Biological invasions are reportedly one of the major contributory factors to biodiversity loss worldwide. The impacts of invasive alien plant (IAP) species on native communities are widely documented in the scientific literature, however, there is still a lack of detailed information on their impacts within the most vulnerable habitats. Riparian habitats are highly dynamic systems and naturally disturbed, making them particularly vulnerable to invasion. Climate change, directly or indirectly, is also predicted to adversely impact river systems, which may subsequently alter invasion rates and the impacts of IAPs. However, the interactions between climate and IAPs and their combined effects on vegetation have rarely been examined. To address these knowledge gaps, this thesis investigates: (1) the role of environmental variables, such as sediment loading or climate-related changes to river flow regime, on the abundance of IAPs within riparian zones; (2) how variation in IAP abundance impacts native vegetation, relative to the effects of native dominant plant species and (3) some of the mechanisms underlying the effects of IAPs in riparian habitats. Historic and recent field survey data were used to investigate changes in riparian vegetation on British rivers during the last 20 years. Analyses indicate that IAPs had a negative but small effect on native plant diversity. Overall, changes in land use and differences in flow regime between recording periods were the most important predictors of plant community change. Specifically, IAPs had a greater probability of being present along lowland rivers that experienced increased frequency of high flow events. On a local scale across rivers in Scotland, the abundance of IAPs was constrained by greater soil moisture in summer, whilst greater abundance was associated with tree-lined banks. Both native dominant species and IAPs negatively affected subordinate species abundance to a greater extent than species richness, although this effect varied spatially with bank elevation. Artificial turf mats were used to quantify viable propagules within riverine sediment deposited over-winter along invaded riverbanks. The data indicate that there is a legacy effect of IAP abundance, with the most invaded sites being associated with higher sediment loading the following year, though, contrary to the general pattern, 12 sediment associated propagules were scarcer at invaded sites. Moreover, lower above-ground native diversity was associated with sites which had been previously invaded. Plant species composition in the propagule bank and above-ground vegetation were highly dissimilar, particularly closest to the water’s edge at highly invaded sites. This suggests that mono-specific stands of IAPs proliferate best under less disturbed environmental conditions, although fluvial disturbance events may be required to create opportunities for initial establishment. The propagule bank contributed very little to the above-ground vegetation, nor did it limit invasion, suggesting that above-ground plant composition is largely dictated by competitive interactions. The findings presented in this thesis suggest that invasion by IAPs is an additional stressor for native vegetation within riparian habitats, modifying above-ground plant communities via competition and suppressing recruitment from the propagule bank. However, native dominant species common in riparian habitats also negatively impact, subordinate species via competition, in some cases equalling the effect of IAPs. Native dominant and IAP species are differently affected by environmental factors operating in the riparian zone, which may provide future opportunities for reducing and managing invasions.

Various aspects of myrmecochory were investigated. Effect of different storage methods on atractiveness of seeds with elaiosomes for ants was examined. Viable seed bank of refuse piles (places where ants deposit unused objects from ant nests, e.g. seeds after the elaiosome was eaten) and places outside them were compared. Chemical content of five major chemical groups, specifically amino acids, free fatty acids, organic acids, polyols and sugars, of elaiosomes and seeds of selected species was determined, and compared with respect to their taxonomic relatedness.

Various aspects of myrmecochory were investigated. Effect of different storage methods on atractiveness of seeds with elaiosomes for ants was examined. Viable seed bank of refuse piles (places where ants deposit unused objects from ant nests, e.g. seeds after the elaiosome was eaten) and places outside them were compared. Chemical content of five major chemical groups, specifically amino acids, free fatty acids, organic acids, polyols and sugars, of elaiosomes and seeds of selected species was determined, and compared with respect to their taxonomic relatedness.

