京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第20962号 / 理博第4414号 / 新制||理||1634(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 湯本 貴和, 教授 古市 剛史, 教授 友永 雅己 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

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Grasslands cover 40% of the earth’s land area and provide numerous valuable ecosystem services. However, climate change, global land use change and increasing intensive anthropogenic interventions make grasslands to one of the most endangered ecosystem types in the world. Effective protection in the future requires a fundamental understanding of the dynamics of grasslands and their major drivers. Field experiments have been conducted for impact analyses, for example, with different management intensities, plant community composition and altered climatic conditions. Complementary, ecological models allow to extend the analysis to long-term effects of changes as well as to a deeper understanding of the underlying ecological processes. In this thesis, an individual-based grassland model and network science were applied to understand the community structure and dynamics emerging from individual plant interactions – in relation to plant traits, ecological processes, environmental and anthropogenic impacts, and the small-scale spatial distribution of plants. In the first study, an individual-based process-oriented grassland model was parameterized to simulate field data of a local biodiversity experiment using the concept of plant functional types. The influence of various functional plant traits and ecological processes on grassland productivity and functional composition were analyzed. Different functional plant traits showed partly contrasting effects on plant growth. With regard to the modeled ecological processes, competition for space between plants affected grassland productivity more than shading of plants. In the second study, the parameterized grassland model was used to analyze the impact of functional diversity, mowing frequency and air temperature on ecological processes that lead to changes in grassland productivity. The model reproduced the increase of biomass yields with functional diversity as observed in the field experiment. Modeled plant competition for space showed to be the dominant process and was responsible for an increase in biomass yields in more frequently mown grasslands. In the third study, an approach to generate a regionally transferable parameterization of the grassland model is presented. The impact of management, environment and climate change on productivity and functional composition of grasslands was analyzed within a German-wide scenario analysis. Management intensity had more influence on grassland productivity than environmental factors and correlations of productivity with environmental factors become stronger in less managed grasslands. Climate change showed to have only a minor influence on simulated vegetation attributes. In the fourth study, network science was applied to forest megaplots to quantify the spatial neighborhood structure of species-rich ecosystems. Networks at the individual-tree and tree-species levels revealed similar structures at three investigated forest sites. Tropical tree species coexisted in small-scale networks and only up to 51% of all possible connections between species pairs were realized. A null community analysis showed that details on the tree position and tree size have no major influence on the network structures identified. In summary, this thesis presents the development of advanced methods and analysis tools as well as their application to vegetation ecosystems with high diversity. Thereby, complex structures and dynamics of ecological systems could be systematically explored by combining ecological models with extensive field measurements.

Human activities are a major driver of biodiversity degradation and loss, especially in tropical forest areas, where forest-fringe towns and villages depend on the forests for their livelihoods. In order to reduce threats that human activities pose to biodiversity, livelihoods support programs are employed as economic incentives for biodiversity conservation. These programs support the livelihoods activities of local communities, with the aim of triggering favorable attitudes and behaviors towards conservation, and ultimately reduce biodiversity degradation. Their effectiveness as conservation tools has not been evaluated. I investigated the effects of livelihoods programs on conservation attitudes and the consequent effects on biodiversity in the Afadjato-Agumatsa and Atewa forest areas in southeastern Ghana. The study areas are coupled human and natural systems, which are excellent for research in the theoretical framework of biocomplexity in the environment. Using literature reviews and field visits, I documented the specific livelihoods support activities (LSAs) used for biodiversity conservation, their historical trend and