The forces of interaction were measured between the siderophore azotobactin and the minerals goethite (FeOOH) and diaspore (AlOOH) in solution using force microscopy. Azotobactin was covalently linked to a hydrazide terminated atomic force microscope tip using a standard protein coupling technique. Upon contact with each mineral surface, the adhesion force between azotobactin and goethite was two to three times the value observed for the isostructural Al-equivalent diaspore. The affinity for the solid iron oxide surface reflected in the force measurements correlates with the specificity of azotobactin for aqueous ferric iron. Further, the adhesion force between azotobactin and goethite significantly decreases when small amounts of soluble iron are added to the system suggesting a significant specific interaction between the azotobactin and the mineral surface. Changes in the force signature with pH and ionic strength were fairly predictable when considering mineral solubility, the charge character of the mineral surfaces, the molecular structure of azotobactin, and the intervening solution. Molecular and quantum mechanical calculations which were completed to further investigate the interaction between azotobactin and iron/aluminum oxide surfaces, and to more fully understand the force measurements, also showed an increased force affinity for Fe over Al. Ab initio calculations on siderophore fragment analogs suggest the iron affinity can be attributed to increased electron density associated with the Fe-O bond compared to the Al-O bond; an observation that correlates with iron's larger electronegativity compared to aluminum. Attachment of the ligand to each surface was directed by steric forces within the molecule and coulombic interactions between the siderophore oxygens and the metals in the mineral. Chelating ligand pairs coordinated with neighboring metal atoms in a bidentate, binuclear geometry. Upon simulated retraction of azotobactin from each surface, the Fe-O(siderophore) bonds persisted into a higher force regime than Al-O(siderophore) bonds, and surface metals were removed from both minerals. Extrapolation of the model to more realistic hydrated conditions using a PCM model in the quantum mechanical calculations and water clusters in the molecular mechanical model demonstrated that the presence of water energetically favors and enhances metal extraction, making this a real possibility in a natural system. / Ph. D.

Die Ausbreitung von Diasporen ist ein zentraler Abschnitt des pflanzlichen Lebenszyklus. Insbesondere Fernausbreitung (>100 m) hat erhebliche Auswirkungen auf eine Vielzahl von biologischen Prozessen, ist aber mit empirischen Messmethoden kaum erfassbar. Ziel der Arbeit ist es daher, die Fernausbreitung von Diasporen aus der Sicht der Mikrometeorologie zu modellieren. Numerische Berechnungsverfahren sind dazu experimentell zu prüfen im Hinblick auf die Wiedergabe der turbulenten Verhältnisse in einem konkreten Messgebiet und an parallel zu diesen Messungen beobachteten Ausbreitungsbedingungen für spezielle Diasporen. Hier wird das Konzept für ein solches Ausbreitungsberechnungsverfahren erläutert und erste Ergebnisse vorgestellt. / Diaspore dispersal is a very important chapter in the life cycle of many plants. In particular long range transport (>100m) has significant effects on many biological processes, but is not ascertainable by empirical measurements. So the objective of this work is to model the long range dispersion of diaspores using micrometeorological methods. Numerical calculations must reproduce the experimentally observed turbulent conditions and the propagation of special seeds within a concrete landscape. The concept of such a numerical model and first results are described here.