Conselho Nacional de Desenvolvimento Científico e Tecnológico / The ecology of regeneration and population dynamics is necessary information to the conservation and management of forest ecosystems. Thus, this study aimed to characterize: a) the mechanisms of natural regeneration in a Mixed Rain Forest remnant and b) the population structure of Araucaria angustifolia (Bertol.) Kuntze, Blepharocalyx salicifolius (Kunth) O. Berg and Ilex paraguariensis A. St.-Hil. Therefore, the structure of the sample project PELD/CNPq was utilized, which was installed in the National Forest of São Francisco de Paula, RS, Brazil to evaluate six clusters (1 ha), with 16 plots (20 x 20m) each, disregarding 10m surround. The mechanisms of regeneration that have been evaluated were seed rain (SR), soil seed bank (SB) and a seedling bank (SeB) (individuals ≥ 30cm in height and diameter at breast height (DBH) <1cm). Additionally, natural regeneration establishment (RNE) (1 ≤ DBH <4.8 cm) sampled in cells of 3.16 x 3.16m, the size class I (SC I) ( 4.8 ≤ DBH <9.55 cm) in plots 10 x 10m and SC II (DBH ≥ 9.55 cm) shown on 96 plots of 20 x 20m were also considered for the study of populations. These data were collected in 2008, also being measured the DBH and sociological position. The mechanisms of regeneration were evaluated for the floristic composition and horizontal structure. In addition, the cluster analysis method TWINSPAN (Two-way Indicator Species Analysis) and canonical correspondence analysis (CCA) were performed for the SeB. The population structure of the three species was examined by characterization of the sociological position, spatial distribution and frequency in size classes. In SR, we sampled 81 species, 46 genera and 28 families, in SB, 103, 65 and 41 and in SeB, 88, 57 and 36, respectively. The analysis by TWINSPAN ordered the formation of three groups in SeB, with their history of interference and environmental conditions. The CCA indicated the slope as environmental variable with the greatest influence on the distribution of plots and species. Group 1 had higher deposition of seeds, however, the lowest densities for the SB and SeB, which is correlated with a decrease in slope. The highest values in SR, SB and SeB were observed in Group 2, with the highest level of anthropogenic interference. Group 3, with lower SR and intermediate density for the other mechanisms, had its plots distributed in accordance with the increase in slope. Allophylus edulis (A. St.- Hil., Cambess & A. Juss.) Radlk., Piper xylosteoides Steud. and Piper aduncum L. showed positive correlation with the increase in slope. The population structure of the three species that were studied showed clustered distribution. Araucaria angustifolia, which is an emerging species, showed high density of individuals regenerating in environments with higher levels of alteration. Blepharocalyx salicifolius and Ilex paraguariensis, have distributed themselves predominantly in the upper and middle strata, and have shown higher population stability within the group on more advanced successional stage. Consequently, it is possible to conclude that the groups determine different patterns in forest regeneration and the structure of populations; therefore, they should be considered in case of possible interventions in the ecosystem. / A ecologia da regeneração e a dinâmica de populações constituem informações necessárias à conservação e ao manejo dos ecossistemas florestais. Assim, este estudo teve como objetivos caracterizar: a) os mecanismos de regeneração natural em um remanescente de Floresta Ombrófila Mista; e b) a estrutura populacional de Araucaria angustifolia (Bertol.) Kuntze, Blepharocalyx salicifolius (Kunth) O. Berg e Ilex paraguariensis A. St. Hil. Deste modo, utilizou-se a estrutura amostral do projeto PELD/CNPq, instalada na Floresta Nacional de São Francisco de Paula, RS, Brasil, sendo avaliados seis conglomerados (1ha), com 16 parcelas (20 x 20m) cada, desconsiderando 10m de bordadura. Os mecanismos de regeneração avaliados foram chuva de sementes (CS), banco de sementes do solo (BS) e banco de plântulas (BP) (indivíduos com altura ≥ 30cm e diâmetro à altura do peito (DAP) < 1cm). Adicionalmente, para o estudo das populações, considerou-se: a regeneração natural estabelecida (RNE) (1 ≤ DAP < 4,8cm) amostrada em células de 3,16 x 3,16m; a classe de tamanho I (CT I) (4,8 ≤ DAP < 9,55cm) em subparcelas 10 x 10m; e a CT II (DAP ≥ 9,55cm) inventariada nas 96 parcelas de 20 x 20m. Esses dados foram coletados em 2008, sendo mensurados o DAP e a posição sociológica. Os mecanismos de regeneração foram avaliados quanto à composição florística e estrutura horizontal. Além disso, para o BP foi realizada a análise de agrupamento, pelo método