Cette étude a pour objectif la compréhension du fonctionnement des géosystèmes d'éboulisation présents dans l'espace supraforestier de la haute montagne des Pyrénées orientales. Quel est le degré d'activité actuel de ces versants d'éboulisation et comment évoluent-ils ? Quelles relations existe-t-il entre les espèces végétales lithophiles qui les colonisent et les dynamiques morphogéniques ? La haute montagne oriento-pyrénéenne se prête bien à ce type de recherche en raison de l'omniprésence des versants dans un contexte morphologique propice à l'éboulisation, de la présence de plantes adaptées aux mouvements des éboulis et des nombreuses études anciennement réalisées dans des disciplines variées. Ce travail de recherche est organisé en quatre axes complémentaires afin de répondre au mieux aux questions précédemment posées. Le premier concerne la géographie des éboulis. Les outils cartographiques et statistiques permettent de mieux comprendre la répartition spatiale de ces versants et de les caractériser. Le deuxième axe traite de l'ambiance climatique d'altitude. Les données climatiques locales et celles recueillies grâce à des enregistreurs de températures mettent en évidence le rôle joué par les paramètres thermiques sur l'efficacité de la gélifraction dans les parois et chicots rocheux, et sur le transit des débris. Le troisième vise à quantifier les dynamiques saisonnières sur les talus d'éboulis et à expliquer les déplacements de matériaux grâce aux paramètres lithologiques, topographiques, climatiques et biologiques. Le fonctionnement global du géosystème prenant en compte les systèmes amont (parois rocheuses et chicots rocheux) et aval (glaciers rocheux, pelouses...) est traité d'une manière plus sommaire que celui des talus d'éboulis. Enfin, le dernier axe concerne l'étude phytogéomorphologique. Les espèces végétales lithophiles sont utilisées à la fois comme outils de mesure des déplacements des matériaux en sub-surface et comme marqueurs de morphodynamiques particulières sur les éboulis.

Dados e análises de gradientes altitudinais em diversidade têm sido fundamentais para o desenvolvimento e avaliação de uma série de teorias gerais da biodiversidade. Esta proposta pretende contribuir para uma melhor compreensão dos padrões de diversidade das samambaias e licófitas ao longo de um gradiente de altitude no Sul do Brasil, abrangendo desde as formações florestais da planície, composta pela floresta de Terras Baixas, Submontana, como por formações florestais de altitude superiores, florestas Montana e Altomontana, e considerando a influência de fatores climáticos e de solo sobre as comunidades de plantas deste grupo. Assim, os objetivos deste estudo são: 1) verificar se a composição e riqueza de espécies varia ao longo do gradiente altitudinal; 2) se caso afirmativo, descrever em qual altitude podemos encontrar a maior diversidade; 3) analisar se existem diferenças ao longo do gradiente nos padrões de distribuição entre comunidade epifítica, terrícola e rupícola; 4) estabelecer quais são os fatores ambientais e de espaço que governam a distribuição das espécies de samambaias e licófitas, sua riqueza e sua cobertura na floresta brasileira ao sul do Atlântico; 5) investigar se espécies epifíticas e terrestres respondem igualmente aos condutores ambientais. Os resultados de um estudo detalhado, bem como de análises morfológicas, ecológicas, pedológicas e geográficas são: 1) a composição e riqueza de espécies variou significativamente ao longo do gradiente altitudinal; 2) riqueza e diversidade foram maiores nas formações superiores (Montana e Altomontana); 3) comunidades ecológicas distintas (epifítica, terrícola e rupícola) diferiram quanto ao padrão de distribuição observado ao longo do gradiente; 4) os fatores ambientais (tais como: insolação solar, precipitação, velocidade do vento, umidade relativa do ar relacionados ao clima e, concentração de matéria orgânica, alumínio, pH e profundidade, relacionados ao solo) apresentam maior importância na variação da distribuição de espécies ao longo do gradiente; 5) a variação da distribuição de espécies epífitas demostrou ser mais relacionada aos condutores ambientais relacionados aos fatores climáticos enquanto que as espécies terrestres, aos fatores edáficos. Processos de nicho (teoria de nicho) são os responsáveis por modular a distribuição e a abundância das espécies de samambaias e licófitas ao longo do gradiente altitudinal da Floresta Atlântica Sul Brasileira. Adicionalmente, é descrito novo registro de Stigmatopteris no Estado do Rio Grande do Sul, Brasil. / Data and analyses of elevational gradients in diversity have been central to the development and evaluation of a range of biodiversity general theories. This proposal seeks to contribute to a better understanding of the ferns and lycophytes diversity patterns along an elevation gradient in Southern Brazil, spanning from Lowland and Submontane to Montane and Upper Montane forest formations, and considering the influence of climatic factors and soil on communities of this group plants. The objectives of this study are: 1) to verify the composition and species richness varies along the altitudinal gradient; 2) if so, describe at what altitude we find the greatest diversity; 3) to analyze if there are differences along the gradient in the distribution patterns of epiphytic community, terrestrial and rupicolous; 4) establish what are the environmental and spatial factors that govern ferns and wealth lycophyte species and its coverage in the Brazilian forest at the South of the Atlantic; 5) investigate if epiphytic and terrestrial species also respond to environmental drivers. The results of a detailed study, as well as morphological, ecological, soil and geographical study are: 1) the composition and species richness