geographical distribution in Ghana. I used ex-post costbenefit analysis to determine socio-economic estimates of the LSAs in the two forest areas. Since communities were not randomly assigned to the interventions, I employed quasi-experimental design to evaluate the effects of LSAs on environmental attitudes. I evaluated the effect of conservation attitudes on biodiversity at two levels. These levels included 1) functional biodiversity at the landscape level represented by mean Normalized Difference Vegetation Index (NDVI) of forest; and 2) compositional biodiversity at the species level represented by species diversity of fruit bats. iv The earliest record of LSAs used for biodiversity conservation in Ghana was in 1993. I identified 71 different activities belonging to eight categories. Some of these activities are beekeeping, animal husbandry, crop farming, and snail rearing. Most LSA programs have been in northern Ghana. There was an increasing tendency to make LSAs part of every conservation program in Ghana and this satisfies the current policy of collaborative conservation. The socio-economic estimates of LSAs included: 1) capital investment; 2) net socio-economic benefits; and 3) the benefit-cost ratio. The per-community values of the three estimates were not different between the two study areas. The per capita values of capital investment and net economic benefit were not significantly different between the two study areas. However, benefitcost ratio per capita was higher in Afadjato-Agumatsa than in Atewa. Estimates of economic returns from LSAs were marginal but the perceptions of success were relatively high. Environmental attitudes in LSA communities and non-LSA communities were not significantly different, and this was confirmed by an estimate of infinitesimal effects of LSAs on forest conservation attitudes. Among LSA communities, benefit-cost ratio of LSAs predicted favorable forest conservation attitudes; and change in pro-conservation attitudes were significantly higher in communities that had active LSAs than in communities which had no active LSA. Mean NDVI of the forests decreased from 1991 to 2000 and decreased further but at a slower rate to 2010. Higher forest conservation attitudes predicted higher mean NDVI in 2010. Higher change in mean NDVI from 1991 to 2000 predicted higher change in mean NDVI from 2000 to 2010. Eleven of the 13 fruit bat species in Ghana were recorded in the study areas. Longer v distances between a local community and its forest predicted higher species diversity of forestspecialist fruit bats. The results indicate that LSAs have become a major contribution to Ghana’s current collaborative forest policy. The fact that perceptions of LSA success were moderate even though the economic returns from them were marginal suggest that other factors such as provision of employment, training in new skills and community cohesion played a part in how communities viewed the success as LSAs. Evaluations of conservation attitudes suggest that just participating in LSAs did not improve attitudes; but higher benefit-cost ratio predicted favorable conservation attitudes, and conservation attitudes were higher in communities that sustained their LSAs. Therefore, it may serve biodiversity conservation to invest in LSAs that can be sustained and involve the least costs to local communities. Primary production of the forests, a proxy for a functional habitat, continued to decrease. Preventing communities from locating closer to forests could improve fruit bat diversity, which contributes to natural forest regeneration. Improving conservation attitudes should be an objective of conservation at the landscape scale. On the basis of the results, I developed a conceptual model for forest biodiversity conservation in a biocomplexity framework. This model could be useful for evaluating conservation in tropical forest areas. Lessons from this study can be applied in other incentive-based conservation programs such as payments for ecosystem services systems and carbon market schemes. I suggest that this study be repeated after a decade and that other socio-political and biogeochemical variables be integrated into future studies.

Global warming and climate change are closely related to the amount of CO2 in the air. Forest ecosystem plays very important role in the global carbon cycle; CO2 from the atmosphere is taken up by vegetation and stored as plant biomass. Therefore, quantifying biomass and carbon sequestration in tropical forests has a significant concern within the United Nations Framework Convention on Climate Change (UNFCC), Kyoto Protocol and Reducing Emission from Deforestation and Forest Degradation (REDD) program for the purpose of the improvement of national carbon accounting as well as for addressing the potential areas for carbon credits, basis for payment for environmental services. The aim of research is to estimate biomass and carbon stocks in tropical forests using remote