Similar forces of selection operating on multiple taxa can cause convergent evolution of quantitative traits. When those same forces of selection occur across an environmental gradient a phenotypic cline may evolve. I have conducted an experiment in central Texas to test whether seed oil melting points and proportions of saturated fatty acids have repeatedly evolved to germinate as predicted by theory. Species with lower seed oil melting points and proportions of saturated fatty acids are expected to germinate at cooler temperatures than ones that germinate at warmer temperatures. Field observations were conducted at two sites for one year to characterize germination temperatures of sixteen species. Gas chromatography was utilized to describe the fatty acid compositions and melting points of those species’ triacylglycerol (oil) stores. The field sites produced conflicting results. At one site, all analyses supported the theory. Whereas evidence from the other site either contradicted expectation or was equivocal. Likewise there was some evidence that plants with annual life histories are under stronger selection to evolve melting points that approximate germination temperature than species with perennial life histories. Finally, evidence was found that a higher level of variation in seed oil melting point and percent of saturated fatty acids is maintained in species that were observed to germinate at low temperatures relative to species observed to germinate at higher temperatures. / text

abstract: Abstract Grammaticalization theory provides a framework for analyzing language change. This thesis uses the concepts relevant to grammaticalization theory in an examination of ‘only’ and ‘just’ to determine how changes in their usage conform to the theory. After an introduction providing a sampling of the myriad ways ‘only’ and ‘just’ are used in Modern English, I provide an overview of grammaticalization theory in Chapter 2. Included in this chapter are a history of the major concepts of grammaticalization theory, an explanation of the commonly-accepted parameters and tools used to test and demonstrate grammaticalization, and a brief discussion of current arguments against grammaticalization theory. Chapter 3 presents my analysis of ‘only’ and ‘just’. I show that ‘only’ has grammaticalized from an adjective to a more grammatical adjective, from an adjective to an adverb, and from an adjective to a complementizer. I then show that ‘just’ has grammaticalized from an adjective to a more grammatical adjective and from an adjective to an adverb. Although some proponents of grammaticalization theory question the grammaticalization pathways leading from adjective to adverb and from adjective to complementizer, I argue that in the cases of ‘only’ and ‘just’ these pathways do indeed exist. I use the Oxford English Dictionary and corpora to support and demonstrate the validity of my argument. / Dissertation/Thesis / Masters Thesis English 2015

Tout observateur peut constater la diversité des milieux sur Terre. La compréhension des liens entre cette diversité des habitats et la biodiversité est l'un des thèmes centraux en Écologie, en Évolution et en Biologie de la conservation. Je m'intéresse dans cette thèse aux conséquences écologiques (à court terme) et évolutives (à plus long terme) de la structuration spatiale et de l'hétérogénéité de l'environnement. Je développe et analyse plusieurs modèles mathématiques, combinant différents formalismes théoriques (dynamiques adaptatives, génétique des populations, génétique quantitative). Ces modèles permettent d'explorer les conséquences de l'hétérogénéité spatiale de l'environnement sur (1) les conditions de persistance des populations ; (2) la coexistence entre différents phénotypes et (3) la dynamique évolutive des populations. Je montre ainsi l'importance (i) de l'intensité et de la place de la migration dans le cycle de vie ; (ii) du type de structure spatiale (explicite ou implicite, continu ou discret) ; (iii) de la forme du compromis évolutif, et donc des coûts d'adaptation à une autre ressource ; et enfin (iv) des éventuels rétrocontrôles démographiques. J'illustre à l'aide des interactions entre hôtes et parasites l'importance des hétérogénéités spatiales. Un premier exemple concerne leur utilisation dans la gestion des pharmacorésistances ou des résistances aux insecticides : une répartition hétérogène du traitement permet d'éviter la propagation de parasites ou de nuisibles résistants. Un second exemple, enfin, illustre comment les hétérogénéités dues à l'auto-structuration spatiale influencent l'évolution de stratégies de défense des hôtes, et permettent l'évolution de défenses altruistes. / Any observer can notice the diversity of habitats on Earth. Understanding the links betweenthis diversity of habitats and biodiversity is a core topic in Ecology, Evolution andConservation Biology.In this thesis, I study the ecological (short-term) and evolutionary (long-term) consequencesof spatial structuring and environmental heterogeneities. I develop and analyzeseveral mathematical models, which combine different theoretical frameworks (adaptivedynamics, population genetics, and quantitative genetics). I explore the consequences ofspatial heterogeneities on (1) the conditions for population persistence; (2) the coexistenceof different phenotypes, and (3) evolutionary dynamics of populations. I show that theresults depend on (i) the life-cycle, and in particular whether migration influences local regulation;(ii) the choice of the spatial structure (explicit or implicit, continuous or discrete);(iii) the shape of the trade-off, and hence the costs of adaptation to another resource, andfinally (iv) the potential demographical feedbacks.I use the specific case of hosts and parasites interactions to illustrate the importanceof spatial heterogeneities. As a first example, I show that a heterogeneous application oftreatments can help prevent the spread of resistant parasites or pests. Secondly, I show howspatial heterogeneities due to self-structuring influence the evolution of host defense strategies,and allow for the evolution of altruistic defense strategies.

