The hybridizing Daphnia mendotae and D. dentifera (Crustacea: Cladocera) are sympatric throughout much of North America, and are considered a cryptic species complex due to their lack of phylogenetically informative morphological characters. They appear to have no biological mating barriers: hybrids may dominate or coexist with either or both parental species, and are sexually competent, forming both F2 hybrids and back-crosses. Nevertheless, the two species remain distinct. There is observational evidence that separation may be enforced by adaptation to different predation regimes: D. mendotae, with its greater anti-predator morphological plasticity, may out-compete D. dentifera under intense invertebrate predation, while the smaller D. dentifera may be better adapted to avoid predation by visually feeding fish. We tested this idea by examining whether D. mendotae and D. dentifera differ in ecologically relevant life history and morphological traits. We performed a replicated life history experiment involving 6 replicates of 6 clones within each species, and measured time until first reproduction, fecundity, and juvenile and population growth rates. In parallel, we examined whether these species differed in morphological traits predicted to arise from adaptation to different predator types, and tested the ecological relevance of these traits by exposing Daphnia to predation by the invasive cladoceran Bythotrephes longimanus. Finally, we examined the plasticity of life history, morphology, and susceptibility to predation by rearing Daphnia under exposure to Bythotrephes chemical cues for two generations. D. mendotae and D. dentifera differed across almost all measured life history and morphological traits in directions that accord with our hypotheses, strongly suggesting that their species boundaries are maintained by adaptation to different predation regimes. Plastic reaction to Bythotrephes, however, was weak and inconsistent, suggesting that these species either do not detect or respond to this recent invader, or that their responses are manifested in other ways. / Thesis (Master, Biology) -- Queen's University, 2011-12-23 20:31:14.76

Evidence for the repeated, independent evolution of hot- and cool-season breeding populations across colonies of the Band-rumped Storm-petrel has garnered much interest from seabird ecologists and evolutionary biologists, alike. Sympatric seasonal populations have been shown to be each other’s closest relatives, implying that ecological divergence into two seasonal foraging types has taken place several times independently across the species’ range, each time in the absence of geographical barriers to gene flow. I present data on the molecular genetic characteristics of a unique year-round breeding population in the Cape Verde archipelago. Using mitochondrial control region sequence and five microsatellite loci, I characterized genetic relationships among 220 birds breeding in four different months (January, April, June and November) and tested for a relationship between temporal isolation and genetic differentiation. Birds breeding in April, June and November were genetically indistinguishable at microsatellite loci, but control region sequence suggested differentiation between January and other months (pairwise ΦST from 0.19 to 0.46, p≤0.05). No evidence for genetic isolation by time was found. A comparison of birds breeding in June and November revealed significant differences in a suite of morphological traits related to foraging strategy (tarsus length, bill length, bill depth, head length, wing length and tail shape), and even significant variation among birds breeding in the same month, despite evidence for gene flow between these groups. Interpreting these patterns in the context of Band-rumped Storm-petrel populations range-wide, I suggest that divergent selection on standing variation within ancestral populations may be an important mechanism explaining the repeated, independent evolution of conserved seasonal foraging types in this species. / Thesis (Master, Biology) -- Queen's University, 2011-02-28 12:02:44.256

