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Genetic variation in Calamus subinermis was investigated in a provenance cum progeny trial and a study on some quantitative morphological traits. These studies show that variation occurs within and among populations of the species. Genetic gain was estimated for provenance and family selection at a range of selection intensity based on some economic traits. A karyomorphological study on C. subinermis and C. caesius reveals little variation in the chromosome morphology between and within species. The flowering and fruiting phenology of C. subinermis, C. caesius, C. manan and C. trachycoleus was elucidated. Except for the relatively opportunistic C. subinermis, in which floral initiation occurred at about four-month intervals, flowering and fruiting of the Calamus species investigated exhibited a regular annual pattern. Unravelling of the pollination mechanism and floral biology of C. subinermis and C. caesius in this project enables understanding of the breeding system and is important in breeding programmes. Pollination mainly by small noctuid and pyralid moths, which have small foraging ranges, partly accounts for the observed divergence among geographically isolated populations of C. subinermis. The discovery of andromonoecy in C. subinermis, together with the potential bisexuality in the other generally dioecious calamoid genera, calls for modification in the present system of classification for the tribe Calameae, and has implications for natural and artificial pollination. Several rattan mensuration techniques were developed. Growth and yield models based on the Richards function were constructed for C. caesius. These techniques and models will contribute to the conservation with wise use of rattan resources. Recommendations for further research are given for attaining the goal of the project.

The stretch from the riparian zone of a major river, through the mouth and upstream in tributaries forms a range of differing habitats. The aim of this study was to investigate the effect of these different habitats on the riparian frog community in primary rain forest in Borneo; i.e. examine whether any gradients could be demonstrated in terms of species composition, diversity and density of frogs, as well as which environmental characteristics that seem to be of most importance in forming habitats of high conservational value. A major river, Segama, and three of its tributaries were investigated. The surveys were carried out at night by searching for frogs visually and acoustically along the streams, from the shore of the Segama river and 400 metres upstream in the tributaries. In all transects 10 habitat variables were also measured. The results from the study show a significantly lower diversity and abundance of frogs along the shores of Segama and close to the mouths of the tributaries compared to further upstream, and a significant difference in species assemblage. Most species exhibited a negative correlation with the downstream transects or were not found there at all. The habitat characteristics waterfalls and boulders were found to be of most importance for the diversity and abundance of frogs, making these characteristics key components of habitats with high conservational value. Major rivers could also potentially form dispersal corridors for invasive species, which findings of the introduced species Fejervarya limnocharis along the shores of Segama shows.

京都大学 / 0048 / 新制・課程博士 / 博士(農学) / 甲第21470号 / 農博第2313号 / 新制||農||1064(附属図書館) / 学位論文||H31||N5165(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 本田 与一, 教授 舟川 晋也 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DGAM

As florestas tropicais representam grande parte do carbono armazenado na forma de biomassa. São caracterizadas por uma alta taxa de produtividade primária, no entanto, não é muito claro o entendimento de como as florestas tropicais respondem ao balanço de carbono com o avanço das mudanças climáticas. Um dos efeitos relacionados ao aumento de concentração de dióxido de carbono na atmosfera e ao aumento da temperatura ambiental, que pode contribuir para ocorrência de distúrbios na função de assimilação ou emissão deste composto dentro das florestas tropicais, é a temperatura. A temperatura tem sido apontada, como influenciadora nas trocas gasosas das plantas, afetando a fotossíntese e a respiração. Neste sentido, estudos que visam melhorar o entendimento dos processos fisiológicos das plantas, principalmente os que estão envolvidos com a ciclagem do C, são importantes para gerar informações sobre os efeitos das mudanças globais nos padrões de respostas da assimilação de CO2 das plantas ou ecossistemas. O objetivo deste trabalho foi determinar as respostas dos processos de fotossíntese e