Populations in fragmented habitat patches may show ecological differentiation, which has implications for metapopulation viability. In this study we used a common garden with two watering treatments to contrast mean differences in ecophysiological traits and the relationships between traits and performance among seven populations of three alvar species. These species differ in their alvar specialization in Ontario, from almost endemic (Solidago ptarmicoides) to highly confined (Dasiphora fruticosa) to a widespread generalist (Hypericum perforatum). Populations of all species exhibited mean differences in at least one trait: photosynthesis (Amax), growth rate and specific leaf area. More surprisingly, the relationship between functional traits and performance was significantly different among populations in all species, suggesting different strategies for maximizing growth in different environments. The ecological differentiation observed affected all species regardless of distribution and is likely genetically based. This differentiation may destabilize metapopulation dynamics and reduce rates of spread if colonization is negatively impacted.

Populations in fragmented habitat patches may show ecological differentiation, which has implications for metapopulation viability. In this study we used a common garden with two watering treatments to contrast mean differences in ecophysiological traits and the relationships between traits and performance among seven populations of three alvar species. These species differ in their alvar specialization in Ontario, from almost endemic (Solidago ptarmicoides) to highly confined (Dasiphora fruticosa) to a widespread generalist (Hypericum perforatum). Populations of all species exhibited mean differences in at least one trait: photosynthesis (Amax), growth rate and specific leaf area. More surprisingly, the relationship between functional traits and performance was significantly different among populations in all species, suggesting different strategies for maximizing growth in different environments. The ecological differentiation observed affected all species regardless of distribution and is likely genetically based. This differentiation may destabilize metapopulation dynamics and reduce rates of spread if colonization is negatively impacted.

No description available.

Doctor of Philosophy / Department of Division of Biology / John Blair / The feasibility of restoration, which traditionally targets historical conditions, is questionable in the context of global change. To address this, my dissertation investigated (Chapter 2) the patterns of restoration establishment along a chronosequence of restored prairies with respect to nearby remnant prairies, (Chapters 3-4) responses of plant communities in restorations initiated using different methods (levels of species richness and sowing density) to drought, which is projected to increase in frequency, and (Chapters 5-6) the effects of propagule source and variation (mixing among sources) on restoration establishment and the generality of restoration outcomes across variable environments using reciprocal common gardens of multi-species restoration seedings. Chapter 2, published in Restoration Ecology, showed that restoration led to the recovery of desirable characteristics within several years, but restorations utilizing primarily fall-collected seeds likely diminished the representation of early phenology species, so biodiversity may be further enhanced by including early phenology species in seeding mixes. Chapters 3 and 4, published respectively in Ecological Applications and Applied Vegetation Science, examined the establishment of native plant communities after seeding and their responses to experimentally imposed drought. Both high seed mixture richness and high density seeding resulted in greater establishment of native, seeded species compared to low richness and low density treatments, and exotic species were less prevalent in high richness and high density treatments. However, we found little evidence of differential drought resistance, recovery, and resilience among treatments. This result coupled with increases in exotic species following drought suggest that other forms of active management may be needed to produce restored plant communities that are robust to climate change. Chapter 5 (published in Ecosphere) iii and Chapter 6 found that seed source affects individual species establishment, community structure, and productivity. However, there was no consistent advantage for any source, including local sources, across sites or species. This suggests that source effects on single species or effects observed at single locations should not be broadly generalized. Together, this dissertation shows that restoration can recover many characteristics of native prairies and that manipulation of seeding practices (seed mixture richness, seeding density, seed source) influence grassland establishment in terms of productivity, community structure, invasion, and the abundance and survival of individual species.

