Dorken, Marcel E

Hybridisation is an important evolutionary mechanism with diverse outcomes, including the formation of new lineages, the exchange of alleles between species, or their extinction through genetic swamping. In some cases, hybrids exhibit higher fitness than their parental species, which can lead to hybrid invasions that threaten ecosystems. In North America, hybridisation between the native cattail Typha latifolia and the non-native T. angustifolia produces the hybrid T. × glauca, which is a highly impactful invader in wetlands across large areas of southern Canada and northern USA; contrastingly, in Europe and Asia, where its parental species also co-occur, T. × glauca is rare to non-existent. This thesis examines the evolutionary history of T. latifolia and T. angustifolia, which, in addition to being the parental species of T. × glauca, are two of the most globally widespread Typha species, a genus of plants foundational to wetlands. First, we developed genomic resources, including ~12M nuclear SNPs and plastome assemblies to facilitate genetic research on Typha; we also described a cost-effective library preparation and genotyping protocol that makes population genetic studies of freshwater plants accessible. Then, we applied those genomic resources to investigate the roles of drift, selection, and hybridisation in the divergence of T. angustifolia, T. latifolia, and their sister species, T. domingensis and T. shuttleworthii. We found that speciation in these taxa was driven by drift under allopatry, resulting from historical bottlenecks, and that natural selection has played a minimal role in the divergence of these species; additionally, we observed introgressive hybridisation from T. latifolia into T. angustifolia. Finally, we reconstructed the demographic histories of T. angustifolia and T. latifolia from North America and Europe. We observed that reproductive isolation is strong in Europe, where the two species have potentially been sympatric for ~800,000 years, and weak in North America, where they have been sympatric for only a few centuries. Our results exemplify how the divergence and demographic histories of species can correlate with their strength of reproductive isolation. We emphasise that preventing invasions by hybrids will require limiting the movement of Typha and other historically allopatric species, which likely lack reproductive barriers.

Author Keywords: biological invasions, cattails, demographic histories, evolutionary histories, population genomics, reproductive isolation

Diversity of flowers in flowering plants is understood to facilitate mating success, and yet understanding the relationships between plant sexual diversity and mating patterns remains a challenge. In this thesis, I used Sagittaria latifolia, an aquatic plant with unique sexual systems, to investigate relationships between plant sexual diversity, mating patterns, and reproductive success and to understand their evolutionary consequences. First, I measured the magnitude of sexual selection in a dioecious and a monoecious population of S. latifolia, while accounting for size variation. I showed that sexual selection does operate in natural populations of plants. Estimates of sexual selection were similar in both populations, possibly due to the temporal separation of female and male flowering in hermaphroditic S. latifolia, enabling individuals to act temporarily as unisexuals. Second, I examined how seasonal differences in sex allocation and flowering time can affect male mating opportunities, and the effect on reproductive success in hermaphroditic S. latifolia. I found that shifting allocations to be more male-biased with earlier flowering increased male mating opportunities. However, greater mating opportunities did not equate to higher rates of siring, indicating that factors other than temporal overlap contribute to male reproductive success. Finally, I determined the factors affecting mating and reproductive success of males and male-function hermaphrodites, distinguishing between the predictable effects of plant-level traits and population-level features, and unpredictable effects of stochastic factors. Across both populations, we found that plant-level traits had a greater impact on mating and reproductive success than population-level features. Nevertheless, these effects were frequently masked by stochastic factors. Furthermore, unexplained variation in mating and reproductive success may stem from additional plant traits affecting post-pollination gametophyte-sporophyte interactions. My findings indicate that while sexual selection does operate in natural populations of S. latifolia as expected under Bateman's principles, temporal overlap in flowering, plant traits and population features did not explain patterns of male mating or reproductive success, suggesting that post-pollination factors may be influencing mating outcomes.