respiração em função da temperatura foliar de um indivíduo e comunidade em estudo. O trabalho foi desenvolvido na Floresta Nacional (FLONA) dos Tapajós (2º 51 Sul (S), 54º 58 Oeste (O)) localizada ao sul da cidade de Santarém, próximo ao marco km 67 e 83 da rodovia Santarém-Cuiabá BR-163. Para realização das medidas nas folhas das plantas na copa do dossel florestal foram utilizadas cinco torres. Para quantificação da fotossíntese e respiração foliar, utilizou-se um analisador de gás por infravermelho (IRGA) modelo (LI- 6400). Para cada folha observada, obteve-se uma curva de resposta da fotossíntese à radiação fotossinteticamente ativa (A-RFA) e quatro curvas de respostas da fotossíntese em relação a concentração interna de CO2 (A-Ci), em quatro grupos de temperaturas foliares (Tfoliar). As curvas de resposta a radiação fotossinteticamente ativa (RFA), Tfoliar e concentração interna de CO2 (Ci) foram realizadas entre os horários de 07:00 e 14:00 horas. Para o ciclo diurno quantificou-se a fotossíntese (A), condutância estomática (gs), transpiração (E) e déficit de pressão de vapor da folha (DPVf), as medidas foram realizadas ao longo de um dia entre 10:00 e 18:00 horas. A temperatura ótima média da fotossíntese foi de 29,9 oC, e a população apresentou taxa assimilatória líquida ótima de CO2 de 7,2 µmol.m-2.s-1. Valores de temperaturas foliares superiores a 32,9 oC e 32,5 oC, diminuíram a capacidade máxima de transporte de elétrons e a velocidade máxima de carboxilação, respectivamente. A eficiência de carboxilação começou a diminuir a partir de 31 oC. A concentração interna de CO2 diminuiu 32,6% aos 42 oC. Na avaliação do ciclo diurno A e gs decresceram significativamente a partir das 16:00 e 15:00 horas, respectivamente. Quando houve um aumento de 0,34 kPa no DPVf a transpiração diminuiu em 64%. A condutância estomática decresceu com o aumento da temperatura foliar e a respiração foliar foi positivamente relacionada com o aumento da temperatura foliar. / Tropical forests hold large stores of carbon as biomass. They are characterized by large net primary productivity, however, it is not well understood how carbon budgets of tropical forests respond to the advance of climatic changes. Temperature is one of the agents related to the increase of carbon dioxide concentration in the atmosphere and ambient temperatures can contribute to alterations in assimilation or emission of CO2 in tropical forests. Temperature has been pointed as responsible for gas exchange rates in plants, affecting the photosynthesis and respiration. Thus, studies aimed at better understanding of physiological processes of plants, particularly those which involve carbon cycling, are important to generate information on the effects of the global changes in the trends of CO2 assimilation response in plants or in ecosystems. The objective of this work was to determine the photosynthesis processes and respiration responses as a function of leaf temperature of an individual and of the community in study. The study was carried out at the Tapajós National Forest (FLONA, 2º 51` S 54º 58` W) located in the south of Santarém municipality, at kms 67 and 83 of the Santarém- Cuiabá highway (BR-163). Five towers were used to support measurements in leaves in the forest canopy. An infra-red gas analyzer (IRGA, LI-6400 model) was utilized for quantification of leaf photosynthesis and respiration. For each observed leaf, a curve relating the photosynthesis response to the photosynthetically active radiation (PAR) was obtained together with four curves of photosynthesis response in relation to the internal CO2 concentration (Ci) in four groups of leaf temperatures (Tleaf). The response curves of PAR, Tleave and internal CO2 concentration (Ci) were obtained between 07h00 and 14h00. For the diurnal cycle it was quantified photosynthesis (A), stomatal conductance (gs), transpiration (E) and vapor pressure deficit of the leaf (VPDf), and measurements were carried out throughout one day between 10h00 and 18h00. The main optimum temperature of photosynthesis was 29.9 °C, and the community presented optimum CO2 net assimilation rate of 7.2 µmol.m-2.s-1. Leaf temperature values above 32.9 °C and 32.5 °C decreased the maximum capacity of electron transport and the maximum speed of carboxylation, respectively. The carboxylation efficiency starts to decreased at 31 °C. The internal CO2 concentration diminished 32.6% at 42 °C In the evaluation of the diurnal cycle, photosynthesis and stomatal conductance decreased significantly after 16h00 and 15h00 hours, respectively. An increase of 0.34 kPa in VPDf diminished transpiration in 64%. The stomatal conductance decreased with increase of leaf temperature and leaf respiration was positively related to the increase of leaf temperature.