Population genetic analyses can provide information about both neutral and selective evolutionary processes shaping genetic variation. In this thesis, extensive population genetic methods were used to make inferences about genetic drift and selection in spruce species. In paper I we studied four species from the Qinghai-Tibetan Plateau (QTP): Picea likiangensis, P. purpurea, P. wilsonii and P. schrenkiana. Big differences in estimates of genetic diversity and Ne were observed in the more restricted species, P. schrenkiana, and the other more widely distributed species. Furthermore, P. purpurea appears to be a hybrid between P. likiangensis and P. wilsonii. In paper II we used Approximate Bayesian Computation (ABC) to find that the data support a drastic reduction of Ne in Taiwan spruce around 300-500 kya, in line with evidence from the pollen records. The split from P. wilsonii was dated to between 4-8 mya, around the time that Taiwan was formed. These analyses relied on a small sample size, and so in Paper III we investigated the impact of small datasets on the power to distinguish between models in ABC. We found that when genetic diversity is low there is little power to distinguish between simple coalescent models and this can determine the number of samples and loci required. In paper IV we studied the relative importance of genetic drift and selection in four spruce species with differing Ne: P. abies, P. glauca, P. jezoensis and P. breweriana. P. breweriana, which has a low Ne, exhibits a low fraction of adaptive substitutions, while P. abies has high Ne and a high fraction of adaptive substitutions. The other two spruce, however, do not support this suggesting other factors a more important. In paper V we find that several SNPs correlate with both a key adaptive trait (budset) and latitude. The expression of one in particular (PoFTL2) correlates with budset and was previously indentified in P. abies. These studies have helped characterise the importance of different population genetic processes in shaping genetic variation in spruce species and has laid some solid groundwork for future studies of spruce.

UA Open Access Publishing Fund / We examined a secondary contact zone between two species of desert tortoise, Gopherus agassizii and G. morafkai. The taxa were isolated from a common ancestor during the formation of the Colorado River (4–8 mya) and are a classic example of allopatric speciation. However, an anomalous population of G. agassizii comes into secondary contact with G. morafkai east of the Colorado River in the Black Mountains of Arizona and provides an opportunity to examine reinforcement of species’ boundaries under natural conditions. We sampled 234 tortoises representing G. agassizii in California (n = 103), G. morafkai in Arizona (n = 78), and 53 individuals of undetermined assignment in the contact zone including and surrounding the Black Mountains. We genotyped individuals for 25 STR loci and determined maternal lineage using mtDNA sequence data. We performed multilocus genetic clustering analyses and used multiple statistical methods to detect levels of hybridization. We tested hypotheses about habitat use between G. agassizii and G. morafkai in the region where they co-occur using habitat suitability models. Gopherus agassizii and G. morafkai maintain independent taxonomic identities likely due to ecological niche partitioning, and the maintenance of the hybrid zone is best described by a geographical selection gradient model.

