京都大学 / 0048 / 新制・課程博士 / 博士(農学) / 甲第20437号 / 農博第2222号 / 新制||農||1049(附属図書館) / 学位論文||H29||N5058(農学部図書室) / 京都大学大学院農学研究科地域環境科学専攻 / (主査)教授 北山 兼弘, 教授 神﨑 護, 教授 北島 薫 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DFAM

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Expanding anthropogenic development within the tropical forest biome is driving the loss of an irreplaceable global resource. Mega-diverse tropical forests are vital for regulating the global carbon cycle, and are essential for climate change mitigation. Today, over half of the world’s remaining tropical forest is degraded or regenerating secondary forest. Tropical forests are becoming increasingly fragmented through the expansion of agriculture and roads. Landscape-scale flooding of terrestrial habitats caused by dam construction is an emerging driver of habitat loss and fragmentation. Much attention has been paid to the long-term impacts of tropical forest fragmentation for biodiversity, ecosystem functioning, and carbon emissions. Most of our understanding of the impacts associated with habitat fragmentation originates from systems in which the habitat matrix surrounding remnant forest patches is another, albeit low quality, terrestrial habitat. However, dam-induced habitat fragmentation results in remnant terrestrial biological communities becoming isolated on islands within a water matrix. A water matrix presents the worst-case scenario for remnant habitat fragments. In Chapter 2 I synthesise the results of numerous studies reporting the responses of taxonomic groups to isolation on reservoir land-bridge islands, and uncover a globally-applicable pattern of extinction debt acting upon remnant biological communities on reservoir islands. All islands, regardless of taxonomic group, habitat type, or island area lose species as island isolation time increases. Moreover, I show that contrary to existing ecological theory, once terrestrial habitat becomes isolated within a water matrix, it is effectively too isolated for species losses to be buffered by metapopulation dynamics. Dam development is rapidly expanding in the largest remaining tract of intact tropical forest, the Amazon Basin. In Chapters 3 and 4 I study the Balbina mega-dam system in the central Brazilian Amazon. Here, I use detailed field inventories of trees and lianas on islands and in continuous mainland habitat to determine the impact of landscape-scale habitat fragmentation caused by reservoir creation on these taxonomic groups. I find that islands maintain tree communities at significantly lower densities, richness and diversity compared to continuous forest. Furthermore, tree communities on islands exhibit compositional divergence from those found in mainland continuous forest. Island tree assemblages are dominated by low-wood density species, and may be on a trajectory towards communities characteristic of early successional forests with reduced carbon storage capacity. In contrast, liana assemblages remain compositionally intact and are becoming increasingly dominant relative to trees. Thus, lianas appear robust to many of the negative impacts associated with landscape-scale habitat fragmentation. As insular tree communities continue to degrade through area- and edge-effects, lianas may become a key feature of this archipelagic landscape due to their competitive advantage over trees in disturbed forest habitats. Lianas significantly inhibit tree recruitment and carbon storage. Thus, findings from Chapters 3 and 4 provide strong evidence for additional, and currently unaccounted-for biodiversity and carbon impacts associated with tropical dams. As development of tropical forest regions increases, there is an urgent need to reconcile the need for resources with the need for ecosystem service provision, such as carbon storage, particularly as we attempt to mitigate the impacts of rising atmospheric carbon. Recent studies have shown that secondary tropical forests have the potential to rapidly uptake atmospheric carbon, and act as a powerful tool in climate change mitigation policy. Broad-scale estimates of secondary forest carbon uptake are currently based on above-ground biomass alone. In Chapter 5 I present carbon stock estimates of additional tropical forest carbon pools - soil and dead woody biomass - in secondary forests ranging from 40-120 years. I find that soil fertility (nitrogen concentration) is key in determining carbon storage in secondary forests, and that the stability of carbon stocks held in dead woody biomass increases with secondary forest stand age. I highlight the need to integrate detailed site-specific information into broad-scale predictive models of secondary tropical forest carbon sequestration. This thesis links ecological theory and landscape-scale field inventories, to provide new understanding of the long-term costs of tropical forest fragmentation for biodiversity conservation and carbon storage, and provides further evidence of the important role secondary tropical forests may play in carbon sequestration and climate change mitigation.