O papel da Floresta Amazônica na ciclagem do carbono tem sido`freqüentemente relatado, no entanto, pouco se sabe sobre os aspectos que regula nos processos de assimilação e liberação de carbono da biosfera para a atmosfera. O objetivo deste trabalho foi descrever as respostas dos processos de respiração e fotossíntese com a variação da temperatura foliar, utilizando características biofísicas e dados micro-meteorológicos, para predizer a taxa assimilatória de CO2 de um indivíduo ou da comunidade em estudo. A pesquisa foi conduzida na Estação Experimental de Silvicultura Tropical (núcleo ZF-2), área de pesquisa da Coordenação de Pesquisa de Silvicultura Tropical (CPST) do Instituto Nacional de Pesquisas da Amazônia (INPA), utilizando-se de 4 torres para o acesso as copas. Os elementos climáticos foram monitorados por estações micro-meteorológicas Li-1401. As medidas de temperatura foliar foram realizadas com o auxilio de termopares de cobre-contantan, sendo a fotossíntese e a respiração avaliadas com um analisador de gases por infravermelho, com o qual foram obtidas curvas de respostas da fotossíntese a variação de radiação fotossinteticamente ativa (A-RFA), e a variação da concentração interna de CO2 em 4 diferentes temperaturas (A-Ci-T). Os resultados mostraram que: a temperatura foliar está relacionada com a radiação fotossinteticamente ativa (RFA) e a umidade relativa do ar (UR); a temperatura ótima da fotossíntese foi de 31,1 oC, onde a comunidade apresentou 7,6 µmol.m-2.s-1 de assimilação líquida de CO2, e a partir desta temperatura houve uma diminuição da fotossíntese máxima; valores de temperaturas foliares maiores que 34,5 oC diminuíram a velocidade máxima de carboxilação e as acima de 35,7 oC diminuíram a capacidade máxima de transporte de elétrons; a condutância estomática decresceu com o aumento da temperatura foliar, dentro do intervalo de temperatura estudado, indicando que a limitação estomática pode ser o fator que mais afeta a fotossíntese; e a respiração contribuiu para que o balanço de carbono fosse menor com o aumento da temperatura foliar devido a respiração estar positivamente relacionada com a temperatura foliar. A taxa assimilatória de CO2 foi diminuída devido a aumentos da temperatura foliar, sendo principalmente afetada pela diminuição da condutância estomática e do mesofilo e depois por aumentos da respiração foliar. / The role of the Amazon Rain Forest in the carbon budget has been often reported, however little is known about the aspects regulating the processes of carbon assimilation and its release from the biosphere to the atmosphere. The objective of this work was to describe the responses of respiration and photosynthesis processes related to leaf temperature variations by using biophysics characteristics and micro-meteorological data in order to predict CO2 assimilation rates on individual or community level. The research was conducted at the Experimental Station for Tropical Forest research (ZF-2) of the National Institute of Research of the Amazon (INPA). For this study four towers were used to access the canopy. Climatic parameters were monitored by micrometeorological stations Li-1401. The leaf temperature was measured using copper-contantan thermocouples. Photosynthesis and respiration were evaluated with an infra-red gas analyzer, generating light and CO2 curves responses. The results showed that: the leaf temperature was related to the photosynthetic active radiation (PAR) and to the relative air humidity (UR). The optimal photosynthesis temperature was 31,1 ºC where the tree community presented 7,6 µmol.m-2.s-1 of net CO2 assimilation. Above this temperature a reduction of the maximal photosynthesis was determined. Leaf temperatures higher than 34,0 ºC decreased the maximal velocity of RuBP carboxilation and temperatures higher than 35,7ºC decreased the maximal capacity of electrons transportation. The stomatal conductance decreased with increasing leaf temperatures within the temperature interval studied. The results indicated that the stomatal limitation probably represent the main factor that effects photosynthesis. The respiration contributes to reduce the carbon assimilation due to the increase of the leaf temperature because respiration is positively related to leaf temperature. Thus, the CO2 assimilation rates decreased with an increase of leaf temperature and is mainly affected by a decrease of stomatal and mesophyll conductance and by an increase of leaf respiration.

