phylogeographic history. Using the mitogenome and known mitochondrial rates of evolution, we dated the diversification of the subspecies ≈100,000 years ago and found six ancient mitochondrial haplotypes. Implementing coalescent theory to estimate a nuclear species tree, we found evidence of sub-refugia within two or three main refugia and that the hybrid zone originated from a recent post-glacial expansion from one sub-refugia. Haldane’s rule evolved rapidly in C. parallelus, within ≈100,000 years, with demographic processes, such as population expansion. Founder effect reduced genetic diversity in C. parallelus parallelus, with the possibility of fixing incompatible alleles with C. parallelus erythropus resulting in hybrid male sterility.

<p>DNA sequences from the <i>rps16</i> intron and the <i>psbE-petL</i> spacer from the chloroplast genome, the ribosomal nuclear ITS region, and introns from the low copy nuclear genes <i>RPA2</i>, <i>RPB2</i>, <i>RPD2a</i> and <i>RPD2b</i>, are in different combinations used to infer phylogenetic relationships in <i>Sileneae</i> (<i>Caryophyllaceae</i>). Used in concert, the biparentally inherited nuclear regions are useful to distinguish between paralogy due to allopolyploidy and single gene duplications, respectively, because the latter are not expected to give rise to repeated phylogenetic patterns in potentially unlinked sequence regions. In addition, the sequences resolve previously poorly known relationships in the tribe <i>Sileneae</i>. Several independent losses and incomplete concerted evolution are inferred between the two <i>RPD2</i> paralogues in a subgroup of <i>Silene</i>.</p><p>An allopolyploid origin is suggested for the tetraploid <i>S. aegaea</i>, with the maternal ancestor from the diploid <i>S. pentelica</i> lineage, and the paternal contributor from the diploid <i>S. sedoides</i> lineage.</p><p><i>Silene involucrata</i> originated as an allotetraploid with the diploid lineage of Arctic <i>S. uralensis</i> as cytoplasmic donor and the diploid Siberian/Northeast Asian <i>S. ajanensis</i> lineage as pollen donor. A subsequent allopolyploidization with the <i>S. ajanensis</i> lineage as pollen donor and the tetraploid <i>S. involucrata</i> lineage as cytoplasmic donor resulted in the hexaploid lineage of <i>S. sorensenis sensu lato</i>.</p><p>A monophyletic origin of the North American polyploids is rejected. One lineage consists of tetraploid <i>S. menziesii</i> and its diploid allies. A separate lineage leads to a clade consisting of both diploid and polyploid Arctic, European and Asian taxa in addition to the majority of the North American polyploids. The tetraploid <i>S. californica</i> and the hexaploid <i>S. hookeri</i> are derived from separate allopolyploidization events between these two lineages.</p>

DNA sequences from the rps16 intron and the psbE-petL spacer from the chloroplast genome, the ribosomal nuclear ITS region, and introns from the low copy nuclear genes RPA2, RPB2, RPD2a and RPD2b, are in different combinations used to infer phylogenetic relationships in Sileneae (Caryophyllaceae). Used in concert, the biparentally inherited nuclear regions are useful to distinguish between paralogy due to allopolyploidy and single gene duplications, respectively, because the latter are not expected to give rise to repeated phylogenetic patterns in potentially unlinked sequence regions. In addition, the sequences resolve previously poorly known relationships in the tribe Sileneae. Several independent losses and incomplete concerted evolution are inferred between the two RPD2 paralogues in a subgroup of Silene. An allopolyploid origin is suggested for the tetraploid S. aegaea, with the maternal ancestor from the diploid S. pentelica lineage, and the paternal contributor from the diploid S. sedoides lineage. Silene involucrata originated as an allotetraploid with the diploid lineage of Arctic S. uralensis as cytoplasmic donor and the diploid Siberian/Northeast Asian S. ajanensis lineage as pollen donor. A subsequent allopolyploidization with the S. ajanensis lineage as pollen donor and the tetraploid S. involucrata lineage as cytoplasmic donor resulted in the hexaploid lineage of S. sorensenis sensu lato. A monophyletic origin of the North American polyploids is rejected. One lineage consists of tetraploid S. menziesii and its diploid allies. A separate lineage leads to a clade consisting of both diploid and polyploid Arctic, European and Asian taxa in addition to the majority of the North American polyploids. The tetraploid S. californica and the hexaploid S. hookeri are derived from separate allopolyploidization events between these two lineages.