Des affleurements alumineux à kaolinite, andalousite, pyrophyllite et diaspore se répartissent en six groupes principaux situés dans l'unité 1 d'Avène (Nord-est de la Montagne Noire). Ces roches massives qui contiennent essentiellement de la silice, de l'alumine et de l'eau sont bordées par une enveloppe sériciteuse et silicifiée à tourmaline et pyrite. Le passage de la métarhyolite encaissante, d'âge Précambrien ou Cambrien inférieur, aux faciès à silicates et hydrates d'alumine est graduel. Il se fait par une disparition progressive des feldspaths (métarhyolites sans feldspath) puis de la foliation métamorphique avec apparition d'andalousite. Les zonalités observées se rencontrent dans d'autres sites mondiaux (Caroline du Nord, Maroc) à altération hautement alumineuse où la roche encaissante est une rhyolite et les altérites présentent les mêmes types de minéraux que les affleurements alumineux de la Montagne Noire. L'étude pétrogénétlque montre que l'essentiel des transformations est antérieur au métamorphisme hercynien (phase hydrothermale majeure) et que les zonalités ainsi que la diversité des faciés observés peuvent s'expliquer par un modèle génétique unique présentant des affinités avec le modèle génétique des porphyres cuprifères. La phase hydrothermale majeure se caractérise par la superposition de deux épisodes d'altération qui se produisent entre le Cambrien inférieur et le Cambrien moyen dans un contexte géodynamique distensif: - l'altération séricitique précoce résulte d'un lessivage de la roche par des fluides acides (pH-4),relativement chauds (T-400'C), sursaturés en silice et contenant principalement du bore, du soufre, du fluor et du sodium. Les feldspaths de la roche, sous faible couverture et peu compactée, (P-30 MPa), sont transformés en muscovite avec un lessivage intense du sodium. La silice, le magnésium et le fer sont lessivés vers les bordures des zones d'apport où cristallisent la dravite et la pyrite. - l'altération argilitique avancée qui se surimpose à la première est liée à une baisse de la température et une augmentation de l'acidité des fluides qui sont enrichis en phosphore. Dans la zone centrale la muscovite est transformée en andalousite avec par .endroit cristallisation de scorzalite (phosphate accidentel) . Les alcalins et la silice sont lessivés vers la périphérie. Un modète génétique général qui explique les principales caractéristiques des sites a été élaboré. Il a la forme d'un ellipsoide de révolution à zone interne d'altération argililique avancée plus ou moins riche en andalousite et à enveloppe externe d'altération séricitique. Les zones silicifiées se répartissent dans l'enveloppe séricitique et à la base (voies d'apport des fluides) de l'ellipsoide. La topographie des sites riches en filons de quartz recoupe le modèle à sa base alors que la topographie des sites à filons de quartz rares et riches en andalousite recoupe le modèle dans sa partie haute. Les autres particularités s'expliquent par une évolution des paragenèses originelles dépendant du contexte géologique local ou régional: -au cours du métamorphisme hercynien les altérites sont reprises dans la schistosité de flux ce qui explique leur aspect stratiforme actuel et les déformations observées sur certains sites. -les sites de bordure du bassin houiller sont soumis à l'altération antéstéphanienne qui se manifeste par des phénomènes de bauxitisation localisés avec oxydation superficielle et altération de l'andalousite en kaolinile et boehmite. La boehmite est ensuite déstabilisée en diaspore sous faible couverture houillère. - les sites de Serviès et du Lissobre situés plus au Nord n'ont pas été soumis à l'altération antéstéphanienne ce qui explique l'absence de boehmite et de diaspore. A la fin du Stépthanien un contexte distensif associé à un réchauffement régional (métamorphisme tardistéphanien) provoque des circulations de fluides ( T-200'C) qui conduisent à la cristallisation de pyrophyllite à partir de la kaolinite dans les fractures des altérites. Au Permo-Trias des circulations de fluides liées à une période de distension provoquent l'altération de la scorzalite et la cristallisation de barytine en conditions supergènes.

Questions: 1. Are there differences among species in their preference for coniferous vs. deciduous forest? 2. Are tree and shrub species better colonizers of recent forest stands than herbaceous species? 3. Do colonization patterns of plant species groups depend on tree species composition? Location: Three deciduous and one coniferous recent forest areas in Brandenburg, NE Germany. Methods: In 34 and 21 transects in coniferous and deciduous stands, respectively, we studied the occurrence and percentage cover of vascular plants in a total of 150 plots in ancient stands, 315 in recent stands and 55 at the ecotone. Habitat preference, diaspore weight, generative dispersal potential and clonal extension were used to explain mechanisms of local migration. Regression analysis was conducted to test whether migration distance was related to species’ life-history traits. Results: 25 species were significantly associated with ancient stands and ten species were significantly more frequent in recent stands. Tree and shrub species were good colonizers of recent coniferous and deciduous stands. In the coniferous stands, all herbaceous species showed a strong dispersal limitation during colonization, whereas in the deciduous stands generalist species may have survived in the grasslands which were present prior to afforestation. Conclusions: The fast colonization of recent stands by trees and shrubs can be explained by their effective dispersal via wind and animals. This, and the comparably efficient migration of herbaceous forest specialists into recent coniferous stands, implies that the conversion of coniferous into deciduous stands adjacent to ancient deciduous forests is promising even without planting of trees.

