Traditionally, the study of biodiversity has focused on quantifying patterns of species diversity, or species richness, by simply counting the number of species across environmental gradients. This approach has been fundamental to ecological investigations and thinking with regards to identifying patterns of biodiversity. Although species diversity is the most commonly used dimension of biodiversity, species diversity alone does not provide a mechanistic understanding of biodiversity gradients. By also quantifying the genetic and phylogenetic diversity of a population, community or ecosystem, ecologists can become more informed on the relationships organisms have with one another, as well as their potential to adapt to changes in their environment. While each of these approaches provides methods for characterizing biodiversity, they do not offer direct insight into what species do, how they function, or how they will respond to changes in their environment. Functional, or trait-based ecology, provides an informative alternative to species-centric approaches that seeks to understand patterns of biodiversity in terms of functional traits. Functional traits capture fundamental tradeoffs in life history strategies that can be used to determine species ecological roles and can be used to scale from organisms to ecosystems to ask broad ecological questions. The overarching goal of my dissertation is to add additional links to trait-based ecology by identifying potential sources of trait variation across different spatial and temporal gradients between varying levels of biological organization. By assessing variation across spatial-temporal scales, I tested two prominent assumptions of trait-based ecology. First, I tested the trait-environment assumption wherein traits affect ecosystem processes. Therefore, there should be a predictable relationship between traits, their environment, and ecosystem function across large ecological gradients and between broad taxonomic groups. Second, I tested the assumption that interspecific trait variation exceeds intraspecific trait variation; thus, the species mean trait value captures much of the variation for a given trait. My study systems include the latitudinal diversity gradient of North America, forests of varying successional age in the tropical dry forests of Costa Rica, and a subalpine meadow of Colorado. First, we collected leaf trait data and soil microbial data at six sites across the latitudinal diversity gradient to test a central hypothesis of trait-based ecology, primarily that shifts in plant traits associated with decomposition and nutrient availability ramify to influence microbial functioning. We found that changes in plant traits not only reflect nutrient limitation across broad ecological gradients, but also have important regional effects on biogeochemical processes, microclimates, and energy fluxes that influence microbial diversity. Furthermore, changes in plant function correspond to changes in bacterial functional traits related to carbon, nitrogen, and phosphorus cycling, although only fungal functional traits related to nitrogen cycling change across the gradient. Our results represent one of the first comparisons of functional diversity within and across bacterial, fungal, and plant communities across a latitudinal gradient. Next, we collected leaf functional trait and abiotic data across a 110-year chronosequence within a tropical dry forest in Costa Rica. We focused on six leaf functional traits for woody plants within 14 plots that have varying times since disturbance in the tropical dry forests of Guanacaste, Costa Rica. When we compare species composition and community function, we find that older tropical dry forest communities differ significantly from younger forests in species composition, above ground biomass, and functional traits. Species in younger forests have traits better adapted to hotter temperatures and increased drought. For example, young forests are characterized by thicker leaves with higher water use efficiency. In contrast, older forests have thinner broader leaves more susceptible to desiccation. Interestingly, in contrast to expectations, variation in these functional traits does not generally change through succession. This means that the different species within each community are converging on similar functional strategies. Our results also suggest that regenerating tropical dry forests are resilient and can be restored within a human lifetime. Finally, we evaluated patterns of trait variation within and between three years to understand the widely-ignored source of temporal variation associated with seasonality and test the assumption that interspecific trait variation exceeds intraspecific variation and the species means account for the overall variation of a trait. To do this, we collected leaf data from eight species at a local site in Colorado throughout the growing season, over three years, as well as extracted data from a global database and made comparisons to assess sources trait variation. We found that, although the timing of collection influences one’s ability to capture fine-scale processes occurring on short time scales, collecting data locally throughout the growing season and across multiple years does not significantly influence species ranking. However, species ranking is not conserved for comparisons between local and global databases. This suggests that extra care should be taken when applying global data for species-specific studies and that ‘snap-shot’ sampling designs may over- or underestimate community trait distributions, reducing predictability. Overall, this body of work extends beyond understanding patterns of species diversity through the inclusion of species function. It contributes to our understanding of variation in biodiversity across broad ecological gradients and between diverse taxonomic groups, how communities assemble via functional traits, and the importance of temporal variation on functional traits for detecting fine-scale patterns.

