A limitação de dispersão de sementes tem sido empiricamente investigada como um mecanismo equalizador das diferenças competitivas entre espécies que coexistem em comunidades vegetais. Além da limitação espacial, as variações temporais da dispersão também podem ser importantes para a coexistência das espécies. O objetivo do estudo foi verificar a relação entre limitação de dispersão espacial (LDe) e limitação de dispersão temporal (LDt) e as características das espécies que influenciam as duas limitações. Em uma floresta da planície costeira, a chuva de sementes foi amostrada, ao longo de 36 meses, em 40 coletores (20m2) distantes 100m entre si e da borda do fragmento. A abordagem de seleção de modelos mistos foi utilizada para testar a relação entre a limitação de dispersão e a massa média das sementes, a síndrome de dispersão, a altura máxima local e a frequência de distribuição dos indivíduos adultos. Os resultados encontrados mostram que as proporções de espécies analisadas limitadas espacial (90,3%) e temporalmente (70,9%) foram altas e a correlação entre LDe e LDT também foi alta (Spearman = 0,8). Tanto para LDe como LDt, foram selecionados a massa média das sementes, a altura máxima e a frequência de distribuição de adultos. Em geral, as relações encontradas indicam que espécies com sementes maiores, com menor altura máxima e menor frequência de distribuição dos indivíduos adultos são mais limitadas espacial e temporalmente. Apesar desses fatores terem sido selecionados, houve uma grande variação nos efeitos para as espécies analisadas. O estudo reforça o fato do mecanismo de limitação de dispersão espacial ser frequentemente encontrado em comunidades arbóreas e apresenta uma abordagem temporal para o estudo da limitação de dispersão. A redução das interações competitivas interespecíficas, decorrente das altas limitações de dispersão observadas no estudo, se contrapõe às teorias amplamente aceitas (Janzen-Connell) que indicam os mecanismos de dispersão eficientes de sementes como uma forma de minimizar a competição intraespecífica. A importância relativa dos mecanismos de dispersão e de limitação de dispersão para a manutenção da diversidade em comunidades deveria ser estudada para avaliar em quais situações ou para quais conjuntos de espécies a coexistência é mediada pela ausência das interações intraespecíficas ou interespecíficas / The seed limitation has been empirically investigated as an equalizing mechanism of the competitive differences between species that coexists in plant communities. In addition to the spatial limitation, the temporal variations of seed dispersal can also be important to the species coexistence. The aim of the study was to investigate the relation between spatial seed limitation and temporal seed limitation and the species characteristics that influence both limitations. In a coastal plain forest, the seed rain was sampled over 36 months in 40 traps (20 m2), 100 m distant from each other and from the forest edges. The generalized mixed-effect models approach was used to test the relation between seed limitation and the average mass of the seeds, the dispersal syndrome, the trees local maximum height, and the adult trees distribution frequencies. The results show that the proportion of the analysed species spatial limited (90,3%) e temporal limited (70,9%) were high and the correlation between spatial seed limitation and temporal seed limitation were also high (Spearman = 0,8). To explain spatial seed limitation and temporal seed limitation were selected the average mass of seeds, the trees local maximum height and the adult trees distribution frequencies. Usually those relations indicate that species with lagger seeds, lowest maximum tree height and lowest adult trees distribution frequencies are more limited spatially and temporally. Although these factors have been selected, there was a great variation in the effects of the species analysed. The study supports the fact that the spatial seed limitation mechanism is frequently found in tree communities and shows a temporal approach to the seed limitation study. The reduction of interspecific competitive interactions, due to the high seed limitation observed in the study, is opposed to widely accepted theories (Janzen-Connell) that indicates that the efficient seed dispersal mechanism are a strategy to minimize the intraspecific competion. The relative importance of dispersal mechanisms and seed limitation on maintaning diversity in trees communities should be studied to evaluate which situations or which species sets the coexistence is mediated by the absence of intraspecific or interspecifir interactions

<p>The distribution of plant species and functional traits in alpine and arctic environments are determined by abiotic conditions, but also by biotic interactions. In this thesis, I investigate interactions among plants and herbivory effects on plant community composition and plant functional traits in three different regions: Swedish Lapland, Beringia (USA/Russia) and Finnmark (Norway). Reindeer grazing was found to be extensive in southern Lapland and had limited effects on plant community composition and seedling germination. However, reindeer presence was found to influence plant functional traits, particularly in the subalpine birch forest. Tall herbs were lower and had lower SLA when reindeer were present, while small herbs showed an opposite pattern. The contrasting effects on the two herb groups are probably explained by a competitive release for small herbs when the tall herbs are suppressed by reindeer. Rodents had the largest relative impact on plant community composition in southern Lapland and this is consistent with the study from Finnmark, where rodents heavily affected dwarf shrubs on predator-free islands. With no predators present, vole densities increased profoundly and almost depleted some dwarf shrub species. These results support the idea that small mammals in arctic and alpine tundra are controlled by predators (i.e. top-down). However, a decrease in the nutritional quality in a sedge after defoliation gives support for the idea that small mammals are regulated by plant quality (i.e. bottom-up). In Beringia, small and large herbivores differed in the relation to plant community composition, since large herbivores were related to species richness and small herbivores were related to plant abundance. Plant functional traits were related only to large herbivores and standing crop of vascular plants.</p>