Current knowledge about processes that generate long-distance dispersal of plants is still limited despite its importance for persistence of populations and colonization of new potential habitats. Today wild large mammals are presumed to be important vectors for long-distance transport of diaspores within and between European temperate forest patches, and in particular wild boars recently came into focus. Here we use a specific habit of wild boar, i.e. wallowing in mud and subsequent rubbing against trees, to evaluate epizoic dispersal of vascular plant diaspores. We present soil seed bank data from 27 rubbing trees versus 27 control trees from seven forest areas in Germany. The mean number of viable seeds and the plant species number were higher in soil samples near rubbing trees compared with control trees. Ten of the 20 most frequent species were more frequent, and many species exclusively appeared in the soil samples near rubbing trees. The large number of plant species and seeds – approximated > 1000 per tree – in the soils near rubbing trees is difficult to explain unless the majority were dispersed by wild boar. Hooked and bristly diaspores, i.e. those adapted to epizoochory, were more frequent, above that many species with unspecialised diaspores occurred exclusively near rubbing trees. Different to plant species closely tied to forest species which occur both in forest and open vegetation, and non-forest species were more frequent near rubbing trees compared with controls. These findings are consistent with previous studies on diaspore loads in the coats and hooves of shot wild boars. However, our method allows to identify the transport of diaspores from the open landscape into forest stands where they might especially emerge after disturbance, and a clustered distribution of epizoochorically dispersed seeds. Moreover, accumulation of seeds of wetness indicators near rubbing trees demonstrates directed dispersal of plant species inhabiting wet places between remote wallows. / Das aktuelle Wissen über Prozesse, die zur Fernausbreitung von Pflanzen führen, ist trotz ihrer Bedeutung für das Überleben von Populationen und die Besiedlung neuer potenzieller Habitate noch immer sehr begrenzt. Wildlebende Großsäuger sind heutzutage vermutlich wichtige Vektoren für den Ferntransport von Diasporen innerhalb und zwischen den einzelnen Waldflächen in Mitteleuropa, und speziell das Wildschwein (Sus scrofa L.) spielt dabei offenbar eine herausragende Rolle. Wir nutzen hier ein spezifisches Verhalten des Wildschweins – Suhlen im Schlamm und nachfolgendes Scheuern an sogenannten Malbäumen – um die epizoochore Ausbreitung von Gefäßpflanzen-Diasporen einzuschätzen. Dargestellt werden die Ergebnisse von Samenbank-Untersuchungen von 27 Malbäumen im Vergleich zu 27 Kontrollbäumen aus sieben Waldgebieten in Deutschland. Sowohl die mittlere Zahl lebensfähiger Samen als auch die Artenzahl waren höher in Bodenproben neben Malbäumen. Zehn der 20 in der Samenbank verbreitetsten Pflanzenarten hatten hier ihren Schwerpunkt, und viele Arten kamen ausschließlich in den neben Malbäumen gewonnenen Proben vor. Die große Zahl von Pflanzenarten und Samen – zumindest > 1000 pro Baum – im Boden an Malbäumen lässt sich nur durch die Aktivität der Wildschweine erklären. Mit Haken oder Borsten ausgestattete, d.h. an Epizoochorie angepasste Diasporen waren häufiger, aber auch viele Arten mit unspezialisierten Diasporen kamen ausschließlich in der Samenbank bei Malbäumen vor. Anders als weitgehend an Wald gebundene Pflanzenarten waren solche, die sowohl im Wald und im Offenland vorkommen, sowie nicht im Wald vorkommende Arten häufiger neben Malbäumen als neben Kontrollbäumen. Diese Befunde stimmen mit denen früherer Untersuchungen von Diasporenladungen im Fell und in den Hufen geschossener Wildschweine überein. Unsere Methode erlaubt darüber hinaus aber die Identifizierung des Diasporentransports aus dem Offenland in die Waldbestände, wo sie insbesondere nach Störungen keimen dürften, sowie einer ungleichmäßigen Verteilung epizoochor ausgebreiteter Diasporen. Außerdem zeigt die Akkumulation von Samen von Nässezeigern neben den Malbäumen eine gezielte Ausbreitung nasse Standorte bewohnender Pflanzenarten zwischen entfernt gelegenen Suhlen.