varied significantly along the altitudinal gradient; 2) richness and diversity were higher in the Montane and Upper Montane formations; 3) distinct ecological communities (epiphytic, terrestrial and rupicolous) to differ on the distribution pattern observed along the gradient; 4) the environmental factors (such as solar insolation, precipitation, wind speed, relative humidity related to the climate, and concentration of organic matter, aluminum, pH and depth, related to soil) have greater importance in the variation species distribution along the gradient; 5) the variation of the distribution of epiphytes demonstrated to be more related to environmental drivers that are related to climatic factors, while terrestrial species are related to the soil factors. Niche processes (niche theory) are responsible for modulate distribution patterns and species abundance along elevational gradient in the Southern Brazilian Atlantic Forest. In addition, we present the new registration Stigmatopteris in the State of Rio Grande do Sul, Brazil.

Natural abundance of 15N varies greatly and unpredictably within and between environments. The unpredictable nature of 15N limits the use of N isotope natural abundance (d15N) in tracing the flow and fate of N in environments. Recent investigations have, however, revealed consistent and repeatable patterns of 15N in some ecosystem components. These patterns suggest that d15N may yet provide a tool to investigate and illuminate ecosystem N cycling processes. Identifying and quantifying the sources of isotopic variation must precede any significant advance in the application of this technique, and to this end an assessment of isotopic variation associated with major ecosystem components has been carried out in this thesis. d15N patterns have been established, hypotheses proposed and tested, and conclusions about the application of the technique are presented. 15N patterns in surface and groundwater were measured in a variety of different land-use catchments in an attempt to identify distinct isotopic 'fingerprints'. High levels of 15N variation were measured in both stream and groundwaters, resulting in strongly overlapping land-use 'fingerprints'. Environmental 15N variation in streams and groundwaters was found to be too great to differentiate between land-uses based on d15N alone. In contrast, the artificially 15N enriched signature of effluent N was used to trace its flow and fate, following irrigation, in a forested catchment. The effluent d15N signature allowed it to be traced into the major ecosystem components, permitting a first order N budget to be determined for effluent N storage and loss. N sources with significantly different 15N signatures to that of 'background ecosystem N' can therefore be used to trace the flow and fate of N in ecosystems. During the course of this work a number of higher and lower order plants were observed to have highly depleted (lt; -8 ) d15N signatures. Epiphytes and lithophytes, strongly reliant on atmospheric N sources, were consistently depleted in 15N, with signatures as low as -24 , measured in a range of environments. A similar level of depletion was measured in a wide range of plants growing in early primary succession sites (as low as -22.3 ), which could not be accounted for by any abiotic or biotic factor or significantly depleted N source. The absence of any measurable driver of depletion suggested a universal fractionating mechanism which acts in a wide range of environments and vegetation types. Diffusive uptake of atmospheric NH3(g) and the proportional uptake of a supplied N source were two proposed mechanisms that could theoretically account for the level and universal nature of depletion. Diffusive uptake of atmospheric NH3(g) was tested as a primary fractionating mechanism in plants. Strongly N deficient plants were capable of utilising NH3(g) as a nutritional source, but the level of 15N depletion measured in these plants closely approximated that of the inherent NH3(g) d15N signature. No significant additional fractionation is associated with NH3(g) diffusive uptake. Diffusive uptake of atmospheric NH3(g) by plants cannot alone account for the level of depletion measured in early primary succession plant communities. Proportional uptake of a N source as a primary fractionating mechanism was tested by growing plants in various concentrations and rates of applied N. Fractionation attributed to the proportional uptake of a supplied N source, as a consequence of P limitation or rapid flow over roots, resulted in a significant level of 15N depletion in plants. The level of depletion attributed to this mechanism was, however, not sufficient to account for the level measured in early primary succession plant communities. Individual 15N fractionating mechanisms cannot alone explain the level of depletion observed in early primary succession plants, however a combination of fractionating mechanisms can. Fractionation attributed to the proportional uptake of an already depleted N source, i.e., wet deposited N, largely accounts for the level of depletion measured in early succession plant communities. This two-step fractionation model can act on both higher and lower plants, independent of ecosystem biotic and abiotic factors. Additional, and less dramatic fractionations attributed to atmospheric NH3(g) uptake, mycorrhizal associations, internal remobilisation, and taxon-specific N acquisition strategies, will contribute to the level of d15N depletion. This thesis presents the first extensive survey of highly depleted d15N signatures in terrestrial vegetation. Furthermore, thorough testing of theoretically plausible mechanisms has resulted in a full account of the highly depleted d15N signatures measured in a wide range of vegetation types and environments.