New Zealand's five endemic mudfish (Neochanna spp.) species have distributions that differ both geographically and by habitat type. Differences in habitat preferences between species have led to the proposal of an evolutionary series within the group. A morphological cline can be observed from the galaxiform Chatham Island and Canterbury species inhabiting lakes and streams, respectively, to the anguilliform Northland and brown mudfishes of ephemeral wetlands. Morphological specializations proposed for wetland dwelling include loss of pelvic fins, reduced eyes, enlarged nostrils, development of caudal flanges, and elongation of dorsal and anal fin bases to become almost confluent with the caudal fin. Another expectation of adaptation to wetland dwelling is specializations in respiratory physiology to obtain oxygen from highly hypoxic or acidic waters, and the ability to cope with seasonal exposure to air during the drought season. Expected respiratory specializations to wetland dwelling include high oxygen affinity haemoglobins, high levels of cooperative oxygen binding, the presence of multiple haemoglobins and the ability to aestivate and survive long periods of emersion. The four mainland Neochanna species were examined to determine if differences in haemoglobin expression as well as differences in haemoglobin oxygen binding correlated with differing habitats and treatments. Whole blood oxygen affinity was determined at several pH levels (6.5, 7.0, 7.5 and 8.0) and temperatures (10'C, 15'C and 20'C), as well as different treatments (aestivating, fasting and control) using a Hemox analyzer. The presence of multiple haemoglobins was determined by isoelectric focusing. All four species displayed high oxygen affinities (p50 = 6.5 to 9.5 mm Hg at pH 7.5 15'C), moderate levels of cooperativity (Hill coefficients = 1.75 to 2.00 at pH 7.5 15'C), pH sensitivity (Bohr coefficients = -0.62 to -0.94 between pH 7.5 and 7.0 at 15'C), temperature sensitivity (ΔH = -2.20 to -15.78 k cal mol-1 between 10'C and 15'C) and the presence of multiple haemoglobins. Black, brown and Northland mudfish were able to survive aestivation for six weeks but there were no changes between air-breathing and water-breathing individuals with respect to oxygen binding characteristics. Although there is evidence of habitat specialization in haemoglobin physiology between mudfish species, differences between species did not correlate with the evolutionary series proposed for specialization to dwelling in ephemeral wetlands and latitudinal distributions of mudfish species appear to strongly dictate oxygen binding properties of mudfish whole blood.

Doctor of Philosophy / Department of Biology / Mark Ungerer / Elucidating the molecular basis of adaptive phenotypic variation represents a central aim in evolutionary biology. Using the model plant species Arabidopsis thaliana, I studied the intra-specific variation in freezing tolerance among natural accessions across its native range. Considerable variation in freezing tolerance among 71 selected accessions was observed both with and without a prior cold acclimation treatment, suggesting that both differences in cold-acclimation capacity and in intrinsic physiology contribute to this variation. A highly significant positive relationship was observed between freezing tolerance and latitude of origin of these accessions. This clinal pattern of variation is found to be attributable, at least in part, to relaxed purifying selection on CBF/DREB1 genes in the species’ southern range. These CBF/DREB1 genes encode transcriptional activators that play a critical role in the ability of A. thaliana plants to undergo cold acclimation and thereby achieve maximum freezing tolerance. Relative to accessions from northern regions, accessions of A. thaliana from the southern part of their geographic range exhibit significantly higher levels of nonsynonymous polymorphisms in coding regions of CBF/DREB1 genes. Relaxed selection on the CBF/DREB1s in southern accessions also has resulted in mutations in regulatory regions that lead to abrogated expression. These mutations in coding and regulatory regions compromise the function of CBF/DREB1 transcriptional activators during the cold acclimation process, as determined by reductions in rates of induction and maximum levels of expression in the downstream genes they regulate. These mutations could be selective neutral or beneficial in southern accessions depending on whether there is an allocation cost associated with cold acclimation. The fitness benefit and possible allocation cost of cold acclimation was examined in freezing and freezing-free environments using natural accessions exhibiting contrasting abilities of cold acclimation as well as transgenic CBF gene over-expression or knockdown/knockout lines. The extent to which cold acclimation benefits the plant in presence of freezing temperature is revealed, but a cost of cold acclimation wasn’t detected in the absence of freezing temperature under our experimental design, which suggests that these mutations in CBF genes in southern accessions might be neutral to natural selection.

This thesis deals with the relationship between the place of origin and the extend of cold hardiness of fire bug Pyrrhocoris apterus (Linnaeus, 1758). Thirteen populations from twelve European localities were used in the experiment. Linear regressions were used to relate the values of SCP (10th percentile, first quartile, mean and median) of the individual populations to minimum and average winter temperatures of given localities. The relationship between air temperatures and SCP values were significant provided that the data of all populations were used. After the removal of three southern non-diapausing populations this relationship lost its significance.

No description available.

京都大学 / 新制・課程博士 / 博士(人間・環境学) / 甲第23263号 / 人博第978号 / 新制||人||232(附属図書館) / 2020||人博||978(吉田南総合図書館) / 京都大学大学院人間・環境学研究科相関環境学専攻 / (主査)教授 瀬戸口 浩彰, 教授 加藤 眞, 教授 市岡 孝朗 / 学位規則第4条第1項該当 / Doctor of Human and Environmental Studies / Kyoto University / DFAM