La problématique de ce travail concerne des cas concrets d'application de l'approche paléobiologique chez les ammonites. Une attention spéciale a été donnée à la variabilité intraspécifique et ses conséquences taxinomiques, ainsi qu'à l'étude des patterns et processus évolutifs des groupes étudiés, afin d'en tirer des généralités. Les études paléobiologiques sur des exemples très variés montrent que la variabilité et le degré de sa prise en compte par les auteurs a une influence particulière sur les conceptions taxinomiques, et par conséquent sur la biostratigraphie et sur la compréhension des mécanismes évolutifs. Les lois de Westermann semblent être prépondérantes pour expliquer la variabilité intraspécifique d'un grand nombre de groupes d'ammonites, mais elles ne sont pas la seule source de covariation et leur influence doit être relativisée face à d'autres facteurs habituellement peu pris en compte. Les facteurs de la variabilité intraspécifique reconnus concernent : (1) le dimorphisme, (2) une variabilité dipolaire dans l'application stricte de la première loi de Westermann, (3) des variation par rapport à l'énoncé général de cette loi, (4) une variabilité tripolaire, (5) des relations hétérochroniques morphologiques dépendantes ou indépendantes, (6) un facteur d'atténuation de l'ornementation, et (7) différents facteurs d'enroulement liés ou non à d'autres caractères morphologiques et ornementaux chez les ammonites hétéromorphes. / The issue of this work involves concrete case of application of the palaeobiological approach in ammonites. Special attention was given to intraspecific variability and its taxonomic consequences and the study of evolutionary processes and patterns of the groups studied, in order to draw generalizations. Paleobiological studies on a variety of examples show that the degree of variability and its consideration by the authors has a particular influence on taxonomic concepts, and consequently on the biostratigraphy and the understanding of evolutionary mechanisms. Westermann laws seem to be dominant to explain intraspecific variability of a large number of ammonites groups, but they are not the only source of covariation and their influence must be relativized against other factors usually not taken into account. Factors of intraspecific variability recognized are: (1) dimorphism, (2) a dipolar variability in the strict application of the first Westermann’s law, (3) changes compared to the general statement of the law, (4) tripolar variability, (5) heterochronic relationships morphological dependent or independent, (6) an attenuation factor of ornamentation, and (7) various coiling factors related or not to other morphological and ornamental characters in heteromorphic ammonites.

