Population structure, gene flow and dispersal are some of the most commonly estimated population parameters in population genetics, evolutionary biology and conservation genetics. The primary aim of thesis is to test the precision and accuracy of genetic estimates of population structure, gene flow and dispersal. The controlled replicated Drosophila melanogaster experiments of known effective population size (Ne = 14.3) and dispersal rate (m = 0.0025 - 0.04) all adhered to Wright??s demographic island model. Three statistical approaches were empirically tested: 1) the conversion of population structure to gene flow using FST, RST, SHUA and PhiST ; 2) the private alleles method to estimate gene flow; 3) a Bayesian assignment method to estimate dispersal (BAYESASS 1.2). Even in the best-case scenario, almost all current methods except SHUA significantly underestimate population structure, and consequently overestimate gene flow and dispersal when applied to real populations. It was crucial to ensure that the manipulated rate of gene flow was correctly defined. This led to three supporting investigations of hybrid performance, inversion polymorphisms and effective population size. The hybrid performance investigation demonstrated that the manipulated rate of gene flow had not been unexpectedly inflated by hybrid vigour or reduced by breakdown. These experiments also demonstrated that close inbreeding is not a necessary precondition for hybrid vigour or breakdown, which is important for conservation strategies involving induced dispersal. The investigation of inversion polymorphisms ensured that the manipulated rate of gene flow was not affected by selection on inverted regions. The effective population size investigation used a temporal estimation method to confirm that the Ne was accurately predicted by an N:Ne ratio of 0.286. Additionally this experiment showed that the single-sample estimation methods implemented by ONeSAMP or LDNE resulted in downwardly biased estimates of Ne in structured populations. In conclusion these results call into question the confidence that biologists may have in some of the most widely used molecular tools in conservation biology.

Population structure, gene flow and dispersal are some of the most commonly estimated population parameters in population genetics, evolutionary biology and conservation genetics. The primary aim of thesis is to test the precision and accuracy of genetic estimates of population structure, gene flow and dispersal. The controlled replicated Drosophila melanogaster experiments of known effective population size (Ne = 14.3) and dispersal rate (m = 0.0025 - 0.04) all adhered to Wright??s demographic island model. Three statistical approaches were empirically tested: 1) the conversion of population structure to gene flow using FST, RST, SHUA and PhiST ; 2) the private alleles method to estimate gene flow; 3) a Bayesian assignment method to estimate dispersal (BAYESASS 1.2). Even in the best-case scenario, almost all current methods except SHUA significantly underestimate population structure, and consequently overestimate gene flow and dispersal when applied to real populations. It was crucial to ensure that the manipulated rate of gene flow was correctly defined. This led to three supporting investigations of hybrid performance, inversion polymorphisms and effective population size. The hybrid performance investigation demonstrated that the manipulated rate of gene flow had not been unexpectedly inflated by hybrid vigour or reduced by breakdown. These experiments also demonstrated that close inbreeding is not a necessary precondition for hybrid vigour or breakdown, which is important for conservation strategies involving induced dispersal. The investigation of inversion polymorphisms ensured that the manipulated rate of gene flow was not affected by selection on inverted regions. The effective population size investigation used a temporal estimation method to confirm that the Ne was accurately predicted by an N:Ne ratio of 0.286. Additionally this experiment showed that the single-sample estimation methods implemented by ONeSAMP or LDNE resulted in downwardly biased estimates of Ne in structured populations. In conclusion these results call into question the confidence that biologists may have in some of the most widely used molecular tools in conservation biology.

Speciation can be viewed as the formation of reproductive barriers between different populations. This thesis investigates patterns of natural and sexual selection shaping reproductive barriers between two hybridizing flycatchers (i.e. collared – and pied flycatchers). Behaviorally driven sexual isolation depends on both the availability of conspecific mates and on discrimination ability of individuals. My results demonstrate that these two factors may also interact. Discrimination abilities may change in response to the relative frequency of two interbreeding species. The underlying reason appears to be that male pied flycatchers have a song that incorporates more elements of the song characteristics of male collared flycatchers into their own song repertoires when occurring in areas inhabited predominantly by collared flycatchers. I investigated selection pressures acting on hybrids. In migratory species, hybrid fitness might be reduced as a consequence of intermediate suboptimal migration routes (extrinsic post zygotic isolation). Comparison of stable isotope signatures of revealed that parental species have separate wintering grounds, but hybrids appear to winter at the same location as pied flycatchers. A possible dominance effect in the inheritance of migration direction may hence reduce this potential cost. This interpretation is supported by the absence of a reduction in juvenile to adult survival of hybrids. By further comparing male hybrid fitness to that of the parental species, using lifehistory data, I demonstrate that hybrid males experience a moderate reduction in fitness (mainly through a sexually selected disadvantage). Sexual selection acting on male hybrids can play a major role in the speciation process because when the same characters affect assortative mating as well as hybrid fitness, reinforcement of reproductive barriers becomes more likely. Even when reproductive isolation is completed- the fate of newly formed species may be uncertain since they may strongly compete for ecological space. Long-term persistence of ecologically similar, species requires that there are spatial or temporal variation in their relative fitness. The growth of nestling pied flycatchers is less affected by harsh environmental conditions. We suggest that a regional co-existence of the two flycatcher species is due to a lifehistory trade-off between interference competitive ability and robustness to a harsh conditions. Overall, the studies in this thesis reveal the complexity of the interactions between mate choice and competition in shaping sexual signals. Furthermore, it suggests that natural selection is moderate on hybrid males and that sexual selection may have strong implications for the maintenance of species integrity.