Dados e análises de gradientes altitudinais em diversidade têm sido fundamentais para o desenvolvimento e avaliação de uma série de teorias gerais da biodiversidade. Esta proposta pretende contribuir para uma melhor compreensão dos padrões de diversidade das samambaias e licófitas ao longo de um gradiente de altitude no Sul do Brasil, abrangendo desde as formações florestais da planície, composta pela floresta de Terras Baixas, Submontana, como por formações florestais de altitude superiores, florestas Montana e Altomontana, e considerando a influência de fatores climáticos e de solo sobre as comunidades de plantas deste grupo. Assim, os objetivos deste estudo são: 1) verificar se a composição e riqueza de espécies varia ao longo do gradiente altitudinal; 2) se caso afirmativo, descrever em qual altitude podemos encontrar a maior diversidade; 3) analisar se existem diferenças ao longo do gradiente nos padrões de distribuição entre comunidade epifítica, terrícola e rupícola; 4) estabelecer quais são os fatores ambientais e de espaço que governam a distribuição das espécies de samambaias e licófitas, sua riqueza e sua cobertura na floresta brasileira ao sul do Atlântico; 5) investigar se espécies epifíticas e terrestres respondem igualmente aos condutores ambientais. Os resultados de um estudo detalhado, bem como de análises morfológicas, ecológicas, pedológicas e geográficas são: 1) a composição e riqueza de espécies variou significativamente ao longo do gradiente altitudinal; 2) riqueza e diversidade foram maiores nas formações superiores (Montana e Altomontana); 3) comunidades ecológicas distintas (epifítica, terrícola e rupícola) diferiram quanto ao padrão de distribuição observado ao longo do gradiente; 4) os fatores ambientais (tais como: insolação solar, precipitação, velocidade do vento, umidade relativa do ar relacionados ao clima e, concentração de matéria orgânica, alumínio, pH e profundidade, relacionados ao solo) apresentam maior importância na variação da distribuição de espécies ao longo do gradiente; 5) a variação da distribuição de espécies epífitas demostrou ser mais relacionada aos condutores ambientais relacionados aos fatores climáticos enquanto que as espécies terrestres, aos fatores edáficos. Processos de nicho (teoria de nicho) são os responsáveis por modular a distribuição e a abundância das espécies de samambaias e licófitas ao longo do gradiente altitudinal da Floresta Atlântica Sul Brasileira. Adicionalmente, é descrito novo registro de Stigmatopteris no Estado do Rio Grande do Sul, Brasil. / Data and analyses of elevational gradients in diversity have been central to the development and evaluation of a range of biodiversity general theories. This proposal seeks to contribute to a better understanding of the ferns and lycophytes diversity patterns along an elevation gradient in Southern Brazil, spanning from Lowland and Submontane to Montane and Upper Montane forest formations, and considering the influence of climatic factors and soil on communities of this group plants. The objectives of this study are: 1) to verify the composition and species richness varies along the altitudinal gradient; 2) if so, describe at what altitude we find the greatest diversity; 3) to analyze if there are differences along the gradient in the distribution patterns of epiphytic community, terrestrial and rupicolous; 4) establish what are the environmental and spatial factors that govern ferns and wealth lycophyte species and its coverage in the Brazilian forest at the South of the Atlantic; 5) investigate if epiphytic and terrestrial species also respond to environmental drivers. The results of a detailed study, as well as morphological, ecological, soil and geographical study are: 1) the composition and species richness varied significantly along the altitudinal gradient; 2) richness and diversity were higher in the Montane and Upper Montane formations; 3) distinct ecological communities (epiphytic, terrestrial and rupicolous) to differ on the distribution pattern observed along the gradient; 4) the environmental factors (such as solar insolation, precipitation, wind speed, relative humidity related to the climate, and concentration of organic matter, aluminum, pH and depth, related to soil) have greater importance in the variation species distribution along the gradient; 5) the variation of the distribution of epiphytes demonstrated to be more related to environmental drivers that are related to climatic factors, while terrestrial species are related to the soil factors. Niche processes (niche theory) are responsible for modulate distribution patterns and species abundance along elevational gradient in the Southern Brazilian Atlantic Forest. In addition, we present the new registration Stigmatopteris in the State of Rio Grande do Sul, Brazil.