sensing data for dry forest of central highlands of Vietnam. This result showed that from satellite images of the SPOT, satellites could build the land cover map, carbon map and biomass map of Yok Don National Park, Central Highlands of Vietnam. Through which also the biomass (above ground biomass and below ground biomass) of each type of forest can be calculated. For instance the biomass of the dry forest (Dry Dipterocarp Forest) is 153.49 tones x ha-1, biomass of rich forest is 343.35 tones x ha-1, biomass of medium forest is 210.34 tones x ha-1 and biomass of poor forest & scrub are 33.56 tones x ha-1. / Sự ấm lên toàn cầu và biến đổi khí hậu có liên quan chặt chẽ với tổng lượng CO2 trong không khí. Hệ sinh thái rừng có vai trò rất quan trọng trong chu trình các bon toàn cầu; khí CO2 trong khí quyển được hấp thụ bởi thảm thực vật dưới dạng sinh khối. Vì vậy, việc xác định sinh khối và carbon tích trữ trong rừng nhiệt đới đã có được sự quan tâm đáng kể trong Công ước của Liên hiệp quốc về biến đổi khí hậu (UNFCC), Nghị định thư Kyoto và Chương trình giảm phát thải từ phá rừng và suy thoái rừng (REDD) gần đây, nhằm cho mục đích cải thiện việc tính toán lượng các bon tích trữ cũng như giải quyết các vấn đề tiềm năng cho tín dụng các bon, làm cơ sở cho việc thanh toán cho các dịch vụ môi trường. Mục đích của nghiên cứu này là ước lượng sinh khối và các bon lưu trữ trong các khu rừng nhiệt đới bằng cách sử dụng dữ liệu viễn thám, mà ở nghiên cứu này là cho rừng khộp Tây Nguyên của Việt Nam. Kết quả cho thấy rằng, từ ảnh vệ tinh SPOT có thể xây dựng bản đồ lớp phủ thực vật, bản đồ các bon và bản độ sinh khối của Vườn quốc gia Yok Đôn, Tây Nguyên Việt Nam. Qua đó đã tính toán được sinh khối (bao gồm cả trên mặt đất và dưới mặt đất) như: đối với sinh khối của rừng khô cây họ dầu (Dry Dipterocarp Forest) là 153,59 tấn/ha, sinh khối rừng giàu là 343,35 tấn/ha, sinh khối rừng trung bình là 210,34 tấn/ha và sinh khối rừng nghèo&cây bụi là 33,56 tấn/ha.

No description available.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第19360号 / 理博第4122号 / 新制||理||1593(附属図書館) / 32374 / 新制||理||1593 / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 中川 尚史, 教授 中務 真人, 教授 曽田 貞滋 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DFAM

Tropical forests are storehouses of more thanhalf of the world‘s biodiversity and play a key role in global carbon, water and energy cycles. However, as a consequence of rapid anthropogenic climate change, biodiversity and climate functions of these forests are under a threat. Climate is changing not only in mean state but its variability is increasing, with extreme events such as droughts, heat waves and storms also rising. Water is fundamental to plants‘ existence, and in the tropics, is a key determinant of plant species‘richness, composition, growth and survival. There is thus an increasing interest in understanding how changing rainfall may cause functional changes in forests or change their species composition. Therefore, the overarching goal of thisdissertation was to understand the impact of water variability on tropical forest tree growth and vulnerability to drought.Forest tree growth along spatial and temporal rainfall gradientsObservational studies that measure whole forest tree growth along spatial or temporal gradients of rainfall are the most common way of formulating forest growth response curves to water availability, when manipulative experiments are cost-prohibitive or impractical (fire or large mammal disturbance). In the tropics, since very few species show anatomically distinct tree rings, estimating tree growth from trunk diameter is the standard practice to obtain growth patterns across species. However, this method—of equating woody growth to diameter change--is susceptible to bias from water-induced stem flexing. In the absence of bias correction, temporal variability in growth is likely to be overestimated and incorrectly attributed to fluctuations in resource availability, especially in forests with high seasonal and inter-annual variability in water. This problem has been largely ignored in the absence of any corrective measure and due to under-appreciation of the magnitude of error. While diameter re-censuses in permanent sampling plots (PSPs) have been most commonly done at 3-5 year scale (using a graduate tape), increasingly they are done at seasonal and annual scales (using band dendrometers) to closely match variation in rainfall, the scales at which hydrostatic bias may be greater in magnitude relative to woody growth. Besides, along a spatial rainfall gradient, inter-annual variability in water may vary, causing systematic differences in the hydrostatic bias for forests along the gradient. Therefore, one broad objective of this thesis was to evaluate the problem of hydrostatic bias in whole forest growth-rainfall relationship at annual and supra-annual scales, for temporal as well as spatial rainfall gradients and propose and test a novel corrective solution.Further, it