La biodiversité des forêts tropicales humides (FTH) est inégalement répartie dans les trois principales régions des tropiques : les Néotropiques, l’Asie du sud-est et l’Afrique tropical. Même si les forêts africaines contiennent une diversité en espèces végétales et un endémisme importants, leur richesse spécifique reste faible par rapport aux autres massifs forestiers tropicaux. Cette disparité serait liée à l’impact négatif d’événements d’extinction durant le Cénozoïque plus intenses dans les forêts africaines. A l’échelle du bassin du Congo, les niveaux de diversité important et l’endémisme des forêts africaines seraient liés à l’hypothèse d’existence de forêts refuges du passé. Cette hypothèse suggère une contraction des forêts en ‘refuges’ durant les événements climatiques du Dernier Maximum Glacier (DMG, entre 24’000 et 12’000 Ans).L’objectif principal de cette thèse est d’étudier l’évolution des forêts africaines en réponse aux changements climatiques sur des pas de temps anciens (millions d’années) et plus récent (milliers d’années). Pour cela nous étudions deux groupes de palmiers Africains : les rotins de la sous-tribu des Ancistrophyllinae et le genre de sous-bois des FTH d’Afrique Centrale Podococcus composé de deux espèces P. barteri et P. acaulis. Le premier groupe nous renseigne sur les événements d’extinctions anciens durant le Cénozoïque, alors que le second permet d’étudier la réponse des FTH face aux changements climatiques du Pléistocène. L’approche utilisée dans cette thèse consiste à 1) reconstruire l’histoire évolutive des rotins, et 2) inférer la dynamique évolutive des deux espèces de Podococcus dans le temps. Cinq objectifs spécifiques structurant deux grandes parties de la thèse ont été retenus. Première partie : (i) reconstruire les relations phylogénétiques chez les Ancistrophyllinae à l’aide de marqueurs chloroplastiques et nucléaires, (ii) estimer les âges de divergences puis tester l’impact des événements d’extinction chez les Ancistrophyllinae. Seconde partie : (iii) construire les modèles de niches écologiques et inférer les zones de stabilité écologique des deux espèces de Podococcus depuis le DMG, (iv) reconstruire les relations phylogéographiques chez Podococcus à partir de plastomes entier obtenus avec du Séquençage Nouvelle Génération ‘NGS’ et (v) évaluer la concordance entre la diversité génétique et la stabilité écologique des populations de Podococcus.Les résultats suggèrent que les espèces d’Ancistrophyllinae se sont probablement diversifiées de manière constante mais avec un taux d’extinction assez élevé ponctué par un ou plusieurs événements d'extinction sévères au cours du Cénozoïque. Les Ancistrophyllinae ont divergé durant l’Eocène et la plupart des espèces sont apparues après 10 Ma durant la fin du Miocène et du début du Pliocène. Chez le genre Podococcus, une corrélation significative entre la diversité génétique unique et la stabilité écologique a été démontré appuyant l’hypothèse des forêts refuges. Les populations écologiquement stables et diversifiées sont principalement localisées dans les zones montagneuses comme les Monts de Cristal et les Monts Doudou au Gabon, mais aussi dans les forêts littorales atlantiques au Gabon et au Cameroun. Par contre la plupart des populations datent d’avant le DMG ce qui traduit une réponse des espèces sur plusieurs cycles glaciers. Les résultats de cette thèse montrent que les changements climatiques anciens et plus récents ont fortement influencé l’évolution et la dynamique des espèces forestières d’Afrique Centrale. De plus, nos résultats ont permis d’identifiés les forêts côtières comme prioritaires pour la conservation de la diversité en Afrique Centrale. / Tropical rain forests (TRF) biodiversity is not equally distributed between the three main tropical regions: Neotropics, South East Asia and tropical Africa. Even though African rain forests display high levels of diversity and endemism, this diversity is low compared to that of the other tropical regions. One of the main hypotheses advanced to explain this lower species diversity is that the African flora has undergone higher extinction rates during the Cenozoic. Across the Congo basin, high levels of species diversity and endemism is thought to be linked to the hypothesis of forest refugia. This