In order to investigate the dispersion of diaspores, the joined BioWind-project has been launched from the universities of Regensburg and Leipzig. The aim of this collaboration was to investigate dispersal of seeds by wind with respect to different atmospheric stability conditions. In the year 1972, Kaimal et al. proposed a standardized spectra of turbulence for each meteorological parameter and the resulting flows. Due to the fact, that the data of Kaimal et al. (1972) were collected from a measuring field whose surface was almost perfectly flat without any disturbances, the question was raised whether these spectra are also valid for the atmosphere over inhomogeneous terrain of the BioWind-project. To be consistent, a turbulence spectrum for each parameter was produced and a qualitative agreement with the standardized spectra was achieved. A systematic lower variance in the high-frequency region was determined for all meteorological parameters. / Um die Ausbreitung von Pflanzensamen (Diasporen) zu untersuchen, wurde das Projekt BioWind von den Universitäten Regensburg und Leipzig ins Leben gerufen. Ziel dieser Zusammenarbeit war es die Verbreitung von Pflanzensamen (Diasporen) durch Wind, unter Berücksichtigung verschiedener atmosphärischer Stabilitätsbedingungen, zu ermitteln. Im Jahre 1972 haben Kaimal et al. erstmals standardisierte Turbulenzspektren für verschiedene meteorologische Parameter und die daraus resultierenden Flussgrößen vorgestellt. Aufgrund der Tatsache, dass die von Kaimal et al. (1972) genutzten Daten auf Messplätzen ermittelt wurden, welche nahezu eben und störungsfrei waren, stellte sich die Frage, ob sich die ermittelten Spektren auch auf die inhomogenen Messfelder des BioWind-Projektes anwenden lassen. Eine qualitativ gute Übereinstimmung für die ermittelten Turbulenzspektren wurde erreicht. Die untersuchten Gröÿen zeigen allerdings eine systematisch niedrigere Varianz im hochfrequenten Turbulenzbereich.

Zugl.: Freiburg (Breisgau), Univ., Diss.

Orientador: Paulo Sérgio Moreira Carvalho de Oliveira / Tese (doutorado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-17T09:19:12Z (GMT). No. of bitstreams: 1 Bottcher_Claudia_D.pdf: 2277526 bytes, checksum: fd9c885535eb1d45ead71daf14f43c8a (MD5) Previous issue date: 2010 / Resumo: A grande quantidade de frutos carnosos e a diversidade e abundância de formigas em ambientes tropicais faz com que a interação entre formigas e diásporos seja bastante comum. Atualmente sabe-se que estas interações envolvem dezenas de espécies de plantas e formigas em uma comunidade ecológica. Dentre as espécies da fauna de formigas que interagem com diásporos, Pachycondyla striata e Odontomachus chelifer destacam-se como dispersores de sementes caídas no solo e as características físicas e químicas de seus ninhos freqüentemente promovem o estabelecimento de plântulas. Contudo, apesar da reconhecida importância dessas formigas para algumas espécies de plantas, nada se sabe sobre a influência desses diásporos no desenvolvimento e reprodução de suas colônias. Com base nesse cenário, esta tese foi dividida em duas partes relacionadas. Na primeira parte, Capítulo I, investigamos as interações entre formigas e diásporos não-mirmecocóricos em três diferentes fisionomias florestais da Ilha do Cardoso (restinga, planície e encosta), sudeste do Brasil. Durante dois anos de amostras mensais 517 interações envolvendo 53 espécies de formigas e 48 espécies de diásporos foram observadas. A maior parte das interações ocorreu em floresta de planície (54.62%), e menos da metade ocorreu nas demais áreas, encosta (31.33%) e restinga (14.06%). Myrmicinae teve o maior número de espécies em interação (39 espécies), totalizando 436 interações (82.20%). Seis espécies da subfamília Ponerinae foram responsáveis por 33 interações (6,4%). Myrtaceae foi a família de planta mais numerosa nas interações (14 espécies). Na segunda parte da tese, capítulos II e III, foi dado ênfase às duas espécies de formigas da subfamília Ponerinae (Pachycondyla striata e Odontomachus chelifer) onde investigamos a importância dos frutos na ecologia de forrageamento e desenvolvimento das colônias. No capítulo II nós estudamos como a estrutura dos ninhos, demografia das colônias, ritmo diário de atividades, dieta e área de vida dessas espécies variam espacial e