At broad scales, community ecologists study how biogeographic factors like environmental dissimilarity and geographic distance influence community assembly and composition. At small scales, community ecologists study how one or several species interact to determine habitat partitioning and coexistence. In this dissertation, I present studies at both scales. Chapter One investigates community assembly across the Caribbean, Central, and South American radiations of Anolis lizards and Eleutherodactylid frogs to test whether oceanic islands are unique in their assembly processes. Such uniqueness is suggested by high levels of endemism on islands; however, comparable levels of endemism can be found in mainland communities. I modeled the rate of species turnover between mainland communities, with respect to geographic distance and environmental dissimilarity, and then used the mainland model to predict turnover among islands. Turnover among island communities was significantly higher than predicted from the mainland model, confirming the long-held but untested assumption that island assemblages accumulate biodiversity differently than their mainland counterparts. Chapter Two reviews the evidence for ecological character displacement (ECD), an evolutionary process whereby two resource competitors diverge from one another in phenotype and resource use, facilitating coexistence in a community. I find that, despite current scientific opinion, the evidence for ECD is equivocal; most cases of ECD pattern fail to rule out processes alternative to resource competition that could create the same pattern. I conclude that better evidence may come from real time tests of ECD. Chapters Three and Four describe just such a test in small island populations of Anolis carolinensis. In Chapter Three, I find that small island populations of A. carolinensis that have come into sympatry with a novel competitor, the invasive A. sagrei, shift their habitat use to become more arboreal, compared to allopatric populations. Consistent with prediction, individuals from sympatric populations have larger toepads with additional adhesive scales - a common adaptation to arboreality in Anolis. In Chapter Four, I describe a common garden experiment that finds that the observed toepad divergence is an evolved response, suggesting rates of divergence for toepad area and scale number on par with well known examples of contemporary evolution.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第22286号 / 理博第4600号 / 新制||理||1660(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)教授 工藤 洋, 教授 田村 実, 准教授 高山 浩司 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Penstemon is one of North America's largest endemic genera with over 280 described species. These species are distributed throughout most of North America from the Arctic northern latitudes to tropics of Central America. The genus has historically been divided into six subgenera, but has recently been reorganized into two subgenera following some recent phylogenetic studies. I made a comprehensive assessment of the Penstemon genus' geographic distribution utilizing herbaria databases by ecoregion to discuss the general ecologic adaptations of each historic subgenera. I also assessed the Penstemon genus' bee pollinator diversity utilizing online databases of bee specimen collections associated with Penstemon flowers. I investigated the efficacy of utilizing the plastid genomes (plastomes) of 29 species in the Lamiales order, including five newly sequenced Penstemon plastomes, for analyzing phylogenetic relationships and resolving problematic clades. I compared whole-plastome based phylogenies to phylogenies based on individual gene sequences (matK, ndhF, psaA, psbA, rbcL, rpoC2, and rps2) and concatenated sequences. I found that my whole-plastome based phylogeny had higher nodal support than all other phylogenies, which suggests that it provides greater accuracy in describing the hierarchal relationships among taxa as compared to other methods. I found that the genus Penstemon forms a monophyletic clade sister to, but separate from, the Old World taxa of the Plantaginaceae family included in our study. My whole-plastome based phylogeny also supports the rearrangement of the Scrophulariaceae family and improves resolution of major clades and genera of the Lamiales. I evaluated 16 accessions of P. cyaneus with 14 accessions of closely related Penstemon species in common garden in two distinct environments in Aberdeen, ID and Provo, UT during 2018 and 2019. I evaluated the accessions for key commercial seed production traits including survival, plant height, number of stems, and seed production. Both common gardens received supplemental irrigation during 2018, but I withheld irrigation during 2019. Plant survival in our Aberdeen, ID site was not significantly different between years, but survival was a significantly lower in 2019 than in 2018 at our Provo, UT site. The mean survival for P. cyaneus accessions ranged from 56% to 94%, and the mean seed production ranged from 91.2 kg/ha to 397.6 kg/ha. I recommend developing a commercial seed source derived from pooling germplasm of six accessions (PECY3-367, PECY3-371, PECY3-376, PECY3-443, PECY3-457, and PECY3-458). These accessions had mean survival rates of 82-94%, and seed production of 196.2-397.6 kg/ha.