La compréhension de la coexistence des espèces végétales et leur l’assemblage dans les communautés naturelles reste un des objectifs fondamentaux de l’écologie, et c’est un enjeu majeur dans le contexte actuel des changements globaux. Ce travail de thèse a consisté à apporter des éléments de compréhension quant aux effets des facteurs abiotiques et biotiques pour l’assemblage des communautés d’arbres en milieu tempéré. Ces questions nécessitent d’être considérées sur le long terme, ce qui est particulièrement difficile en milieu forestier du fait de leurs dynamiques lentes. C’est pourquoi nous avons utilisé ici une approche originale basée sur un modèle mécaniste de dynamique forestière. Dans la filiation des études empiriques basées sur les traits fonctionnels en écologie des communautés, nous avons cherché à identifier les signatures fonctionnelles du filtre environnemental et de la compétition dans l’assemblage des communautés forestières, et ce le long d’un gradient de productivité en Europe centrale. Nos résultats montrent que le filtre environnemental est un déterminant majeur de la structure des communautés en restreignant la gamme de variation des traits, avec des effets plus marqués dans les milieux très contraignants et peu productifs. Par ailleurs, nos résultats montrent que la compétition pour la lumière mène à une convergence de la structure fonctionnelle des communautés, convergence d’autant plus forte que les milieux sont productifs. Dans le but d’approfondir la compréhension des mécanismes d’assemblages et de coexistence, nous avons montré qu’une augmentation de la variabilité climatique interannuelle n’aboutit pas systématiquement à une perte de diversité, et pourrait même favoriser la coexistence en modifiant le pouvoir compétitif relatif des espèces au cours du temps. Enfin, nous avons montré que les différences de valeurs de traits des espèces d’arbres pouvaient favoriser leur coexistence via des réponses différentielles des espèces à la variabilité climatique interannuelle et aux conditions de lumière. L’ensemble de ces résultats apporte de nouveaux éléments quant à la compréhension de la structuration des communautés forestières, et propose une vision intégrative des processus complexes d’assemblage et de coexistence au sein des communautés. / Understanding the coexistence of plant species and their assembly within natural communities remains one of the fundamental goals of ecology, and this major challenge appears even more crucial in the current context of global change. The aim of this work was to better understand how the abiotic and biotic factors determined tree communities’ assembly in temperate forests. These questions can only be considered on the long term, which is especially difficult for forest ecosystems because of their slow dynamics. Therefore we used an original approach based on a forest dynamic model. Following trait-based empirical studies in community ecology, we tried to identify the functional signatures of environmental filtering and competition in forest communities along a productivity gradient in Central Europe. Our results showed that environmental filtering is a major determinant of community structure as it reduced the trait range all along the gradient, with stronger effects in harsh sites with low productivity. Further looking for identifying underlying causal processes, we showed that competition for light led to a convergence of functional community structure, and that this convergence increased with site productivity. Furthermore, our results suggested that inter-annual climatic variability does not systematically result in biodiversity loss, and could even promote species coexistence through changes in the hierarchy of competitors over time. Finally, we showed that differences in trait values of tree species could promote coexistence through differential species responses to inter-annual climatic variability and to light conditions. These findings bring new insights regarding the understanding of tree communities’ structure, and they provide an integrative understanding of the complex processes driving community assembly and species coexistence in natural forests.

Tree growth influences forest community dynamics and responses to environmental variations, but currently is not well understood. Tree growth in highly diverse wet tropical forests have been well studied and characterised compared to the species-poor dry tropical forests. Thus, it is not clear if growth rates and community dynamics of dry forests are similar to those of wet forests, given the longer dry season, greater rainfall variability, more open canopy and lower number of species in dry forests. This thesis focuses on identifying important factors that influence tree diameter growth rates in the dry tropical forest at Mudumalai, southern India, and also compares growth patterns at this dry forest with those at moister forests. The thesis thus contributes towards closing the gap in understanding of tree growth patterns across the tropics. An initial analysis involving matrix-based population projections of four common canopy species at Mudumalai showed that variations in diameter growth have the potential to drastically modify population trajectories of dominant species. Thus the main focus of this thesis is aimed at identifying the important intrinsic and extrinsic factors affecting growth in this dry forest, as this information could be useful for future management of the forest. The second important