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京都大学 / 新制・課程博士 / 博士(情報学) / 甲第24037号 / 情博第793号 / 新制||情||134(附属図書館) / 京都大学大学院情報学研究科社会情報学専攻 / (主査)教授 大手 信人, 准教授 小山 里奈, 教授 北島 薫 / 学位規則第4条第1項該当 / Doctor of Informatics / Kyoto University / DGAM

African clawed frogs (Xenopus) have been scientific and medical model species for decades. These frogs present many curious features, and their genomic history is no exception. As such, a variety of evolutionary genomic questions can be addressed with these species in a comparative framework, owing to the great array of genetic tools available and a large number of abundant species. The sex chromosomes of this group are evolutionarily young, and this thesis establishes that there has been an additional change in what constitutes the sex chromosomes in one species of Xenopus. This allows us to compare the evolutionary trajectory of newly established sex chromosomes. By exploring the genetic content of these systems, profiling their recombinational activity, and assessing the extent of nucleotide divergence between the sex chromosomes, we find that sex chromosome evolution may be predictable in some aspects, and highly unpredictable in others. In addition, this genus is uncharacteristic for vertebrates in the frequency with which lineages undergo whole genome duplication. In this thesis, we explore the selective dynamics operating on duplicate genes over time, and the rate at which duplicate copies are purged from the genome from multiple Xenopus species. These investigations provide an animal perspective on the subject of biased subgenome evolution, characteristic of allopolyploids. The last two chapters of this thesis redefine the species boundaries for the most intensively studied Xenopus species (X. laevis), and explore the genetic extent of hybridization between the common X. laevis and the endangered X. gilli. Overall, this thesis provides a broad look at several aspects of Xenopus evolutionary genomics, providing novel contributions to the fields of sex chromosome research, whole genome duplication, and speciation and hybridization. / Thesis / Doctor of Philosophy (PhD)

The Roanoke logperch (Percina rex) is a federally endangered large darter that occurs only within the Roanoke and Chowan drainages of Virginia. This dissertation examines multi-scale habitat use patterns by logperch in three river systems in Virginia, including comparisons among rivers and life stages. The first study in this dissertation compares microhabitat use patterns of logperch among the Roanoke, Pigg, and Nottoway rivers. My objectives are to: 1) compare available microhabitat and microhabitat use by logperch among these rivers; and 2) examine the transfer of habitat models among rivers. Habitat availability in the three rivers indicates that the Nottoway River is least impacted by human activity, while the Pigg River is most impacted. The Roanoke and Pigg rivers are found within the same region of Virginia and share many habitat characteristics. Logperch consistently use silt free, loosely embedded gravel in all rivers and can occupy a variety of depths and velocities to accommodate substrate requirements. Microhabitat models transfer better between the similar Pigg and Roanoke rivers. The second study in this dissertation compares micro- and meso-habitat use patterns by Roanoke logperch in the Roanoke and Nottoway rivers. My objectives are to: 1) compare micro- and meso-habitat use patterns of logperch in the Roanoke and Nottoway rivers; and 2) examine transfer of habitat models at both scales. An increase in scale from micro- to meso- habitat did not improve model transfer. Habitat selectivity and transfer was strongest at the microhabitat scale. Logperch appear to be microhabitat substrate specialists and mesohabitat generalists. The final study in this dissertation examines ontogenetic patterns of habitat use by Roanoke logperch in the Roanoke and Nottoway rivers. My goals are to: 1) examine habitat use by three age classes of logperch and 2) compare ontogenetic patterns of habitat use between the Roanoke and Nottoway rivers. In the Roanoke River, adult and subadult logperch primarily used run and riffle habitat, often over gravel substrate. Subadults were found in lower water velocities and more embedded microhabitats than adults. Young-of-year logperch were found in shallow, stagnant backwaters and secondary channels. In the Nottoway River, both adult and subadult logperch were found over sand and gravel in deep, low velocity pools and runs. Subadults were observed in slightly more silted, lower velocity habitat. Younger age classes of logperch appear to be more vulnerable to sedimentation caused by human activity. Evidence in this dissertation strongly indicates that logperch have strict substrate requirements and the distribution of habitat types and pathways of dispersal will be critical for completion of the logperch life cycle. A watershed-level conservation approach that addresses sediment loading and preserves ecological processes that provide ephemeral, seasonal, and persistent types of habitat required over logperch ontogeny will be most effective for management geared towards the recovery of this endangered species. / Ph. D.