O presente trabalho analisou as fases gametofítica e tetrasporofítica de morfos verde, vermelho e marrons de Gracilaria domingensis em laboratório. A capacidade reprodutiva foi testada considerando-se: i, número de cistocarpos diferenciados; ii, número de carpósporos e tetrásporos liberados; iii, diâmetro desses esporos; e iv, sobrevivência de carpósporos e tetrásporos, esse último em diferentes condições de irradiância, nutrientes, e radiação ultravioleta. A capacidade somática foi testada em diferentes fases do desenvolvimento por meio da análise dos seguintes parâmetros: i, taxas de crescimento em diferentes condições nutricionais; ii, taxas de crescimento em diferentes condições de radiação ultra-violeta; iii, atividade da enzima nitrato redutase em diferentes condições nutricionais; iv, taxas de fotossíntese; e v, síntese de aminoácidos semelhantes a micosporinas (MAAs) em diferentes condições de radiação UV-B. Uma das únicas diferenças observadas na capacidade reprodutiva entre as linhagens foi o maior período de liberação de carpósporos derivados dos cruzamentos envolvendo apenas gametófitos vermelhos. Essa característica disponibilizaria uma maior quantidade de propágulos dessa linhagem, trazendo vantagens competitivas em relação às demais no ambiente natural. Plantas verdes mostraram maiores valores de fotossíntese máxima, maior síntese de MAAs quando expostas à radiação UV-B, e maior teor de proteínas solúveis totais quando comparadas às plantas vermelhas. Essas respostas sugerem adaptações a ambientes oligotróficos e com intensa luminosidade. Foi observada heterose nas linhagens de tetrasporófitos marrons (VmVd ou VdVm) com relação a pelo menos um dos seguintes parâmetros: fotossíntese máxima e eficiência fotossintetizante; taxas de crescimento; conteúdo de proteínas solúveis totais; atividade da NR; e sobrevivência de carpósporos. Esse vigor híbrido, pelo menos quanto a alguns aspectos, favoreceria a manutenção do alelo verde na natureza. As linhagens de tetrasporófitos marrons apresentaram desempenho distinto quando comparadas entre si com relação a sobrevivência de esporos e taxas de crescimento, indicando que os dois genótipos se expressam de forma diferente frente a condições abióticas distintas. O número de carpósporos liberados foi semelhante ao de tetrásporos considerando-se a massa fresca das plantas férteis, porém, a sobrevivência desses últimos foi sempre maior. O desempenho somático e reprodutivo de tetrasporófitos foi maior que o de gametófitos na maior parte das condições testadas, independentemente da linhagem. Esses resultados demonstraram que as diferentes fases do histórico de vida de G. domingensis têm características metabólicas distintas, o que confere às plantas uma maior plasticidade fenotípica. As diferenças detectadas entre as linhagens no presente trabalho foram discretas. Caso as vantagens proporcionadas pelo alelo verde fossem muito superiores às apresentadas pelo alelo vermelho, ou vice-versa, seria esperado que, ao longo do tempo, uma das duas formas excluísse a outra. A coexistência dos morfos, porém, indica que cada um deles deve ocupar um nicho ligeiramente distinto do outro, o que confere à espécie vantagens frente a ambientes heterogêneos e/ou mudanças ambientais, possibilitando uma maior capacidade de adaptação. / This work investigated gametophytic and tetrasporophytic phases of green, red and brown morphs of Gracilaria domingensis in laboratory. The reproductive performance was tested considering the following: i, number of cystocarps produced; ii, number of carpospores and tetraspores released; iii, diameter of these spores; and iv, survival of carpospores and tetraspores in different nutrient, irradiance and UV-B conditions. The somatic performance of different life phases was tested considering the following: i, growth rates on different nutritional conditions; ii, growth rates on different UV-B conditions; iii, nitrate reductase (NR) activity on different nutritional conditions; iv, photosynthetic rates; and v, mycosporine-like amino acids (MAAs) synthesis on different UV-B conditions. Carpospores originated by the cross of red males x red females were released by a longer period of time when compared to the other carpospores strains. This was almost the only difference found in the reproductive performance among different strains, and could make a greater number of the red strain propagules available for settling. This result could represent competitive advantages in the natural environment. Green plants showed greater values of maximum photosynthesis, a greater MAAs synthesis when exposed to UV-B radiation, and greater amounts of total soluble proteins when compared to the red plants. These responses suggest adaptations to high irradiances and oligotrophic environments. Heterosis was observed in one of the two brown tetrasporophyte strains considering, at least, one of the following parameters: maximum photosynthesis and photosynthetic efficiency; growth rates; total soluble proteins content; NR activity; and survival of carpospores. The heterosis related to these aspects could favor the maintenance of the green allele in nature. The two brown tetrasporophytic strains showed different performance considering the survival of spores and growth rates, which indicates that the two genotypes are expressed in different ways depending on the abiotic conditions. The number of spores released was the same for carpospores and tetraspores when expressed by the fresh biomass of the fertile plants. However, survival of tetraspores was always higher. The somatic and reproductive performance of tettrasporphytes were higher than the gametophytes ones for most of the conditions tested, regardless the strain. These results demonstrate that the different life phases of G. domingensis have specific metabolic characteristics, which contributes to a higher phenotypic plasticity of this species. The differences found among the strains were slight. If the green allele promoted very superior advantages when compared to the ones promoted by the red allele, or vice-versa, it would be expected that, within time, one of the two morphs would exclude the other. The coexistence of the morphs, however, indicates that each one of them must occupy a slightly different niche, which provides advantages to the species concerning heterogonous environments, and /or environmental changes, and enable it with a greater adaptative capacity.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第22033号 / 理博第4537号 / 新制||理||1652(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 森 哲, 教授 沼田 英治, 教授 中務 真人 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