sazonalmente em restinga e planície. Operárias de O. chelifer e de P. striata caçam de modo oportunista incluindo uma ampla variedade de itens alimentares, forrageando mais intensamente no período úmido quando alados são mais abundantes e as condições ambientais são mais favoráveis. A atividade de forrageamento apresentou variação entre as espécies de formigas, o que poderia ser associada com fatores físicos e disponibilidade de alimentos, mostrando que o comportamento de forrageamento de O. chelifer e P. striata corresponde a variabilidade temporal e espacial da Mata Atlântica. Para testar a hipótese de que o consumo de diásporos não-mirmecocóricos beneficiaria colônias de formigas, no Capítulo III nós testamos o efeito do consumo de sementes de C. canjerana, com arilos ricos em lipídeos, em colônias de O. chelifer sob condições controladas em laboratório. Constatamos que larvas de formigas alimentadas com diásporos de C. canjerana foram em média maiores e se desenvolveram melhor que larvas controle. Em conclusão, os resultados desta tese reforçam a importância da dispersão secundária por formigas para sementes primariamente dispersas por vertebrados. Os dados evidenciam também a importância do estudo da ecologia de forrageamento e da influência do consumo de frutos na sobrevivência e crescimento de colônias de formigas. / Abstract: The large number of fleshy fruits and the diversity and abundance of ants in tropical environments makes the ant-diaspore interaction very common. Currently it is known that these interactions involve a large number of plants and ants species in an ecological community. Among the ant species interacting with diaspores, Pachycondyla striata and Odontomachus chelifer stand out as main dispersers of seeds encountered on the forest ground. Moreover, the physical and chemical characteristics of their nests often promote the establishment of seedlings. However, despite the recognized importance of these ants for some species of plants, nothing is known about the influence of the consumption of fleshy diaspores in the development and reproduction of the ant colonies. Based on this scenario, this thesis was divided into two related parts. In the first part, Chapter I, we investigated the interactions between ants and nonmyrmecochorous diaspores in three different physiognomic forest of Cardoso Island (sandy, lowland and hillside), southeastern Brazil. During two years of monthly samples 517 interactions involving 53 ant species and 48 species of seeds were observed. Most interactions occurred in lowland Forest (54.62%), and less than half occurred in the two other areas, hillside (31.33%) and sandy Forest (14.06%). Myrmicinae had the highest species number in interaction (39 species), totaling 436 interactions (82.20%). The six species of the Ponerinae subfamily were observed exploring diaspores accounted for 33 interactions (6.4%). Myrtaceae was the most numerous plant family in interactions (14 species). In the second part of this thesis, Chapter II and III, we emphasized two species of Ponerinae (Pachycondyla striata and Odontomachus chelifer), investigating the importance of fruits in their foraging ecology and the development of their colonies. In Chapter II we studied how nest structure, colonies demography, daily activities, diet and home range of these two species vary spatially and seasonally in sandy and lowland forest. Odontomachus chelifer and Pachycondyla striata workers hunt opportunistically including a wide variety of food items. They forage more intensively during the wet season when alates are more abundant and environmental conditions are more favorable. The foraging activity showed variation among the ant species, which could be associated with physical factors and food availability, showing that the foraging behavior of O. chelifer and P. striata correspond to temporal and spatial variability of the Atlantic Forest. To test the hypothesis that the consumption of non-myrmecochorous diaspores benefit ant colonies, in Chapter III we tested the effect of consumption of seeds of Cabralea canjerana, with lipid-rich arils, in colonies of O. chelifer under controlled laboratory conditions. We found that ant larvae fed with diaspores of C. canjerana were on average larger and grew better than control larvae. In conclusion, the data presented here reinforce the importance of secondary dispersal by ants for seeds primarily dispersed by vertebrates. Our results also indicate the importance of studying the foraging ecology and the influence of fruit consumption on survival and growth of colonies of ants. / Doutorado / Ecologia / Doutor em Ecologia