also examines if growth-diameter relationship vary along the spatial gradient, which it may arise due to differences in light environments and/or disturbance history and species composition. The missing link of Eco-hydrology Differential responses of tree species in terms of growth and survival to variation in water that they can access, the proximate cause is likely shaped through their life-history strategies, the ultimate cause. However, we neither know the depths at which the diverse tree species in a forest draw water from and its dynamics, nor variation in water at those depths vis-à-vis rainfall patterns—for lack of appropriate methods. This has been a key missing link in understanding how water shapes trees‘ life-history strategies, their demographic trade-offs and co-existence, and also our predictive ability to determine species-specific responses to changing rainfall patterns, especially droughts. Since droughts are highly stochastic events and trees‘ responses to their drought ―experiences‖ may be revealed at decadal scales, long-term evaluations are key. Therefore, the second broad objective of this thesis was to develop a framework to determine trees’ water uptake depths, variation in water availability at those depths and trees’ demographic responses over multiple decades. From this, to understand how belowground hydrology shapes drought-vulnerability, demographic trade-offs and coexistence of forest tree species. This thesis titled—Eco-Hydrology of a Seasonally Dry Tropical Forest: Tree Growth, Belowground Water Dynamics and Drought-Vulnerability—is organized as follows: Chapter 1 lays down an introduction to the thesis, followed by a description of the study site and datasets used in the thesis in Chapter 2. This thesis uses a variety of methods and multiple datasets, all of which are from the protected Seasonally Dry Tropical Forests of the Western Ghats in southern India in the Mudumalai and Bandipur National Parks. It is then followed by three data chapters: Chapter 3 describes the seasonal fluctuations in a five year long (1980-1985) tree diameter time series (using dendrometers) of a Seasonally Dry Tropical Forest in Bandipur National Park to illustrate the issue of hydrostatic stem-flexing. It investigates the possibility that band dendrometers may themselves underestimate stem shrinkage at diurnal or seasonal scale. It also evaluates if there could be a best season and time of the day for undertaking forest diameter censuses that can minimize hydrostatic bias. Chapter 4(published in Forest Ecology and Management)measures the hydrostatic bias in a sample of trees in a 50 ha PSP of a Seasonally Dry Tropical Forest in Mudumalai National Park, and proposes a novel way to correct this bias at the whole community level in the 20 year long 4-year interval growth time series. Chapter 5 (in review with Environmental Research Letters) investigates and presents two new confounding factors in growth-rainfall relationships along a spatial rainfall gradient: hydrostatic bias and size-dependency in growth rates. For this it evaluates forest tree growth estimates in seven 1-ha PSPs (~800 trees, 3-year annual time series 9using dendrometers) along a 1000 mm rainfall gradient spanning a mesic savanna-moist forest transition in Mudumalai National Park. Using the period for which seasonal diameter time series was available (2 yrs), it evaluates if the extent of seasonal fluctuations systematically vary along the gradient—most likely due to hydrostatic stem flexing. It also describes the presence of an anomalous size-diameter relationship in the mesic savanna from a large plots (50 ha PSP, diameter records using graduated tape). These observations are then used to draw insights for ―space for time‖ substitution modeling. Chapter 6 (in prep for Nature Plants) analyses belowground water environments of trees over two decades (1992-2012), a period that includes a prolonged and intense drought, in the 50 ha PSP of a Seasonally Dry Tropical Forest in Mudumalai. It uses a locally parametarised dynamic hydrological model in which site rainfall is also a forcing variable. It then develops a novel dynamic growth model and inversely estimates water uptake depths for adult trees of all common species (include ~9000 trees) in the PSP from their above-ground growth patterns over two decades vis-à-vis belowground water availability at multiple depths. It then examines if species‘ water uptake depth obtained thus is a predictor of their drought-driven mortality. Finally, this is used to evaluate the hydrological niche partitioning tree species operate under and how that drives their water uptake strategies, demographic trade-offs, and drought-vulnerability. Summarizes the thesis and suggests future directions