The southeastern United States is a global biodiversity hotspot but has experienced severe declines of natural ecosystems. The southeast is currently facing widespread change, particularly from an increasing human population and climate change, that is likely to impact all remaining natural areas to some degree. In this work, I examine some of the challenges currently facing imperiled species of reptiles and amphibians in this region. The work is focused on two species, the Reticulated Flatwoods Salamander (Ambystoma bishopi) and the Eastern Indigo Snake (Drymarchon couperi) both of which are listed on the U.S. Endangered Species List. Chapter 1 used Light Detection and Ranging (LiDAR) data to measure wetland bathymetry (basin shape) in flatwoods salamander breeding wetlands. Bathymetry data were used to construct stage–area relationships for each wetland, and a history of water level monitoring data were applied to these relationships to build multi-year time series of flooded area metrics. These metrics were then combined with an assessment of vegetation characteristics to map potentially suitable habitat for flatwoods salamander breeding within each wetland. Chapter 2 focused on flatwoods salamander phenology (the timing of life history events) in response to climate change. I quantified flatwoods salamander movements into and out of breeding wetlands over a 10-year period (2010–2020), identifying temperature and precipitation patterns that were conducive to salamander movements. I then used future climate projections to forecast movement opportunities for flatwoods salamander from 2030–2099 and used an existing hydrologic model built on the same climate data to understand how phenology may interact with hydrology. Overall, only a small number of years are likely to have an ideal intersection of phenology and hydrology as has been observed during recent breeding seasons. Chapter 3 described the construction of a stochastic Integral Projection Model for flatwoods salamanders. I integrated the projections from Chapter 2 with the population model to estimate the viability of two flatwoods salamander populations from 2030–2099 under multiple climate change scenarios. The results indicated that approximately half of the examined scenarios resulted in a high probability (>0.5) of extinction when considering both wetland hydrology and salamander phenology. In Chapter 4, I described the creation of a stochastic Integral Projection Model for indigo snakes. I then demonstrated the utility of this model by examining the effects of initial population size, road density, and removal of individuals to support a captive colony on indigo snake populations. I found that high road densities and high collection rates would likely lead to population declines, although the rate of declines and extinction risk varied across scenarios. Taken together, these projects highlight some of the challenges currently facing herpetofauna in the southeastern United States, demonstrate the difficulty in conserving these often-overlooked species, and provide useful tools for ongoing conservation efforts focusing on these two imperiled species. / Doctor of Philosophy / We are in the midst of a global biodiversity crisis, with rates of species extinction far exceeding normal levels. Species loss is largely driven by global change attributable to human activities. A rapidly changing world can make it challenging to effectively conserve and manage imperiled species. In this work, I studied two species found only in the southeastern United States that are listed on the U.S. Endangered Species List. Chapters 1–3 focused on the Reticulated Flatwoods Salamander (Ambystoma bishopi), while Chapter 4 focused on the Eastern Indigo Snake (Drymarchon couperi). In Chapter 1, I used high resolution elevation data to map the shape of flatwoods salamander breeding wetlands. These data were then used to estimate flooded areas across multiple years. Flooded area metrics were combined with vegetation measurements to map potential flatwoods salamander breeding habitat. In Chapter 2, I examined how flatwoods salamander movements may respond to climate change. I identified time periods and weather conditions that coincided with flatwoods salamander movements into and out of breeding wetlands. I then projected potential movement opportunities based on multiple future climate scenarios for each breeding season from 2030–2099. My results showed that few years are likely to be ideal for flatwoods salamander reproduction, which is similar to trends observed in recent years. In Chapter 3, I built a population model for flatwoods salamanders. I then combined the model with predictions made in Chapter 2 to estimate the probability that populations would go extinct by the end of the century. The results indicated that the two flatwoods salamander populations examined had a high probability (>0.5) of extinction in about 50% of the climate scenarios. In Chapter 4, I constructed a population model for indigo snakes using a variety of available data. I used this model to examine the effects of road density, initial population size, and removal of individuals to support a captive colony on indigo snake populations. The results suggested that populations experiencing high road densities or high collection rates were likely to decline over time. These projects highlight some of the difficulties in conserving often-overlooked reptiles and amphibians in the southeastern United States and provide important tools for ongoing conservation projects working with these two imperiled species.