Abstract Both demographic histories and natural selection complicate the speciation process. There is a need to jointly study the effects of natural selection on so called magic traits that can cause reproductive isolation such as climatic adaptation, and its interaction with neutral demographic histories. Closely related incipient coniferous species offer us a great system for this effort. I used genetic variation at one set of climate-related candidate genes and another set of reference loci and cytoplasmic genomic fragments of two closely related parapatric pine species: Pinus massoniana Lamb. and Pinus hwangshanensis Hisa. Population genetic analyses were used to measure genetic variation and detect signals of ancient and recent selection. Speciation parameters including migration rates and divergence times at candidate genes and reference loci were compared under the Isolation with migration model. Hierarchical Approximate Bayesian Computation (ABC) was used to define demographic and speciation models. Intra- and interspecific genetic variation at cytoplasmic and nuclear intronic sequences were compared between parapatric populations and allopatric populations to distinguish the effects of introgression and incomplete lineage sorting in generating shared genetic variation between the species. The results showed that ancient selection were shared by the lineages leading to the species while recent selection has been species-specific. Candidate genes had significant lower migration rates compared to reference loci. Recent differential climatic selection might counteract against gene flow at underlying genes, which therefore favors divergence between the two pines through ecological speciation. Shared mitotypes were randomly distributed across species’ ranges, which therefore supported the incomplete lineage sorting hypothesis, but the shared nuclear intronic variation distributed more frequently in parapatric populations than in allopatric populations, supported the introgression hypothesis. ABC and species’ distribution modeling also supported the secondary gene flow model. The three genomes had different rates of mutation and gene flow might mirror different phases of the speciation continuum. The results in this thesis are valuable for understanding evolution in general and for other applied purposes such as tree breeding and climate change adaptation. / Tiivistelmä Luonnonvalinta ja populaatioiden historian demografia tekevät lajiutumisesta monimutkaisen tapahtumaketjun. Luonnonvalinnan ja demografisten tekijöiden vuorovaikutusta on paras tutkia samanaikaisesti, kun tarkastellaan lajiutumiseen vaikuttavia ominaisuuksia. Tällaisia ovat esimerkiksi ilmastoon sopeutumiseen liittyvät ominaisuudet. Lähisukuiset havupuulajit tarjoavat erinomaiset mahdollisuudet tähän työhön. Tutkin geneettistä muuntelua yhtäältä ilmastosopeutumiseen liittyvissä ns. ehdokasgeeneissä ja toisaalta neutraaleiksi oletetuissa verrokkigeeneissä sekä sytoplasman genomeissa kahdessa lähisukuisessa mäntylajissa Pinus massoniana Lamb. ja Pinus hwangshanensis Hisa, joiden populaatiot esiintyvät joskus erillään toisistaan (allopatrisesti), toisinaan vierekkäin (parapatrisesti). Mittasin muuntelun määrää ja etsin merkkejä valinnan vaikutuksesta. Vertasin erilaisia lajiutumismallien parametrejä verrokki- ja ehdokasgeeneissä. Käytin simulaatioita etsiäkseni parhaat demografiset ja lajiutumiseen liittyvät mallit. Vertasin kloroplastien ja mitokondrioiden genomien sekvenssien lajinsisäistä ja lajien välistä muuntelua allopatrisissa ja parapatrisissa populaatioissa tutkiakseni onko lajien yhteinen muuntelu seurausta siitä että lajien eriytymisestä on kulunut vain vähän aikaa vai siitä että sen jälkeen on tapahtunut geenivirtaa. Kauan sitten tapahtunut valinta on vaikuttanut samalla tavalla kumpaankin lajiin, osin koska tutkimus kohdistui myös niiden yhteiseen edeltäjälinjaan. Äskettäinen valinta taas oli suuremmassa määrin kummallekin lajille ominaista. Viime aikojen ilmastoon liittyvä valinta on voinut vähentää geenivirtaa ehdokasgeeneissä, mikä voisi edistää ekologista lajiutumista. Tuman DNA:n muuntelu jakautuminen tuki sitä mahdollisuutta että lajien yhteinen geneettinen muuntelu johtuu äskettäisestä geenivirrasta, ei vain siitä että lajiutuminen on niin varhaisessa vaiheessa. Mitokondrioiden geeneissä lajeilla yhtä paljon yhteistä muuntelua sekä allopatrisissa että parapatrisissa populaatioissa, mikä tukee sen sijaan eriytymisen jälkeistä epätäydellistä muuntelun erilaistumista. Eri genomit heijastavat lajiutumisprosessin eri vaiheita. Väitöskirjan tulokset ovat osaltaan tuottaneet uutta tietoa lajiutumisesta ja valinnasta. Lisäksi niillä on merkitystä ilmastomuutoksen vaikutusten ymmärtämisessä ja metsänjalostuksessa.

Phylogenetics is the research of ancestor-descendant relationships among different groups of organisms, for example, species or populations of interest. The datasets involved are usually sequence alignments of various subsets of taxa for various genes. A major task of phylogenetics is often to combine estimated gene trees from many loci sampled from the genes into an overall estimate species tree topology. Eventually, one can construct the tree of life that depicts the ancestor-descendant relationships for all known species around the world. If there is missing data or incomplete sampling in the datasets, then supertree methods can be used to assemble gene trees with different subsets of taxa into an estimated overall species tree topology. In this study, we assume that gene tree discordance is solely due to incomplete lineage sorting under the multispecies coalescent model (Degnan and Rosenberg, 2009). If there is missing data or incomplete sampling in the datasets, then supertree methods can be used to assemble gene trees with different subsets of taxa into an estimated species tree topology. In addition, we examine the performance of the most commonly used supertree method (Wilkinson et al., 2009), namely matrix representation with parsimony (MRP), to explore its statistical properties in this setting. In particular, we show that MRP is not statistically consistent. That is, an estimated species tree topology other than the true species tree topology is more likely to be returned by MRP as the number of gene trees increases. For some situations, using longer branch lengths, randomly deleting taxa or even introducing mutation can improve the performance of MRP so that the matching species tree topology is recovered more often. In conclusion, MRP is a supertree method that is able to handle large amounts of conflict in the input gene trees. However, MRP is not statistically consistent, when using gene trees arise from the multispecies coalescent model to estimate species trees.