Author Keywords: Phenology, Plant mating, Plant reproduction, Sagittaria latifolia, Sex allocation, Sexual selection

Natural populations of flowering plants rarely have perfectly uniform distributions, so trends in pollen dispersal should affect the size of the pollination neighbourhood and influence mating opportunities. Here I used spatial analysis to determine the size of the pollination neighbourhood in a stand of the herbaceous, wind-pollinated plant (Typha latifolia; broad-leaved cattail) by evaluating patterns of pollen production and seed set by individual cattail shoots. I found a positive correlation between pollen production and seed set among near-neighbour shoots (i.e., within 4 m2 patches of the stand; Pearson's r = 0.235, p < 0.05, df = 77) that was not driven by a correlation between these variables within inflorescences (Pearson's r = 0.052, p > 0.45, df = 203). I also detected significant spatial autocorrelations in seed set over short distances (up to ~ 5 m) and a significant cross-correlation between pollen production and seed set over distances of < 1 m indicating that the majority of pollination events involve short distances. Patterns of pollen availability were simulated to explore the shape of the pollen dispersal curve. Simulated pollen availability fit actual patterns of seed set only under assumptions of highly restricted pollen dispersal. Together, these findings indicate that even though Typha latifolia produces copious amounts of pollen, the vast majority of pollen dispersal was highly localized to distances of ~ 1 m. Moreover, although Typha latifolia is self-compatible and has been described as largely selfing, my results are more consistent with the importance of pollen transfer between nearby inflorescences. Therefore, realized selfing rates of Typha latifolia should largely depend on the clonal structure of populations.

Author Keywords: clonal structure, correlogram, dispersal curves, pollination, spatial analysis, Typha latifolia

The structure and composition of the landscape can facilitate or impede gene flow, which can have important consequences because genetically isolated groups of individuals may be prone to inbreeding depression and possible extinction. My dissertation examines how landscape structure influences spatial patterns of genetic differentiation and diversity of American marten (Martes americana) and Canada lynx (Lynx canadensis) in Ontario, Canada, and provides methodological advances useful for landscape geneticists. First, I identified the effects of map boundaries on estimates of landscape resistance, and proposed a solution to the bias: a buffer around the map boundary. Second, I assessed the sensitivity of a network-based estimate of genetic distance, conditional genetic distance, to incomplete sampling. I then used these landscape genetic tools in a pairwise, distance-based analysis of 653 martens genotyped at 12 microsatellite loci. I evaluated whether forest management in Ontario has influenced the genetic structure of martens. Although forest management practices had some impact, isolation by distance best described marten gene flow. Our results suggest that managed forests in Ontario are well connected for marten and do not impede marten gene flow. Finally, I used a site-based analysis of 702 lynx genotyped at 14 microsatellite loci to investigate spatial patterns of genetic diversity and differentiation at the trailing (contracting) edge of the lynx distribution in Ontario. I analyzed harvest records and found that the southern edge of lynx range has contracted by >175 km since the 1970s. I also found that neutral genetic diversity decreased towards the trailing edge, whereas genetic differentiation increased. Furthermore, I found strong correlations between gradients of lynx genetic structure and gradients of climate and land cover in Ontario. My findings suggest that increases in winter air temperature, decreases in snow depth, and loss of suitable habitat will result in further loss of genetic diversity in peripheral populations of lynx. Consequently, the adaptive potential of lynx populations on the southern range periphery could decline. In conclusion, my dissertation demonstrates the varying influences that contemporary landscape structure and climate gradients can have on genetic diversity and differentiation of different species.

Author Keywords: Circuitscape, genetic network, landscape genetics, Lynx canadensis, Martes americana, range shift

Sexual reproduction in flowering plants depends on investment in reproduction, the mode of pollen transfer, the availabilities of nutrient resources and potential mates, and the spatial scales over which these processes take place. In this thesis, I studied the general reproductive biology of Shepherdia canadensis (L.) Nutt. (Elaeagnaceae) and the suite of pollinators that visit the plants in Ivvavik National Park, Yukon, Canada. Across ten sites, I found that S. canadensis females were larger than males, but males produced more flowers than females at most sites. Males typically occurred at higher frequencies than females with the average male to female sex ratio being 1.19 ± 0.08 (mean ± SE, n = 10 sites). Both shrub size and flower production were significantly influenced by interactions between soil nitrogen and sex. Insect visitors to S. canadensis flowers were primarily ants and flower flies (Syrphidae), but exclusion experiments indicated that visitation by flying insects yielded greater fruit production than visitation by crawling insects. I found that fruit set was limited by the density of males within populations, but only over small distances (4-6 m). This is the first study to demonstrate that female reproductive success of a generalist-pollinated dioecious plant is limited by the density of males over small spatial scales.