aim of the thesis was to find out if growth rates are influenced by different sets of factors in tropical dry versus moist forests. A large permanent 50ha plot vegetation monitoring plot was set up in 1988-89 in the Mudumalai dry deciduous forest, and was subsequently monitored annually by staff of the Centre for Ecological Sciences. Data used in this thesis represent a 12-year interval between 1988 and 2000. Girth measurements were obtained from all woody tree stems ≥1cm in diameter every four years during this 12 year interval, which provided three census intervals of diameter increment data on >13,000 trees. For the comparison between dry and moist deciduous forests, data were obtained from a similar large plot maintained and monitored at the Barro Colorado Island (BCI) in Panama. Influences of the intrinsic factors, tree size, individual identity, species identity and growth form, were examined using t-tests, Wilcoxon signed ranks tests, linear regressions, analysis of variance (ANOVA), principal components analysis (PCA) and cluster analysis. Among the intrinsic factors tested, species identity explained approximately 20% of growth rates at the community level, while tree diameter explained less of growth variation, and growth form had a minor influence on growth. Growth rates also were examined for variations across the three census intervals, and for relationships with rainfall and survival from fire. Statistical tests included t-tests, Wilcoxon and other non-parametric sign tests, logistic regression and ANOVA. Most species and individuals showed significant reductions of growth in the second census interval (1992-1996), and growth rates of most trees were positively related to rainfall. Growth rate variations generally were not related to survival from fire, and few species were capable of escaping fire mortality by fast growth. Spatial environmental influences were tested in the commonest fifteen species, using five habitat categories, local elevation, slope, aspect, and the biotic neighbourhood variables of local conspecific and heterospecific density. Statistical tests included analysis of covariance, multiple linear regression and redundancy analysis. The tests were quadrat-based or individual-based, and species' growth responses were tested at different levels of distance and spatial scale. Topographic features and habitat categories had ephemeral effects on species growth. Only the most dominant species, Lagerstroemia microcarpa, showed consistent conspecific neighbour density effects. Redundancy analysis using a subset of common species and environmental factors did not reveal common growth responses to spatial environmental factors. Comparison of factors influencing growth at Mudumalai versus at BCI using multiple factor ANOVA and multiple linear regressions showed a similar influence of temporal variation at the two sites, but stronger and more widespread influence of tree size (diameter) at BCI. The greater influence of tree size at BCI may be related to greater light limitation in this dense moist forest. Spatial environmental factors had weak influences at both plots. Species were less differentiated from each other at the more diverse BCI plot compared to the relatively species-poor Mudumalai plot, suggesting that species' growth niches may be weakly related to diversity across tropical forests. Overall the results showed that among the factors tested species identity and census intervals were the most important influences on diameter growth at the Mudumalai dry deciduous forest. Tree diameter was less important and less consistent in affecting growth at the Mudumalai dry forest, contrary to expectations based on moist tropical forests where this relationship has been established previously. When comparing Mudumalai and BCI, the relative importance of different factors was different at the two sites, and the most important difference was a dominant influence of light limitation at the wetter forest in Panama. In terms of management applications, this study showed that fires at Mudumalai might be an inescapable source of mortality for many vulnerable species, and improved fire management is crucial for long term survival of species in this dry forest. At a larger scale, light and other environmental variables were found to influence growth differently at Mudumalai compared to BCI. This suggests that location-specific responses may be important for projections of tree biomass and carbon sequestration, especially under future climatic change scenarios.

Orientadores: Luiza Sumiko Kinoshita, Fernando Roberto Martins / Tese (doutorado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-15T14:38:51Z (GMT). No. of bitstreams: 1 Urbanetz_Catia_D.pdf: 5450468 bytes, checksum: 808c9a2f01bb127e923015f3c5db764d (MD5) Previous issue date: 2010 / Resumo: O objetivo principal desse trabalho foi investigar a influência do microrelevo, de nutrientes e de fatores granulométricos do solo na determinação da estrutura espacial da Floresta Ombrófila Densa Atlântica. Foram instalados três blocos de 40 parcelas em uma encosta, em solo argiloso e em cotas altitudinais distintas. Também foram instalados três blocos de 20 parcelas em uma planície, a 1,5 km da encosta, em intervalos de 1,5 km entre cada bloco na direção do mar e em solo arenoso. Foram marcados e medidos todos os indivíduos arbóreos com PAP ? 