5 Abstract Morphological traits give information about the adaptations of the studied species, the resources, which it uses, and they can generally tell us about the use of ecological space. Intraspecific morphological variability reflects various adaptations of local populations thus variability reflect different way of life. The subject of this thesis is an ecologically interesting group of specialized tropical songbirds - Sunbirds (Nectarinidae). The taxon exhibits number of adaptations to specific food collecting a combination of nectarivory and insect hunting. I focused here on the intraspecific variability of three species (Cyanomitra oritis, Cinnyris reichenowi and C. bouwieri) living in different habitats in Cameroon. The aim of this work was to find out in what morphological characters are this species sexually dimorphic. Furthermore, if their populations living in remote locations differ? Finally, if all studied species show similar trend of intraspecific variability of the traits, which could have common ecological causes. The following traits were measured: wing and tail feather length, beak dimensions, total body weight and other characters. The analysis is based on field data collected between 2003 and 2015 and museum collections. Overall, we collect information about ca. 1200 individuals for...

The family Procyonidae (raccoons, coatis, olingos, ringtails, kinkajous, and their extinct relatives) consists of six extant genera and is restricted to North and South America. Currently recognized fossil species suggest that procyonid diversity was previously much greater, including six extinct genera throughout South America. However, it is unusual that so many confamilial taxa are represented in a relatively brief span of time and restricted geographic region, and, considering that six of ten are based on badly preserved specimens, often fragments of bone with worn teeth, the validity of many of these taxa is suspect. As a step towards reevaluating past procyonid diversity in South America, we sought to identify the degree of intra- and interspecific variation in six molariform teeth of extant Procyon, particularly to identify which teeth are potentially most useful for identifying fossil procyonids. The six molariform cheek teeth analyzed consistently yielded smaller intra- than interspecific variation, permitting high accuracy of taxonomic classification. However, this accuracy varied by tooth, and the upper and lower first molars proved to be the most reliable. Thus, these particular teeth should be preferred, if available, as bases for recognizing extinct species of procyonids or reevaluating currently recognized extinct species, as a means to prevent nomina dubia.

Genetic improvement has been an integral part of loblolly pine plantation forestry in southern United States for about 60 years with focus on improving timber yield, wood quality, and pest and disease resistance. Advances in techniques of genetic selection, breeding, and propagation of planting material have made it possible to achieve genetic gains that are likely to result in significant changes in loblolly pine stand dynamics. Height-age relationships, height and diameter relationships and distributions, and intraspecific competition were investigated in second generation open-pollinated, controlled-pollinated, and clonal loblolly pine with an objective of characterizing the nature and magnitude of changes in these characteristics due to genetic improvement and clonal forestry. Genetic improvement and/or clonal forestry had no practical effect on parameters of the height-age and height-diameter relationships beyond the effect on the asymptote parameter of the Chapman-Richards and Korf equations that were used to model these relationships. Genetic improvement resulted in an increase in the mean of height distribution without a corresponding increase in the mean of the diameter distribution, but had no effect the variance and skewness of the distributions. Thus, growth and yield models in which basal area is a function of height at a specific age (site index) are likely to over predict genetic gains in basal area growth and volume yield. Increase in stand density resulted in an increase in variance of the diameter distribution of non-clonal stands but had no effect on the variance of the diameter distribution of clonal stands. Thus, diameter distribution of clonal stands may differ from that of non-clonal stands after crown closure despite the distributions not being different before and during early stages of crown closure. This study also evaluated methods that may be used to predict height growth of new genetic varieties and those that may be used to asses intraspecific competition in forest stands. Mixed-model approach of calibrating a height-age relationship to a new loblolly pine clone gave biased estimates for clones that were at the extremes of the distribution of the groups. The use of maximum likelihood with simulated annealing (MLSA) to evaluate competitive interactions among trees in loblolly pine stands gave non-unique estimates of optimum competitor selection radius. A simpler technique that uses Pearson correlations was proposed and was found to work better than MLSA. / Ph. D.

京都大学 / 0048 / 新制・課程博士 / 博士(農学) / 甲第19324号 / 農博第2145号 / 新制||農||1036(附属図書館) / 学位論文||H28||N4952(農学部図書室) / 32326 / 京都大学大学院農学研究科地域環境科学専攻 / (主査)教授 縄田 栄治, 教授 北山 兼弘, 教授 北島 宣 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DFAM

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