hypothesis suggests a contraction of rain forests in 'refugia' during the Last Glacial Maximum (LGM, between 24’000 and 12’000 Years).The aim of this thesis is to study the evolution of African rain forests in response to climate change on ancient (million years) and more recent (thousands of years) time scales. For this we study two groups of African palms: the rattans of the sub-tribe Ancistrophyllinae and the understory genus of Central African TRF Podococcus with two species P. barteri and P. acaulis. The first group provides information about ancient extinctions events during the Cenozoic, while the second is used to study the response of TRF during Pleistocene climate change in Central Africa. The approach used in this thesis is to 1) reconstruct the evolutionary history of African rattans, and 2) to infer the evolutionary dynamics of the two species of Podococcus through time. The five specific objectives structuring two main parts of the thesis are: First part: (i) reconstruct phylogenetic relationships among Ancistrophyllinae using plastid and nuclear markers, (ii) estimate divergence times and test the impact of extinction events on Ancistrophyllinae. Second part: (iii) build ecological niche models and infer the areas of habitat stability of the two species of Podococcus from the LGM, (iv) reconstruct phylogeographic relationships of populations of Podococcus from whole plastomes obtained using Next Generation Sequencing and (v) assess the correlation between genetic diversity and ecological niche stability of all populations of Podococcus.Results suggest that Ancistrophyllinae has possibly undergone a constant diversification with a relatively high extinction rate punctuated by one or more severe extinction events during the Cenozoic. Ancistrophyllinae diverged during the Eocene with most species originating to the Late Miocene after 10 Ma. In the genus Podococcus, a significant correlation between the unique genetic diversity and habitat stability was demonstrated supporting the hypothesis of forests refugia in Central Africa. Ecologically stable and diverse populations are mainly located in mountainous areas such as Monts de Cristal and Monts Doudou in Gabon, but also in the Atlantic coastal forests in Gabon and Cameroon. In contrast, most of the populations predate the LGM reflecting that species respond over several glacial cycles. Results of this thesis highlight that ancient and recent climate changes have strongly influenced the evolution and dynamics of Central African forests. In addition, our results allowed the identification of African Atlantic coastal forests as an important conservation priority.

Made available in DSpace on 2016-04-28T14:56:37Z (GMT). No. of bitstreams: 1 2009 - Tokitika Morokawa.pdf: 5650089 bytes, checksum: a8d1865101e3a9effdd865554b069415 (MD5) Previous issue date: 2009-02-05 / A model of the growth dynamics of forests was developed applying Bertalanffy growth model (BGM) y=a[(1 e bt)c] for individual trees. Based on linearized relationship with diameter and height equation LnH=b0+LnD and volume equation V=b0Db1Hb2 mutually compatible equations for diameter, basal area, height and volume were generated, adjusting parameters a and c of BGM. Additionally, it was developed a methodology based on DBH measurements in two occasions and the estimate value of parameter a to get the values of parameter c and b of BGM. The data of the inventory of 1,061 trees over 40cm DBH of 67.5 hectares of tropical rain forest of Aquariquara Extractive Reserve, located in State of Rondonia, Brazil, were used to test the developed model. These data were grouped in nine maximum diameter (Dmax) classes. The tree survival was estimated based on the frequency of trees by relative age class. The input and output of each tree was adjusted considering a closed and stable population with replacement of a dead tree by the ingrowth of the same specie of tree with 40cm DBH. Applying the model to such data and simulating the growth dynamics of the forest in a time frame of 1500 years (T1 → 1,500), the difference of simulated annual average, in relation to the observed values in the year T0 of the forest inventory was -2.09% in age, -0.33% in height, -0.96% in diameter, -3.41% in basal area, -3.81% in volume and +1.81% in merchantable