15 cm. A área total amostrada foi de 1,8 ha. Foram feitas análises de agrupamento e ordenação a partir de matrizes de abundância de espécies para verificar a presença de padrões, grupos ou gradientes. Foi aplicado o Mantel simples para testar a correlação entre uma matriz de dissimilaridades das parcelas (índice de Bray-Curtis e abundância de espécies) e uma matriz de dados abióticos (distância Euclidiana simples e nutrientes no solo, altitude e declividade). Foi aplicado o teste de Mantel parcial para verificar se existia correlação entre estas matrizes mesmo quando levadas em conta as distâncias geográficas. Foi feita a Análise Canônica de Correspondências (CCA) a partir das matrizes de abundância de espécies e dos dados abióticos para verificar a existência de relações entre as mesmas. Foram marcados e medidos 3415 indivíduos, pertencentes a 201 espécies e 55 famílias. Foi encontrada uma diferenciação clara entre a composição e distribuição de abundância de espécies da floresta da encosta e a da floresta da planície. Entretanto, ocorreram espécies comuns à ambas as formações. Provavelmente, as espécies comuns possuem uma plasticidade ecológica ampla e são capazes de tolerar as condições mais extremas de escassez de nutrientes e alagamento da planície. Os resultados da CCA e dos testes de Mantel simples e parcial sugerem que parte da diferenciação entre a floresta da encosta e a da planície provavelmente está relacionada com o teor de areia, Magnésio e pH do solo, declividade e altitude. Apesar disso, a maior parte da inércia encontrada não pôde ser explicada pelas variáveis abióticas mensuradas. A composição de espécies da planície não se mostrou uniforme, uma vez que o techo mais próximo do mar se diferenciou claramente dos demais. Os resultados da CCA indicaram que isso pode estar relacionado com um gradiente de areia no solo. A composição florística e a estrutura da floresta de encosta se mostrou mais uniforme do que a da planície. Não ocorreram correlações significativas entre a abundância de espécies na encosta e as variáveis abióticas na encosta quando foi levada em conta as distâncias geográficas. Possivelmente, eventos estocásticos tais como dispersão ou interferência antrópica ou outras variáveis bióticas e abióticas não mensuradas podem ter determinado a variação na estrutura espacial encontrada na encosta. O determinismo ecológico parece exercer um papel mais importante na escala da paisagem do que na escala local no presente estudo. Estudos complementares enfocando outros aspectos tais como dispersão, competição ou outras variáveis abióticas não mensuradas podem ser úteis para auxiliar no esclarecimento das principais causas dos padrões espaciais encontrados que não ficaram claras no modelo. / Abstract: The aim of this study was to investigate the influence of soil nutrients and microrelief on the variation in the spatial structure of the Atlantic Dense Ombrophilous Forest vegetation. Three blocks of 40 plots were established in a steep slope forest, a clay soil, at three altitudinal levels. Three other blocks of 20 plots were established in a lowland forest, in a poor sandy soil. Area "D" is 1,5 km from the steep slope, and areas "E" and "F" at further 1,5 km intervals in the direction of the coast. All the trees individual with PBH ? 15 cm were collected. The total area sampled was 1.8 ha. Classification and ordination analysis were applied to an abundance matrix in order to analyse the similarities among samples, patch structure or gradients. Floristic dissimilarities among plots (Bray-Curtis index computed with species abundance in each plot) were correlated with environmental distances among plots (Euclidian distance index computed with nutrients soil data, altitude and declivity in each plot) and geographic distances among plots (Euclidian distance index) through Mantel and partial Mantel tests. Canonical Correspondence Analysis (CCA) was applied to species abundance and abiotic data to detect species-environment relations. It was found 3415 individuals of 201 species and 55 families. A clear separation among plots of the steep slope forest and the lowland forest was found, although, there were species common to both vegetation types. The CCA and partial Mantel test results suggested that part of this spatial structure differentiation seems to be related to sand percentage, Magnesium and pH of the soil, declivity and altitude. Nevertheless, a large part of the inertia could not be explained by the abiotic data. The species common to both areas probably have a broad ecological plasticity and can tolerate the extreme condictions of poor nutrients avaiability and flooding of the lowland. The lowland forest flora was not uniform and two communities were recognised in that area. The CCA results indicate that these could be related with the texture gradient in the lowland soil. The floristic composition and the structure of the steep slope forest was more regular than the lowland forest. There were no correlation among species abundance and abiotic factors when geographic space was taken into account. Possibly stochastic events such as dispersion and human impacts or other unmeasured factors could determine the spatial structure found, together with to unexplained inertia. Ecological determinism seems to play a more important role at the landscape scale than at the local scale in this study. Studies focusing other aspects such as dispersion, competition or other unmeasured abiotic data could be helpful in clarifying the main causes of the unexplained patterns found in this study. / Doutorado / Biologia Vegetal / Doutor em Bilogia Vegetal