Dados e análises de gradientes altitudinais em diversidade têm sido fundamentais para o desenvolvimento e avaliação de uma série de teorias gerais da biodiversidade. Esta proposta pretende contribuir para uma melhor compreensão dos padrões de diversidade das samambaias e licófitas ao longo de um gradiente de altitude no Sul do Brasil, abrangendo desde as formações florestais da planície, composta pela floresta de Terras Baixas, Submontana, como por formações florestais de altitude superiores, florestas Montana e Altomontana, e considerando a influência de fatores climáticos e de solo sobre as comunidades de plantas deste grupo. Assim, os objetivos deste estudo são: 1) verificar se a composição e riqueza de espécies varia ao longo do gradiente altitudinal; 2) se caso afirmativo, descrever em qual altitude podemos encontrar a maior diversidade; 3) analisar se existem diferenças ao longo do gradiente nos padrões de distribuição entre comunidade epifítica, terrícola e rupícola; 4) estabelecer quais são os fatores ambientais e de espaço que governam a distribuição das espécies de samambaias e licófitas, sua riqueza e sua cobertura na floresta brasileira ao sul do Atlântico; 5) investigar se espécies epifíticas e terrestres respondem igualmente aos condutores ambientais. Os resultados de um estudo detalhado, bem como de análises morfológicas, ecológicas, pedológicas e geográficas são: 1) a composição e riqueza de espécies variou significativamente ao longo do gradiente altitudinal; 2) riqueza e diversidade foram maiores nas formações superiores (Montana e Altomontana); 3) comunidades ecológicas distintas (epifítica, terrícola e rupícola) diferiram quanto ao padrão de distribuição observado ao longo do gradiente; 4) os fatores ambientais (tais como: insolação solar, precipitação, velocidade do vento, umidade relativa do ar relacionados ao clima e, concentração de matéria orgânica, alumínio, pH e profundidade, relacionados ao solo) apresentam maior importância na variação da distribuição de espécies ao longo do gradiente; 5) a variação da distribuição de espécies epífitas demostrou ser mais relacionada aos condutores ambientais relacionados aos fatores climáticos enquanto que as espécies terrestres, aos fatores edáficos. Processos de nicho (teoria de nicho) são os responsáveis por modular a distribuição e a abundância das espécies de samambaias e licófitas ao longo do gradiente altitudinal da Floresta Atlântica Sul Brasileira. Adicionalmente, é descrito novo registro de Stigmatopteris no Estado do Rio Grande do Sul, Brasil. / Data and analyses of elevational gradients in diversity have been central to the development and evaluation of a range of biodiversity general theories. This proposal seeks to contribute to a better understanding of the ferns and lycophytes diversity patterns along an elevation gradient in Southern Brazil, spanning from Lowland and Submontane to Montane and Upper Montane forest formations, and considering the influence of climatic factors and soil on communities of this group plants. The objectives of this study are: 1) to verify the composition and species richness varies along the altitudinal gradient; 2) if so, describe at what altitude we find the greatest diversity; 3) to analyze if there are differences along the gradient in the distribution patterns of epiphytic community, terrestrial and rupicolous; 4) establish what are the environmental and spatial factors that govern ferns and wealth lycophyte species and its coverage in the Brazilian forest at the South of the Atlantic; 5) investigate if epiphytic and terrestrial species also respond to environmental drivers. The results of a detailed study, as well as morphological, ecological, soil and geographical study are: 1) the composition and species richness varied significantly along the altitudinal gradient; 2) richness and diversity were higher in the Montane and Upper Montane formations; 3) distinct ecological communities (epiphytic, terrestrial and rupicolous) to differ on the distribution pattern observed along the gradient; 4) the environmental factors (such as solar insolation, precipitation, wind speed, relative humidity related to the climate, and concentration of organic matter, aluminum, pH and depth, related to soil) have greater importance in the variation species distribution along the gradient; 5) the variation of the distribution of epiphytes demonstrated to be more related to environmental drivers that are related to climatic factors, while terrestrial species are related to the soil factors. Niche processes (niche theory) are responsible for modulate distribution patterns and species abundance along elevational gradient in the Southern Brazilian Atlantic Forest. In addition, we present the new registration Stigmatopteris in the State of Rio Grande do Sul, Brazil.