The external dispersal ("epizoochory") of vascular plant diaspores (seeds and fruits) by roe deer and wild boar, i.e. the most common wild large mammals with a large home range in central Europe, was investigated in a 6.5-km² forest area in NE Germany dominated by mesic deciduous forests. The study involved brushing out the diaspores from the coats and hooves of 25 shot roe deer and nine wild boar. The results were compared with the forest vegetation of the study area. Whilst wild boar transported large amounts of various diaspores in the coat, the significance of roe deer for epizoochory was low due to their sleek fur and different behaviour compared to wild boar. Altogether, 55 vascular plant species were transported externally. Since only a limited number of seeds came from woodland habitats, the open landscape was at least as important as a source of attached seeds as the forest vegetation. Thus, most plant species occurring in the studied forest area, especially characteristic woodland herbs, showed no adaptations to epizoochorous dispersal, although being very abundant in the herb layer. We conclude that hoofed game play a particular role concerning the dispersal of ruderal and grassland species in the agricultural landscape of central Europe. However, the actual spread of some herb species in forests of northern Germany, e.g. <i>Agrostis capillaris</i>, <i>Brachypodium sylvaticum</i>, <i>Deschampsia flexuosa</i>, <i>Galium aparine</i> and <i>Urtica dioica</i>, may be mainly facilitated by wild ungulates. Though dispersal by large mammals is an important mechanism for long-distance dispersal of plants in general, our results suggest that most of the characteristic herb species of mesic deciduous forests have only low epizoochorous dispersal potentials. The implications for nature conservation and silviculture are discussed. / Die Ausbreitung von Gefäßpflanzen-Diasporen (Samen und Früchte) durch äußerliche Anhaftung ("Epizoochorie") an Rehen und Wildschweinen, den beiden häufigsten Schalenwild-Arten in Mitteleuropa, wurde im 6,5 km² großen Forst Brieselang bei Berlin (Bundesland Brandenburg) untersucht, in dem mesophile Laubwälder vorherrschen. Dazu wurden die Felle und Hufe von 25 geschossenen Rehen und neun Wildschweinen ausgekämmt und die Diasporen anschließend bestimmt. Die Ergebnisse wurden mit der Waldvegetation verglichen. Während Wildschweine große Mengen verschiedener Diasporentypen transportierten, war die Bedeutung von Rehen für die Ausbreitung von Pflanzen auf Grund des glatten Fells und der im Vergleich zum Wildschwein unterschiedlichen Verhaltensweisen wesentlich geringer. Insgesamt wurden 55 Phanerogamenarten epizoochor transportiert. Da nur ein kleiner Teil der ausgebreiteten Pflanzen Waldhabitate bevorzugt, war das Offenland eine mindestens ebenso wichtige Quelle anhaftender Diasporen wie die Waldvegetation. Die meisten Waldpflanzenarten wurden nicht ausgebreitet; insbesondere solche Arten, die ausschließlich in Wäldern wachsen, wurden nicht nachgewiesen. Viele Pflanzenarten sind – vermutlich auf Grund ihrer Diasporenmorphologie – weitgehend vom Transport ausgeschlossen, obwohl sie sehr häufig in der Krautschicht des untersuchten Waldes vorkommen. Daher ist Schalenwild in der Agrarlandschaft Mitteleuropas vermutlich vor allem für die Ausbreitung von Ruderal-, Segetal- und Grünlandpflanzen von Bedeutung. Die Ausbreitung einiger Pflanzenarten der Krautschicht in norddeutschen Wäldern z.B. <i>Agrostis capillaris</i>, <i>Brachypodium sylvaticum</i>, <i>Deschampsia flexuosa</i>, <i>Galium aparine</i> und <i>Urtica dioica</i>, könnte jedoch wesentlich auf Schalenwild zurückgehen. Obwohl Großsäuger insgesamt ein wichtiger Vektor für die Fernausbreitung von Pflanzen sind, zeigt unsere Studie, dass die meisten charakteristischen Waldbodenpflanzen mesophiler Laubwälder kaum ausgebreitet werden, also nur ein geringes epizoochores Ausbreitungspotenzial aufweisen. Die Bedeutung der Ergebnisse für den Waldnaturschutz und den Waldbau wird diskutiert.