Western prairie clover [Dalea ornata (Douglas ex Hook.) Eaton & J. Wright] is a perennial legume that occurs in the northern Great Basin, Snake River Basin, and southern Columbia Plateau, whereas Searls prairie clover [Dalea searlsiae (A. Gray) Barneby], also a perennial legume, occurs in the southern Great Basin and surrounding areas. Understanding the genetic and ecotypic variation of these prairie clovers is a prerequisite for developing populations suitable for rangeland revegetation in the western USA. DNA sequences of internal transcribed spacer (ITS/5.8S) and trnK/matK were used to study the phylogeny of these species. The species were distinguished by DNA sequences from both regions and conserved haplotypes were observed between and within species. Common-garden plots of 22 collections of western prairie clover from Idaho, Oregon, and Washington and 20 collections of Searls prairie clover from Utah and Nevada were established in northern Utah for phenotypic evaluation. Significant variation was detected among the collections for all traits measured in the common gardens for both species. Flowering date was correlated with collection-site temperature and elevation in western prairie clover collections, whereas biomass-related traits were closely related with collection-site precipitation in Searls prairie clover. Population structure from amplified fragment length polymorphism (AFLP) markers resulted in two distinct, genetically differentiated groups and a third admixed group in western prairie clover, and flowering date played a significant role in discriminating those genetic-based groupings of collections. For western prairie clover, two populations are recommended for development, one from the Deschutes River watershed and another from the remaining collections. For Searls prairie clover, two genetically different groups of collections were identified from southern Utah and eastern Nevada and from northwestern Utah. Three western Nevada collections exhibited close association with eastern Nevada and southern Utah groups for AFLP-markers but with collections from northwestern Utah for phenotypic traits. Strong isolation by distance was observed for Searls prairie clover collections suggesting that genetic drift and gene flow are major factors for determining population structure in this species. As a result, two regional seed sources should be developed for Searls prairie clover, one from northwestern Utah and the other from eastern Nevada and southern Utah.

Two separate experiments were designed to assess the value of Lacy Buckwheat (Eriogonum corymbosum ) as a low water landscape plant. Low water use landscapes can contribute to water conservation in arid climates. Developing a palette of plants that are both attractive and drought tolerant can promote the acceptance of low water use landscapes as an alternative to the traditional bluegrass landscapes of the Intermountain West. Eriogonum corymbosum is an attractive subshrub species native to low rainfall areas of the Colorado Plateau. A strip plot design containing four repetitions with four randomly assigned plants each of Eriogonum corymbosum , Eriogonum thompsoniaeand the control species Cornus sericea `Kelseyi' was established to determine E.corymbosum tolerance to frequent irrigation. Two water treatments were assigned to the repetitions for each species. One treatment was watered by a drip irrigation system with sixteen liters of water every three days; the other treatment was not watered. Stomatal conductance (Gs) and plant water potential were assessed weekly for each species from June through August for the years 2009 and 2010. In 2009 and 2010 bothEriogonum accessions showed no significant difference with the water treatment/accession interaction. NeitherE. corymbosum accession exhibited differences in stomatal conductance or water potential between the wet and dry treatments for the length of study season over both years.Cornus sericea `Kelseyi' showed less ability to withstand the prolonged dry frequencies. Eriogonum corymbosum has many aesthetic qualities, in addition to being drought tolerant, such as long duration late season blooming of yellow and white flowers, and an appealing hemispherical crown shape. A second study was designed to investigate the morphological diversity of thirteen Eriogonum accessions collected in the state of Utah and established in a common garden. Nineteen different variables made up of both quantitative and qualitative morphological characteristics comprised of leaf, canopy and floral characteristics were selected to compare between and within accessions. These characteristics were observed or measured, then used in a Multidimensional Preference analysis (MDPREF) to facilitate the selection of potential cultivars. The MDPREF is useful in selecting accessions with unique combinations of ornamental characteristics that could have a marketable advantage.

Plants adjust to challenging environments by genetically fixed changes and phenotypically plastic response. Alpine environments pose multiple challenges to plant life including cold, high irradiance and short vegetative period. To survive such specific conditions, plants often significantly alter their morphology. In my thesis I studied to which extent specific traits of alpine ecotypes repeatedly appear among independently formed alpine populations and to which extent these changes represent fixed genotypic differentiation vs phenotypic plasticity. To address these questions I performed an experiment in which Arabidopsis arenosa plants from sixteen populations belonging to two ecotypes (alpine and foothill) were grown in conditions resembling alpine vs foothill conditions. Specifically, I modified levels of irradiance and temperature and complemented alpine-like and foothill-like treatment by additional two extreme treatments to reach full-factorial design. I used discriminant and classificatory analysis to examine the overall morphological differentiation characterised by set of twenty measured traits. Then I examined variation in each trait by statistical Bayesian model that I designed for this purpose. I found out that although ecotypes are predominantly differentiated by fixed morphological...