In order to prevent species extinctions, conservation strategies need to incorporate the identification and mitigation of the root causes of population decline with an assessment of vulnerability to genetic and stochastic factors affecting small populations. Species or populations with small ranges, such as those on islands, are particularly vulnerable to extinction, and deficient knowledge of these species often impedes conservation efforts. The Bahama Swallow (Tachycineta cyaneoviridis) is an endangered secondary cavity-nester that only breeds on three islands in the northern Bahamas: Abaco, Grand Bahama, and Andros. I investigated questions related to population size and distribution, genetic diversity and population structure, breeding biology, and ecological interactions of the swallow, with the goal of informing the conservation and management of the species. Using several population survey methods on Abaco, I found that swallow site occupancy and density is higher in southern Abaco, especially near roads and pine snags. Future research should prioritize identifying the causes of variable and low population densities in parts of the swallow's range. I used microsatellite markers and morphometrics to assess differences between populations on Abaco and Andros. We found a lack of genetic differentiation (G'ST = 0.03) between populations, but differences in morphology suggest that gene flow might be low enough to enable traits under selection to diverge. By locating and monitoring nests, I found that swallows rely on woodpecker-excavated cavities in pine snags and utility poles, and that swallows nesting in pine snags had higher fledging success (92%) than those nesting in utility poles (50-62%). Using a cavity nest-web approach, I assessed how swallows interact with cavity-nesting birds and resources on Abaco. Hairy Woodpeckers (Dryobates villosus) primarily excavated pine snags, while West Indian Woodpeckers (Melanerpes superciliaris) excavated utility poles in non-pine habitat. Only swallows and La Sagra's Flycatchers (Myiarchus sagrae) used nest sites in the pine forest. Swallows in non-pine habitat face competition for cavities with American Kestrels (Falco sparverius), and non-native House Sparrows (Passer domesticus) and European Starlings (Sturnus vulgaris). These results highlight the importance of pine forest and the Hairy Woodpecker for the persistence of the swallow. / Doctor of Philosophy / In order to prevent species extinctions, conservation strategies need to identify and resolve the problems that cause species to decline, as well as address issues characteristic of small populations. Species or populations with small ranges, such as those on islands, are particularly vulnerable to extinction, and lack of knowledge of these species often impedes conservation efforts. The Bahama Swallow is an endangered species that only breeds on three "pine islands" in the northern Bahamas. The swallow is a secondary cavity-nester, which means that it nests in a cavity, usually either a natural tree hole or a hole created by another bird species. In this study, I investigated where swallows are found on the islands, the genetics and body sizes of populations, nesting biology, and connections with other species, with the goal of providing information for the conservation and management of the species. On Abaco, I found that there are more swallows in the southern part of the island, especially near roads and the dead standing pine trees (pine snags) used for nesting. Future research should assess why there are fewer birds in other parts of the swallow's range. I tested whether the genetics and body sizes of populations on Abaco and Andros are different from each other. Populations were genetically similar, but may be separated enough to result in body size differences. I located swallow nests on Abaco, and found that swallows rely on cavities made by woodpeckers in pine snags and utility poles. By visiting some nests repeatedly, I found that swallows nesting in pine snags were more successful than those nesting in utility poles. I also assessed how swallows interact with the other bird species that create and use nesting cavities. Hairy Woodpeckers usually create cavities in pine snags, while West Indian Woodpeckers use utility poles outside of the pine forest. Swallows nesting in the pine forest compete with fewer bird species for cavities than swallows nesting in other habitats. These results show that managing the forest to retain pine snags and Hairy Woodpeckers is important for the conservation of the swallow.