O modelo tradicional de dinâmica de expansão da floresta com Araucaria sobre os campos do Planalto Sul-brasileiro foi desenvolvido pelo naturalista Roberto Klein (1923-1992), e foi baseado no processo de nucleação florestal em áreas campestres. Estudos visando compreender o processo de estabelecimento de espécies de plantas lenhosas florestais em manchas de floresta com Araucaria isoladas em áreas campestres do Planalto Nordeste do Rio Grande do Sul foram desenvolvidos a partir da análise de padrões de colonização e mecanismos de dispersão de diásporos. Os estudos foram conduzidos no CPCN Pró-Mata PUCRS, em São Francisco de Paula, Estado do Rio Grande do Sul, Brasil. O CCapítulo 1 apresenta os principais eixos teóricos abordados ao longo da tese, seus referenciais históricos e contexto local. No CCapítulo 2 foi avaliado o papel de arbustos e árvores isoladas como sítios de colonização para espécies lenhosas florestais em áreas abertas.Foram encontradas mais plântulas de espécies florestais sob a copa de plantas-berçário (particularmente sob araucárias) do que em áreas abertas e as plântulas apresentaram diásporos dispersos principalmente por vertebrados, indicando a importância de árvores de Araucaria angustifolia como plantas-berçário no processo de colonização de sítios campestres por espécies florestais. Os capítulos 3, 4 e 5 discutem padrões de colonização de manchas florestais por espécies lenhosas florestais e mecanismos de dispersão de espécies florestais colonizadoras de manchas. Foram analisadas manchas florestais de tamanhos diferentes em área de campo nativo. No Capítulo 3 foram analisadas a composição de espécies (abundância relativa), a riqueza, a diversidade e os padrões de colonização em comunidades de plantas jovens nas manchas. Riqueza e diversidade foram positivamente correlacionadas com aárea da mancha; além disso, a ordenação das unidades amostrais de acordo com a abundância relativa de indivíduos jovens de espécies lenhosas indicou um gradiente nas comunidades de plantas jovens associado com a área da mancha. A razão entre a abundância de espécies residentes (presentes nas comunidades tanto como indivíduos jovens quanto como adultos) e colonizadoras (presentes nas comunidades somente como indivíduos jovens) aumentou de plantas-berçário para manchas grandes. Nestas últimas, o predomínio de espécies residentes nas comunidades de indivíduos jovens dificulta a imigração de novas espécies para as manchas, limitando a riqueza e a diversidade. No Capítulo 4 foram analisadas as estratégias de dispersão das espécies lenhosas colonizadoras das manchas.Foi avaliada a influência da filogenia e atributos de diásporos relacionados à atração de dispersores (DAT) sobre o trade-off tamanho de semente/número de sementes (SSNT) em espécies lenhosas colonizando manchas florestais, e em comunidades de plantas jovens presentes em manchas florestais de tamanhos diferentes. A alocação de energia para dispersão (número de sementes) em detrimento de sobrevivência da prole (tamanho das sementes) aumentou o sucesso no estabelecimento de espécies colonizadoras em manchas florestais somente quando as influências da filogenia e do DAT foram controladas através de análise de resíduos. A área da mancha exerceu uma pressão seletiva sobre o tamanho das sementes, especialmente quando as influências de filogenia e DAT foram removidas. No Capítulo 5 foi analisada a influência das interações entre plantas colonizadoras e frugívoros dispersores sobre a colonização de manchas florestais.A maioria das espécies lenhosas colonizando manchas de floresta com Araucaria apresentaram diásporos associados com dispersão por vertebrados. Interações entre plantas e frugívoros envolvendo aves de tamanho médio, a maioria frugívoras, associadas com habitats de borda e hábito migratório correlacionaram-se positivamente com manchas pequenas e grandes, enquanto aquelasenvolvendo aves residentes de tamanho grande, em sua maioria não-frugívoras e habitantes de áreas abertas e florestas associaram-se com plantas-berçário. Nossos resultados sugeriram um trade-off envolvendo a quantidade de reservas contidas na semente e a probabilidade do diásporo ser transportado