The tropical forests on the island of Borneo are among of the richest in the world in terms of tree diversity, and their capacity to store a large reservoir of carbon. The Southeast Asian forests are fundamentally different from Neotropical and African forests, with their single-family dominance by dipterocarp trees, and with inherently greater stature and biomass. The carbon productivity and allocation in Asian tropical forests is still poorly quantified, and their responses to environmental drivers are still poorly understood. Almost all recent advances in tropical forest carbon cycling research have occurred in the Neotropics, with very few studies in Asia. The principal aim of this thesis is to quantify the carbon budget of a lowland dipterocarp forest in the Lambir Hills National Park, Miri, Sarawak, Malaysian Borneo. I examined and explored the productivity and carbon cycling processes and their responses to environmental factors across two major and contrasting soil types, in particular the clay and sandy loam soils. I recorded and analysed the Net Primary Productivity (NPP) and respiration for the above- and below-ground components, and observed the responses to seasonal variation and environmental drivers. Total soil respiration was relatively high and contributed a great deal to ecosystem respiration. Variation in soil respiration rates appeared closely related to soil moisture content. I found a strong diurnal cycle in soil respiration. On the basis of the first soil carbon dioxide (CO2) efflux partitioning study undertaken in a tropical forest, the diurnal cycle in total soil respiration appeared to be entirely driven by the diurnal cycle in litter respiration, and in turn litter is strongly controlled by moisture. There was little seasonal variation in allocation of net primary productivity (NPP), but there was evidence showing potential inter-annual variability for several components of NPP. Further, the allocation of NPP showed a strong seasonal shift between the forest plots on clay and sandy loam soils. Combining all the data measured and obtained in this D.Phil. thesis, the overall carbon budget assessed in this lowland dipterocarp forest showed a high level of agreement with other studies in Asia using micrometeorological techniques and the situation appears to be comparable to tropical forests in Amazonia. The key difference is that the aboveground NPP is higher and is the largest component contributing to the overall carbon budget, with relatively higher carbon use efficiency (CUE). The lowland dipterocarp forest in Lambir shows higher allocation in the above-ground NPP, and there were also differences in NPP and its allocation between sandy and clay-rich plots.

The trait based approach has been proposed as a way to reconcile community ecology. Despite recent advances in trait based ecology, such as the development of global trait databases and standardized methodology for trait collections, it remains unclear to what degree traits vary across individuals, species, and communities. In addition, the drivers of trait variation may shed light on the underlying processes that maintain species diversity and community assembly at local to continental scales yet these have been poorly studied. In this study, I examine both the magnitude of trait variation as well as the patterns of trait variation at local to continental scales in order to understand the drivers of diversity patterns across environmental gradients. First, I quantified the magnitude of trait variation at local scales in a dry tropical forest and determined that intraspecific variation is not negligible and can be quite large for compound-leaved species. However, I showed that the sample sizes necessary for quantifying trait variation are tractable and should encourage the adoption of trait variation in trait based ecology. Second, I tested whether climatic variables are predominantly responsible for observed trait variation across dry tropical forests in the Americas. I showed that climatic variability, specifically variability in precipitation, explained a large degree of observed trait variation across dry tropical forests and may provide a unique approach for classifying dry tropical forests based on their inherent degree of climatic seasonality. Third, I quantified patterns of trait variation at continental scales across elevational gradients at high to low latitudes. I showed that climatic variables largely drive patterns of trait variation at high latitudes while biotic factors largely drive patterns of trait variation at low, tropical latitudes. This finding has implications for understanding large-scale patterns of species diversity across elevational and latitudinal gradients. Finally, I apply trait variation to life history theory by quantifying variation in two life history traits (growth and reproduction) in a tropical tree species using a legacy dataset. I showed that variation in these two life history traits is due to both resource availability and allometric related effects on both traits. In sum, this study advances our understanding of the magnitude and underlying drivers of trait variation at local to continental scales.