Introgression of genes from crops into ruderal populations is a multi-step process requiring sympatry, synchronous flowering, chromosomal compatibility, successful pollination and development of the zygote, germination, establishment and reproduction of hybrid progeny. The goal of this thesis was to generate data on as many steps in this process as possible and integrate them into a predictive statistical model to estimate the likelihood of successful introgression under a range of scenarios. Rape (Brassica napus) and wild turnip (B. rapa var. oleifera) were used as a model system. A homozygous dominant mutation in the rape genome conferring herbicide resistance provided a convenient marker for the study of introgression. Potential differences between wild turnip populations from a wide range of geographic locations in New Zealand were examined. Hand pollination established the genetic compatibility of rape and wild turnip and a high potential for gene introgression from rape to wild turnip. Interspecific hybrids were easily generated using wild turnip as the maternal plant, with some minor differences between wild turnip populations. The frequency of successful hybridisation between the two species was higher on the lower raceme. However, the upper raceme produced more dormant interspecific hybrid seed. Field trials, designed to imitate rare rape crop escapes into the ruderal environment, examined the ability of rare rape plants to pollinate wild turnip plants over four summers. At a ratio of 1 rape plant for every 400 wild turnip plants, the incidence of interspecific hybridisation was consistently low (<0.1 to 2.1 % of total seed on wild turnip plants). There was a significant year effect with the first season producing significantly more seed and a greater frequency of interspecific hybrid progeny than the other years. The frequency of interspecific hybrid progeny increases when the ratio of rape: wild turnip plant numbers increases. The relative importance of anemophily and entomophily in the production of interspecific hybrids was examined. Wild turnip plants produced twice as many seeds with bee pollination relative to wind pollination. However, the frequency of interspecific hybrids under wind pollination was nearly twice that for bee pollination. Light reflectance patterns under UV light revealed a marked difference between wild turnip and rape flowers compared to near identical appearance under visible light. The data indicates that bees are able to distinguish between rape and wild turnip flowers and exhibit floral constancy when foraging among populations with these two species. Hybrid survival in the seed bank, germination and seedling establishment in the field are important components of fitness. Seed banks established in the soil after the field trials described above germinated in subsequent spring seasons. The predominantly brassica weed populations were screened for herbicide resistance and the numbers of interspecific hybrids germinating compared to the original frequency in the field trial results. Frequency of interspecific hybrids was reduced in the populations compared to the original seed deposit. Seed with a known frequency of interspecific hybrid seed was sown in a separate trial, and the frequency of interspecific hybrids compared at 0, 4, 6, and 8 weeks after sowing. Poor germination resulted limited competition between seedlings, however the frequency of interspecific hybrids declined over time indicating low plant fitness. There were no significant population effects on any parameters tested. Interspecific hybrids grown in a glasshouse were backcrossed to the parental species and selfed within the plant and within populations. Pollen from the interspecific hybrids was found to have markedly reduced fertility. Interspecific hybrid plants had low female fertility, with the majority (88%) of the pollinated flowers aborting the siliques. Of the remaining siliques, most (98%) had only one to three seeds per silique. Inheritance of the herbicide resistance gene was regular in backcrosses but highly skewed following self pollination with an excess of herbicide-sensitive progeny. Production of a stochastic predictive model integrated the information acquired over the practical work phase of this thesis and utilised the capabilities of @risk, a new application of a risk analysis tool. The three outputs examined were the number of flowering plants resulting from backcrosses to rape and wild turnip and self pollination of the interspecific hybrid progeny. Five scenarios were modelled and all demonstrated the high likelihood of introgression failure in this system. In all scenarios, >75% of simulations resulted in no interspecific hybrid progeny surviving to flowering in the third generation. In all scenarios, and for all three outputs, the seed set on the interspecific hybrids of the second generation was the major factor that limited the number interspecific hybrid progeny surviving to flowering in the third generation.