trees value. The average values of T1→500, T501→1000 and T1001→1500 periods were equal for all variables by Kruskal-Wallis test (p=0.05). The simulation showed that one hectare of this forest maintains a total stock composed of 15.72 120-year-old trees, accumulated 4.35m2 of basal area and 37.98m3 wood volumes on average. The annual input flow of the system was 0.3457 88-year-old trees which accumulated 0.0441m2 of basal area and volume of 0.3265m3. The output of the system consists of 0.3459 132-year-old trees totaling 0,1127m2 of basal area and 0.9911m3 in volume. The annual input and output rates were, respectively, 2.20 and 2.20% in number of trees, 1.01 and 2.59% in basal area and 0.86 and 2.61% in volume. In addition, the simulation indicated that it is possible to sustain an average annual production of 0.5528m3 of merchantable timber generating R$5.50 (about US$1.49) annual income per hectare of forest. The simulation showed that the model developed describes growth dynamics of tropical forest. However, it needs to be properly adjusted based on real growth of each individual tree and flows of input and output of trees in the system. / Nesta tese foi desenvolvida a modelagem da din?mica de crescimento de uma floresta, aplicando-se o modelo de crescimento de Bertalanffy (MCB) y=a[(1 e bt)c] para ?rvores individuais. Baseando-se nas rela??es linearizadas entre o di?metro e altura da equa??o LnH=b0+LnD e do di?metro com volume V=b0Db1Hb2 foram geradas equa??es de crescimento reciprocamente compat?veis entre di?metro, ?rea basal, altura e volume, ajustando-se os valores dos coeficientes a e c do MCB. Adicionalmente, foi desenvolvida metodologia, baseando-se em medi??es de DAP de cada ?rvore em duas ocasi?es e com a estimativa do coeficiente a, para obter os valores dos coeficientes c e b do MCB. Para testar o modelo desenvolvido foram utilizados numa simula??o os dados de invent?rio de 1.061 ?rvores, com DAP≥40cm, de uma ?rea de 67,5ha de Floresta Omb?fila Aberta da Reserva Extrativista Aquariquara, localizada no munic?pio de Machadinho D?Oeste, estado de Rond?nia. Estes dados foram agrupados em nove classes de di?metro m?ximo (Dmax) compreendidas entre 50 e 240cm, mantendo-se uma amplitude entre elas de 10cm. A taxa de sobreviv?ncia da ?rvore foi estimada baseando-se na freq??ncia de ?rvores por classe de idade relativa. O sistema de entrada (input) e sa?da (output) de cada ?rvore foi ajustado considerando uma popula??o fechada e est?vel com substitui??o de uma ?rvore morta pela entrada de ?rvore da mesma esp?cie com DAP≥40cm. Em rela??o aos valores observados no ano T0 do invent?rio florestal, as m?dias anuais dos valores simulados da din?mica de crescimento dessa floresta num horizonte temporal de 1.500 anos (T1→1.500), apresentaram diferen?as m?dias de -2,09% em idade, -0,33% em altura, -0,96% em di?metro, -3,41% em ?rea basal, -3,81% em volume e +1,81% em valor da ?rvore em p?. As m?dias anuais dos valores simulados dos per?odos T1→500, T501→1.000 e T1.001→1.500 foram iguais para todas as vari?veis pelo teste de repetibilidade de Kruskal-Wallis (p=0,05), mostrando que o modelo ? est?vel no tempo. A simula??o mostrou que esta floresta mantem em m?dia um estoque total de massa em crescimento composta de 15,72 ?rvores.ha-1 com 120 anos de idade acumulando um total de 4,35m2.ha-1 de ?rea basal e 37,98m3.ha-1 de volume de madeira. O fluxo anual de ingresso no sistema foi de 0,3457 ?rvores com 88 anos de idade que somaram 0,0441m2 de ?rea basal e 0,3265m3 de volume, e de sa?da do sistema composto de 0,3459 ?rvores com 132 anos de idade que somaram 0,1127m2 de ?rea basal e 0,9911m3 de volume por hectare, resultando em taxas de ingressos e sa?das anuais, respectivamente, de 2,20 e 2,20% em n?mero de ?rvores, 1,01 e 2,59% em ?rea basal e 0,86 e 2,61% em volume. Al?m disso, a simula??o indicou que ? poss?vel sustentar uma produ??o m?dia anual de 0,5528m3 de madeira de valor comercial gerando renda anual de R$5,50 por hectare (a pre?o corrente de outubro de 2002) de floresta. A simula??o mostrou que o modelo desenvolvido descreve a din?mica de crescimento de floresta tropical, por?m ele necessita ser devidamente calibrado tomando os dados de crescimento real de cada ?rvore individual e dos fluxos de ingressos e sa?das de ?rvores do sistema.