The creation of biodiversity involves the evolution of new species. Recent trends in the study of speciation have increased the emphasis on the role of ecology in adaptation and the evolution of reproductive isolation. This includes examining the relative contributions of different types of selection, the role of gene flow and the genomic changes that occur during ecological speciation. The search for speciation genes continues, however our growing knowledge of how the genome translates into phenotypes means we should now consider a broader molecular basis of speciation, which includes genetic, transcriptomic and potentially epigenetic variation that contribute to phenotypic variation. This thesis addresses the molecular basis of speciation by using three different complementary methods to examine the early stages of ecological speciation and the evolution of premating reproductive isolation between two incipient species of the cactophilic fly, Drosophila mojavensis. First, the genetic basis was examined through the sequencing of two candidate genes underlying reproductive isolation (Chapter 2). Second, the historical biogeography of population divergence was uncovered using multiple sequenced loci (Chapter 3). Lastly, gene expression across the whole transcriptome associated with phenotypic plasticity and mating success was assessed (Chapter 4). Further, the role of epigenetic imprinting in the population divergence of a freshwater fish, Girardinichthys multiradiatus, was examined through sequencing of a well known gene involved in sexual conflict (Chapter 5). These studies find that uncovering the genetic variation underlying speciation is difficult, especially when there is extensive phenotypic plasticity. Further, gene expression plasticity may play an important role in the evolution of premating isolation, and this includes a role for epigenetic mechanisms of gene expression. Additionally, it is important to assess the demographic scenario of population divergence to put into context the ecological and functional data on divergent groups. Through these studies this thesis examines the genetic, expression and epigenetic variation associated with on-going population divergence, and emphasises the need to consider the potential role of the full range of gene expression changes during ecological speciation.

It is now widely accepted that adaptation to different ecological niches can result in the evolution of new species. However, when gene flow persists speciation must overcome the antagonism between selection and recombination: Specifically, if gene flow persists, recombination will break down the genetic associations between alleles that characterise emerging species and cause reproductive isolation. Accordingly, genetic architectures that impede recombination can slow the breakdown of linkage disequilibrium and facilitate speciation. Mimicry in tropical butterflies has long been championed as an example of adaptation driving speciation. In the Neotropical genus Heliconius, distantly related pairs of unpalatable species often converge on the same bright warning-pattern to more efficiently advertise their distastefulness to predators. In contrast, closely related taxa often belong to different mimicry rings. The sister species, Heliconius melpomene and H. cydno are sympatric across much of Central and northern South America. Using artificial butterflies I reveal selection against non-mimetic hybrid colour patterns between these two species. These colour patterns are also used as mating cues and mimetic shifts may cause both pre-mating and post-mating isolation. However, shifts in colour pattern cannot drive reproductive isolation alone; rather, they must be accompanied by corresponding mate preferences. Associations between trait and preference loci may be broken down by mating and subsequent recombination. I demonstrate a genetic linkage between loci for both male and female mate preference and wing colour pattern in Heliconius cydno and H. melpomene. In addition, I present evidence for further associations between alleles affecting hybrid sterility and host-plant use and colour pattern loci. All this implies that linkage between traits that contribute to reproductive and ecological isolation is a general phenomenon in Heliconius with an underlying adaptive basis. Overall these results expose a genetic mechanism that, by impeding recombination, can facilitate speciation in the face of gene flow.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第17774号 / 理博第3897号 / 新制||理||1562(附属図書館) / 30581 / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 渡辺 勝敏, 教授 曽田 貞滋, 教授 疋田 努 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第20208号 / 理博第4293号 / 新制||理||1616(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 渡辺 勝敏, 教授 沼田 英治, 教授 曽田 貞滋 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

No description available.

Parasitic wasps are highly diverse and play a major role in suppression of herbivorous pest populations, but relatively little is known of the mechanisms driving their diversity. Molecular studies indicate that cryptic species complexes resulting from adaptations to specific hosts or host-foodplants may be common. The gregarious endoparasitoid, Cotesia congregata (Braconidae), is a model system for understanding parasitic wasp biology. It is reported to attack at least 15 species of sphingid caterpillars, most of which are plant family specialists. Molecular studies have demonstrated genetic differentiation of two host-foodplant complex sources originating from Manduca sexta on tobacco (MsT) and Ceratomia catalpae on catalpa (CcC). Response to female pheromone and elements of their courtship songs differ. Wasps from both sources mated and produced F1 hybrid offspring in the laboratory; however, 90% of hybrid females resulting from one of the reciprocal crosses failed to produce offspring. I built on this previous work by evaluating an ecological barrier, the evolution of courtship songs within the genus, and patterns of hybrid sterility among four additional host-foodplant complexes, as well as differentiation of their symbiotic bracovirus. Tests of developmental tolerance to nicotine demonstrate that MsT wasps are highly adapted to hosts feeding on tobacco, whereas CcC wasps experience high mortality. Acoustic analysis of courtship songs among host-foodplant sources of C. congregata and eleven additional species of Cotesia demonstrates that songs are species specific and appear to be correlated with genetic relatedness. Cotesia congregata from all sources mated and produced F1 hybrid offspring in the laboratory; however, hybrid females resulting from specific reciprocal crosses failed to produce progeny. Dissections of hybrid females revealed that sterile wasps lacked mature ovaries and functional bracovirus, a symbiotic virus integrated into the wasp genome and necessary to suppress the host immune system. Relative in vivo expression of wasp bracovirus genes differs between MsT and CcC host-foodplant complexes. Cumulatively, these behavioral, ecological, and genetic barriers to reproduction indicate that C. congregata is diverged into two incipient species with limited gene flow, and provides insight into the role of varied reproductive barriers in speciation of parasitic wasps.