Studies of seed dynamics and seedling establishment at the community level in savanna ecosystems are scant, particularly in the tropical savannas of Central Brazil. The Brazilian savannas (cerrado) have the largest diversity of plant species among Neotropical savannas. Cerrado vegetation exhibits consistent changes in tree density and tree size along shallow topographic gradients. Vegetation types differ from closed savannas with high tree density, and a nearly continuous canopy cover in the uppermost portions, to open savannas with scattered short trees in the low portions of the topographic gradients. Whether or not dynamics of seeds and seedlings of woody species are consistent with variations in tree density and tree canopy cover across the cerrado landscape has not been determined, but could potentially influence such spatial patterns of tree density and canopy cover. The main objective of this study was to evaluate seed dynamics (i.e., seed rain, soil seed banks, and seed characteristics) and seedling establishment of woody species in three major cerrado vegetation types (closed, intermediate and open savannas) which differ in tree canopy cover and tree height. These vegetation types are located along topographic gradients of approximately 30 m in elevation and 1000 m in length. I conducted field work from June 2005 to August 2008 at the Instituto Brasileiro de Geografia e Estatística (IBGE) reserve, a field experimental station located 35 Km south of Brasilia, Brazil (15˚ 56' S, 47˚ 63' W, altitude 1100 m). I established permanent plots in each of the three vegetation types to assess seed rain, seed limitation, seed predation, and seed removal. I also quantified density and composition of the soil seed bank in each vegetation type and studied seed characteristics (i.e., kind of dormancy, moisture content, longevity) of 14 common cerrado woody species. In the field, I quantified establishment of woody seedlings in fire-protected savannas as well as in accidentally burned and frequently burned savannas. In the greenhouse I performed experiments to evaluate the effect of litter cover and light level on seedling emergence of 9 common cerrado woody species. Finally, I evaluated the effect of pulses of heat on seed germination of 5 common cerrado woody species under controlled conditions. The overall results of this study show that regardless of tree canopy cover, cerrado vegetation types are limited in seeds and seedlings of woody species, but these limitations are higher in open than in closed savannas. Most woody species do not form persistent soil seed banks along cerrado vegetation types because their seeds are short-lived, predated and quickly removed from the ground. In addition, dispersal season and dormancy appear to control timing of germination at the onset of the rainy season to ensure survival of seedlings in the field. In the absence of fire, establishment of woody seedlings is higher in closed than in open savannas because canopy cover mitigates the stressful environmental conditions of open savannas and thus facilitates establishment of woody seedlings. As a result, closed savannas are likely to maintain higher density of woody elements over time than open savannas. In burned savannas, however, species composition of woody and herbaceous seedlings is significantly affected because establishment of seedlings of woody and herbaceous species decreases but vegetative reproduction increases. This dissertation study indicates that interactions of early life-history stages of woody species with environmental and disturbance factors appear to play a more significant role in maintaining woody cover variations along cerrado topographic gradients than previously recognized.

The distribution of plant species and functional traits in alpine and arctic environments are determined by abiotic conditions, but also by biotic interactions. In this thesis, I investigate interactions among plants and herbivory effects on plant community composition and plant functional traits in three different regions: Swedish Lapland, Beringia (USA/Russia) and Finnmark (Norway). Reindeer grazing was found to be extensive in southern Lapland and had limited effects on plant community composition and seedling germination. However, reindeer presence was found to influence plant functional traits, particularly in the subalpine birch forest. Tall herbs were lower and had lower SLA when reindeer were present, while small herbs showed an opposite pattern. The contrasting effects on the two herb groups are probably explained by a competitive release for small herbs when the tall herbs are suppressed by reindeer. Rodents had the largest relative impact on plant community composition in southern Lapland and this is consistent with the study from Finnmark, where rodents heavily affected dwarf shrubs on predator-free islands. With no predators present, vole densities increased profoundly and almost depleted some dwarf shrub species. These results support the idea that small mammals in arctic and alpine tundra are controlled by predators (i.e. top-down). However, a decrease in the nutritional quality in a sedge after defoliation gives support for the idea that small mammals are regulated by plant quality (i.e. bottom-up). In Beringia, small and large herbivores differed in the relation to plant community composition, since large herbivores were related to species richness and small herbivores were related to plant abundance. Plant functional traits were related only to large herbivores and standing crop of vascular plants.