Author Keywords: dioecy, pollinators, sex ratio, sexual dimorphism, Shepherdia canadensis

Population density regulation is a fundamental principle in ecology, however there remain several unknowns regarding the functional expression of density dependence. One prominent view is that the patterns by which density dependence is expressed are largely fixed across a species, irrespective of environmental conditions. Our study investigated the expression of density dependence in Chlamydomonas reinhartti grown under a gradient of nutrient densities, and hypothesized that the relationship between per capita growth rate (pgr) and population density would vary from concave-up to concave-down as nutrients became less limiting. Contrary to prediction, we found that the relationship between a population's pgr and density became increasingly concave-up as nutrient levels increased. Our results suggest that density dependence is strongly variable depending on exogenous and endogenous processes acting on the population, implying that expression of density regulation depends extensively on local conditions. Population growth suppression may be attributable to environments with high intraspecific competition. Additional work should reveal the mechanisms influencing how the expression of density dependence varies across populations through space and time.

Author Keywords: Chlamydomonas reinhartti, density dependence, logistic model, population dynamics, single species growth, theta-logistic equation

Interspecific hybridization rates can vary depending on genomic compatibilities between progenitors, while subsequent hybrid spread can vary depending on hybrid performance and habitat availability for hybrid establishment and persistence. As a result, hybridization rates and hybrid frequencies can vary across regions of parental sympatry. In areas around the Laurentian Great Lakes, Typha × glauca is an invasive plant hybrid of native Typha latifolia and introduced Typha angustifolia. In areas of parental sympatry in Atlantic Canada and outside of North America, T. × glauca has been reported as either rare or non-existent. I investigated whether the low frequencies of hybrids documented in Nova Scotia, Atlantic Canada, are influenced by reproductive barriers that prevent hybrid formation or environmental factors (salinity) that reduce hybrid performance. I identified an abundance of hybrids in the Annapolis Valley (inland) and a scarcity of hybrids in coastal wetlands through preliminary site surveys throughout Nova Scotia. In Annapolis Valley populations, flowering times of progenitor species overlapped, indicating that asynchronous flowering times do not limit hybrid formation in this region. Viable progeny were created from interspecific crosses of T. latifolia and T. angustifolia from Nova Scotia, indicating that there are no genomic barriers to fertilization and germination of hybrid seeds. Typha × glauca germination in high salinity was significantly lower than that of T. latifolia, but there was no difference at lower salinities. Therefore, while germination of hybrid seeds may be impeded in the coastal wetlands where salinity is high, inland sites have lower salinity and thus an environment conducive to hybrid germination. However, I found that once established as seedlings, hybrids appear to have greater performance over T. latifolia across all salinities through higher ramet production. Moreover, I found that T. latifolia sourced from Ontario had reduced germination and lower survivorship in high salinities compared to T. latifolia sourced from Nova Scotia, which could indicate local adaptation by T. latifolia to increased salinity. These findings underline that interactions between environment and local progenitor lineages can influence the viability and the consequent distribution and abundance of hybrids. This, in turn, can help explain why hybrids demonstrate invasiveness in some areas of parental sympatry but remain largely absent from other areas.

Author Keywords: flowering phenology, Hybridization, invasive species, physiology, pollen compatibility, salinity tolerance

The Kentucky coffee tree (Gymnocladus dioicus) is an endangered tree species native to the American Midwest and Southwestern Ontario. Significant habitat loss and fragmentation due to agricultural, industrial and urban development has caused gradual decline across its native range. The aims of this study were to investigate: (1) patterns of genetic diversity and, (2) genetic differentiation (3) relative levels of sexual vs. clonal reproduction, and (4) potential for reduced genetic diversity at range edge for wild G. dioicus populations. An analysis of variation at nine microsatellite loci from populations in the core of the species distribution in the U.S.A. and 4 regions of Southwestern Ontario indicated that G. dioicus has remarkably high genetic similarity across its range (average pairwise FST= 0.05). Germination trials revealed that the seed coats require highly invasive treatments (e.g. 17.93 mol/L H2SO4) to facilitate imbibition, with negligible germination observed in treatments meant to emulate prevailing conditions in natural populations. Low levels of sexual reproduction, high genetic similarity, and habitat degradation are issues that exist across the entire native range of G. dioicus.