Geographic Information Systems (GIS) and remote sensing technologies were used to identify and describe potential habitat for three species endemic to the Southwestern United States; the Golden-cheeked Warbler (Dendroica chrysoparia), the Black-capped Vireo (Vireo atricapillus), and the Texas kangaroo rat (Dipodomys elator). For each species, the computerized classification of digital satellite imagery was integrated with ancillary spatial information (e.g. soils, geology, and land use) to construct a data base to be used for ecological evaluation as well as habitat protection and management measures.

Thesis (MScConsEcol (Conservation Ecology and Entomology))--University of Stellenbosch, 2006. / Using the World Conservation Union’s (formerly the International Union for the Conservation of Nature) IUCN 2001 categories and criteria, the conservation status of the South African butterfly fauna has been reassessed. This study includes an assessment of the 62 globally threatened South African taxa and 1 that has a marginal distribution in this region.

The capercaillie is a rare species of forest bird which is once again facing extinction in Britain. Working pine forests are thought to represent the future for capercaillie in Scotland. However, few studies have focused on capercaillie within working forests and as a result current management recommendations are based on studies conducted within semi-natural pinewoods. Therefore study of the ecology of capercaillie within working forests is vital for the survival of the species in Scotland. The present study investigated the ecology of capercaillie, based on indirect measures of capercaillie use, within Morangie Forest, a pine plantation managed by the Forestry Commission. Monthly dropping counts were collected over a 30 month period to gain insight into the capercaillie population at Morangie Forest, and to explore the use of faecal counts as a non-invasive monitoring tool. Habitat analysis, dietary analysis of droppings and genetic analysis of feathers were further conducted to fully explore ecology of capercaillie within Morangie Forest. Results showed that the capercaillie population in Morangie Forest was subject to seasonal movements and that movements were not confined to the bounds of the forest. Substantial mixing occurred with capercaillie from neighbouring Novar Forest. In addition, capercaillie within Morangie Forest selected habitat at a radius of 50m and beyond. The population of capercaillie at Morangie Forest was estimated at around 65 individuals based on measures from dropping counts and genetic determination of individuals from feathers. Results also suggested that winter dropping counts could represent a useful tool for forest managers to monitor the size and movement of capercaillie populations frequenting their forests. In addition winter dropping counts could be used in conjunction with habitat data to plan management of habitat for capercaillie. The present study contributes towards future management strategies for the conservation of capercaillie within working forests and suggests further research priorities for capercaillie in Scotland, particularly those focused at a landscape scale.

Biodiversity conservation requires a framework for prioritising limited resources to the many endangered species. One such framework that has seen much attention and is considered extensively in this thesis, is the Noah's Ark Problem (NAP). The NAP combines a biodiversity measure (Phylogenetic Diversity; PD) with species survival probabilities and conservation costs. The aim of the NAP is to allocate the limited conservation resources such that the future expected PD is maximised. Obtaining optimal solutions to the NAP is a computationally complex problem to which several efficient algorithms are provided here. An extension to the NAP is also developed which allows uncertainty about the survival probability estimates to be included. Using this extension we show that the NAP is robust to uncertainty in these parameters and that even very poor estimates are beneficial. To justify using or promoting PD, it must produce a significant increase in the amount of biodiversity that is preserved. We show that the increase attainable from the NAP is typically around 20% but may be as high as 150%. An alternative approach to PD and the NAP is to prioritise species using simple species specific indices. The benefit of these indices is that they are easy to calculate, explain and integrate into existing management frameworks. Here we investigate the use of such indices and show that they provide between 60% and 80% of the gains obtainable using PD. To explore the expected behaviours of conservation methods (such as the NAP) a distribution of phylogenetics trees is required. Evolutionary models describe the diversification process by which a single species gives rise to multiple species. Such models induce a probability distribution on trees and can therefore be used to investigate the expected behaviour of conservation methods. Even simple and widely used models, such as the Yule model, remain poorly understood. In this thesis we present some new analytic results and methods for sampling trees from a broad range of evolutionary models. Lastly we introduce a new model that provides a simple biological explanation for a long standing discrepancy between models and trees derived from real data -- the tree balance distribution.