A limitação de dispersão de sementes tem sido empiricamente investigada como um mecanismo equalizador das diferenças competitivas entre espécies que coexistem em comunidades vegetais. Além da limitação espacial, as variações temporais da dispersão também podem ser importantes para a coexistência das espécies. O objetivo do estudo foi verificar a relação entre limitação de dispersão espacial (LDe) e limitação de dispersão temporal (LDt) e as características das espécies que influenciam as duas limitações. Em uma floresta da planície costeira, a chuva de sementes foi amostrada, ao longo de 36 meses, em 40 coletores (20m2) distantes 100m entre si e da borda do fragmento. A abordagem de seleção de modelos mistos foi utilizada para testar a relação entre a limitação de dispersão e a massa média das sementes, a síndrome de dispersão, a altura máxima local e a frequência de distribuição dos indivíduos adultos. Os resultados encontrados mostram que as proporções de espécies analisadas limitadas espacial (90,3%) e temporalmente (70,9%) foram altas e a correlação entre LDe e LDT também foi alta (Spearman = 0,8). Tanto para LDe como LDt, foram selecionados a massa média das sementes, a altura máxima e a frequência de distribuição de adultos. Em geral, as relações encontradas indicam que espécies com sementes maiores, com menor altura máxima e menor frequência de distribuição dos indivíduos adultos são mais limitadas espacial e temporalmente. Apesar desses fatores terem sido selecionados, houve uma grande variação nos efeitos para as espécies analisadas. O estudo reforça o fato do mecanismo de limitação de dispersão espacial ser frequentemente encontrado em comunidades arbóreas e apresenta uma abordagem temporal para o estudo da limitação de dispersão. A redução das interações competitivas interespecíficas, decorrente das altas limitações de dispersão observadas no estudo, se contrapõe às teorias amplamente aceitas (Janzen-Connell) que indicam os mecanismos de dispersão eficientes de sementes como uma forma de minimizar a competição intraespecífica. A importância relativa dos mecanismos de dispersão e de limitação de dispersão para a manutenção da diversidade em comunidades deveria ser estudada para avaliar em quais situações ou para quais conjuntos de espécies a coexistência é mediada pela ausência das interações intraespecíficas ou interespecíficas / The seed limitation has been empirically investigated as an equalizing mechanism of the competitive differences between species that coexists in plant communities. In addition to the spatial limitation, the temporal variations of seed dispersal can also be important to the species coexistence. The aim of the study was to investigate the relation between spatial seed limitation and temporal seed limitation and the species characteristics that influence both limitations. In a coastal plain forest, the seed rain was sampled over 36 months in 40 traps (20 m2), 100 m distant from each other and from the forest edges. The generalized mixed-effect models approach was used to test the relation between seed limitation and the average mass of the seeds, the dispersal syndrome, the trees local maximum height, and the adult trees distribution frequencies. The results show that the proportion of the analysed species spatial limited (90,3%) e temporal limited (70,9%) were high and the correlation between spatial seed limitation and temporal seed limitation were also high (Spearman = 0,8). To explain spatial seed limitation and temporal seed limitation were selected the average mass of seeds, the trees local maximum height and the adult trees distribution frequencies. Usually those relations indicate that species with lagger seeds, lowest maximum tree height and lowest adult trees distribution frequencies are more limited spatially and temporally. Although these factors have been selected, there was a great variation in the effects of the species analysed. The study supports the fact that the spatial seed limitation mechanism is frequently found in tree communities and shows a temporal approach to the seed limitation study. The reduction of interspecific competitive interactions, due to the high seed limitation observed in the study, is opposed to widely accepted theories (Janzen-Connell) that indicates that the efficient seed dispersal mechanism are a strategy to minimize the intraspecific competion. The relative importance of dispersal mechanisms and seed limitation on maintaning diversity in trees communities should be studied to evaluate which situations or which species sets the coexistence is mediated by the absence of intraspecific or interspecifir interactions

There are generally two contrasting alternatives to what limits recruitment in plants, namely the availability of seeds (seed limitation) or the quality or quantity of suitable sites (microsite limitation). Dust seeds, the smallest existing seeds, lack or have minimal nutrient reserves. During germination and initial development they consequently parasitize on mycorrhizal fungi. This is called mycoheterotrophy, and can vary in degree of fungal dependency in adult plants from full, partial or initial mycoheterotrophy. The aim of this thesis was to investigate the recruitment ecology of mycoheterotrophic Ericaceae (tribe Pyroleae) species with dust seeds, and to determine what limits their recruitment. The investigated species were: Chimaphila umbellata, Moneses uniflora, Orthilia secunda, Pyrola chlorantha, P. minor and P. rotundifolia. This aim was achieved by combining field experiments (seed sowing) with isotope analysis and fungal host pyrosequencing. Results provide evidence that the species in Pyroleae are heterogeneous, not only with regard to their degree of mycoheterotrophy, but also concerning germination and early seedling development. A combination of microsite and seed limitation is thus likely to be of importance for all studied species, but the relative importance of these limitations varies among species. Despite having adaptations for wind dispersal the majority of the seeds were deposited in close vicinity of the seed source. But with high seed production at least some seeds should be able to disperse long-distance. Seedlings of all studied species were found to associate with a wide range of ectomycorrhizal fungi, at least during their initial developmental stages. There seems to be a tendency for host narrowing in some Pyroleae species, but not as strict as the host specialization seen in fully mycoheterotrophic Monotropa hypopitys, supporting the hypothesis of geographical and developmental host shifts. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 3: In press. Paper 4: Manuscript.</p>