This thesis investigates the ecological and evolutionary implications of a recent secondary contact between two closely related bird species: collared (Ficedula albicollis) and pied (F. hypoleuca) flycatchers. Collared flycatchers started to colonize the Swedish island of Öland, where pied flycatchers were already present, in the late 1950s-early1960s. My major aims were to investigate which factors are acting against versus for long-term coexistence between the two species. Specifically, I investigated the relative importance of allopatric divergence, interspecific competition, hybridization and learning in promoting or inhibiting coexistence. The combined effects of interspecific competition and hybridization drives pied flycatchers towards local extinction in their preferred deciduous habitat. However, my results also show that pied flycatchers are better able to tolerate harsh environmental conditions. This trade-off between competitive ability and resilience in the face of harsh conditions facilitates a regional coexistence between the species. Coexistence is furthermore favoured by competition-mediated divergence in breeding habitat choice, timing of breeding and male breeding plumage colouration. Due to interspecific competition, male pied flycatchers are forced to breed in a more mixed forest type with a later peak in food abundance, which is accompanied by a divergence in breeding time between the two species. In areas shared with collared flycatchers, male pied flycatchers with brown plumage coloration, most divergent from that of collared flycatchers, are favoured by selection. In addition to facilitating coexistence, the observed shift in habitat occupancy increases reproductive isolation between the two species. By using cross-fostering experiments I demonstrate that natal habitat imprinting has the potential to additionally speed up habitat segregation. Finally I show that hybrid nestlings express an intermediate response to harsh environments, indicating that another aspect of ecological-based selection may be important in reproductive isolation between the species. In summary, my results show that adaptations during historic allopatry are important both in facilitating coexistence as well as in providing a foundation for further ecological divergence at secondary contact. This is of relevance today as many species are shifting their distributions in response to habitat disturbance and global warming.

Mainly because of the problems with species delineation, the biogeography of microscopic organisms is notoriously difficult to elucidate. In this contribution, variable nuclear and mitochondrial DNA markers were sequenced from individual specimens representing the Echiniscus virginicus complex that are morphologically indistinguishable under light microscopy (five populations from the temperate Eastern Nearctic and 13 populations from the subtropical and tropical zone). A range of methods was used to dissect components of variability within the complex (Bayesian inference, haplotype networks, Poisson tree processes, automatic barcode gap discovery delineations, principal components analysis and ANOVA). We found deep divergence between the temperate Eastern Nearctic E. virginicus and pantropical Echiniscus lineatus in all three genetic markers. In contrast, intraspecific genetic variation was very low, regardless of the geographical distance between the populations. Moreover, for the first time, statistical predictions of tardigrade geographical distributions were modelled. The factor determining the allopatric geographical ranges of deceptively similar species analysed in this study is most likely to be the type of climate. Our study shows that widespread tardigrade species exist, and both geographical distribution modelling and the genetic structure of populations of the pantropical E. lineatus suggest wind-mediated (aeolian) passive long-distance dispersal.