por um frugívoro. No Capítulo 6, uma ampliação do modelo original de Klein é sugerida, e inclui outros níveis tróficos no processo de nucleação de manchas de floresta com Araucaria sobre os campos do Planalto Sul-brasileiro. / The traditional model of expansion dynamics of the Araucaria forest over Campos grassland found in the southern Brazilian highlands was developed by the naturalist Roberto Klein (1923-1992), and was based on the forest nucleation process over grassland sites. Studies concerning the establishment of forest woody plant species into Araucaria forest patches isolated in grassland sites occurring in the Northeast Plateau of the Rio Grande do Sul were developed from analises of plant colonization patterns and diaspore dispersal mecanisms. Studies were conducted at the CPCN Pró-Mata PUCRS, São Francisco de Paula, Rio Grande do Sul State, Brazil. The Chapter 1 presents the main theoretical axes covered along the thesis, its historical backgrounds and its local context. In the Chapter 2 was evaluated the role of isolated shrubs and trees as colonization sites for forest woody species in open areas.We found more forest species seedlings beneath the nurse plant crowns (particularly beneath Araucaria trees) than in open areas, and the seedlings presented diaspores mainly dispersed by vertebrates, indicating the importance of Araucaria angustifolia trees as nurse plants in grassland colonization process by forest species.The chapters 3, 4 e 5 discuss patch colonization patterns and dispersal mecanisms of colonizer species in patches. Forest patches of different sizes and occurring in a native grassland were analyzed. In the Chapter 3 we analyzed the species composition (relative abundance), richness, diversity and colonization patterns in sapling communities in the patches. Richness and diversity were positively correlated with patch area; furthermore, the ordination of the sampling units according to the relative abundance of saplings from woody species has indicated gradient in the sapling communities associated with patch area.The ratio among theabundance of resident species (those present in the communities both as saplings and adult individuals) and the abundance of colonizer species (those present in the communities only as saplings) increased from nurse plants to large patches. In the last, the prevailence of resident adults in the sapling communities made difficult the immigration of new species into the patches, limiting richness and diversity. In the Chapter 4, the dispersal strategies of woody species colonizing patches were analyzed. We evaluated the influence of phylogeny and diaspore attributes related to disperser attraction (DAT) on the seed size/number trade-off (SSNT) in woody species colonizing forest patches, and in sapling communities present in forest patches of different sizes.Energy allocation for dispersal (seed number) in detriment of offspring survival (seed size) increased the establishment success of the colonizer species in forest patches only when the influences of phylogeny and DAT were controlled through residual analysis. Patch area exerted a selective pressure on seed size, especially when the influences of phylogeny and DAT were removed. In the Chapter 5 we analyzed the influence of interactions between colonizer plants and disperser frugivores in the forest patch colonization. Most woody species colonizing Araucaria forest patches presented diaspore associated with vertebrate dispersal. Interactions between plants and frugivores involving medium-sized, mostly frugivorous birds, associated with edge habitats and migratory habit positively correlated with small and large patches, while those involving large-sized, resident, mostly non-frugivorous birds inhabiting open areas and forests were associated with nurse plants. Our results suggested a trade-off involving the seed reserve content and the probability of the diaspore being transported by a given frugivore. In the Chapter 6, an extension of the Klein´s original model is suggested, and includes other trophic levels in the process of Araucaria forest patch nucleation over the Campos grassland of the South Brazilian Plateau.