TWINSPAN (Two-way Indicator Species Analysis) e análise de correspondência canônica (CCA). A estrutura populacional das três espécies foi analisada pela caracterização da posição sociológica, distribuição espacial e frequência em classes de tamanho. Na CS, foram amostradas 81 espécies, 46 gêneros e 28 famílias, no BS, 103, 65 e 41 e no BP, 88, 57 e 36, respectivamente. A análise pelo TWINSPAN determinou a formação de três grupos no BP, com respectivos históricos de interferência e condições mbientais. A CCA indicou a declividade como variável ambiental com maior influência sobre a distribuição das parcelas e espécies. O Grupo 1 apresentou elevada deposição de diásporos, porém, as menores densidades para o BS e BP, estando correlacionado com a diminuição na declividade. Os maiores valores, na CS, BS e BP foram observados no Grupo 2, com maior nível de interferência antrópica. O Grupo 3, com menor CS e densidade intermediária para os demais mecanismos, teve suas parcelas distribuídas de acordo com o aumento na inclinação do terreno. Allophylus edulis (A. St.-Hil., Cambess & A. Juss.) Radlk., Piper xylosteoides Steud. e Piper aduncum L. apresentaram correlação positiva com o aumento na declividade. A estrutura populacional das três espécies estudadas indicou distribuição espacial agregada. Araucaria angustifolia, espécie emergente, apresentou elevada densidade de indivíduos regenerantes em ambientes com maior nível de alteração. Blepharocalyx salicifolius e Ilex paraguariensis distribuíram-se, predominantemente, nos estratos superior e médio respectivamente, e apresentaram maior estabilidade populacional no grupo em estádio sucessional mais avançado. Pelo exposto, concluí-se que os agrupamentos determinam padrões diferenciados, na regeneração da floresta e na estrutura das populações, devendo ser considerados em caso de possíveis intervenções no ecossistema.

Os ecossistemas de ilhas, continentais ou oceânicas, são considerados os mais sensíveis à perturbação humana. A maioria das ilhas costeiras do Brasil sofreu fortes alterações na fauna e flora silvestre. A Ilha Anchieta (Ubatuba, SP) é uma Área Protegida com um longo histórico de perturbação, tendo sido amplamente desmatada no passado e sofrido a introdução de animais. Sabe-se que a fauna possui um papel chave na composição e estrutura da comunidade vegetal, favorecendo algumas espécies e prejudicando outras. Sendo assim, a perda ou alteração dos processos de interações entre animais e plantas afetam na estrutura e composição de espécies. Este estudo analisa as interações antagônicas entre planta e animais como possíveis fatores limitantes no processo de regeneração natural em três ambientes com distintos estágios sucessionais presentes na Ilha Anchieta: campo aberto (CA) e florestas ombrófilas rala (FOR) e densa (FOD). Para isso foram testados nos três ambientes os processos pós-dispersão de sementes: a predação de sementes pós-dispersas, a germinação do banco de sementes e a herbivoria. A predação de sementes apresentou forte efeito espacial (FOD>FOR>CA) e sazonal, com maiores proporções de predação de sementes no período de escassez de alimento (estação seca). Quantitativamente o banco de sementes não representa um fator limitante, porém foi composto por uma baixa diversidade de espécies lenhosas. A capacidade de germinação do banco de sementes do solo foi similar entre os ambientes florestais apresentando menor emersão de plântulas no campo aberto, provavelmente associado ao intenso escoamento superficial no local. Adicionalmente, a mortalidade de juvenis transplantados foi de 72,27%, sendo superior nas parcelas abertas aos vertebrados para todas as espécies em todos os ambientes, demonstrando um forte efeito negativo dos herbívoros vertebrados na comunidade vegetal. Sendo assim, foram detectados distintos gargalos atuando simultaneamente na regeneração natural nos três ambientes da Ilha Anchieta, sendo necessário o estabelecimento de práticas de manejo visando minimizar os efeitos abióticos no campo aberto e floresta ombrófila rala, favorecendo as interações mutualísticas e inibindo a atividade de animais antagônicos nas florestas ombrófilas rala e densa, acelerando assim o processo de regeneração natural na Ilha Anchieta. Considerar simultaneamente os obstáculos no processo de regeneração nos auxiliará a traçar práticas de restauração e recuperação de áreas degradadas mais efetivas e viáveis economicamente. / Island ecosystems, either continental or oceanic, are considered the most sensible to anthropogenic influences. Most Brazilian coastal islands have their original fauna and flora composition altered. The Anchieta Island (southeast Brazil) is a Protected Area that suffered deforestation and introduction of alien species. It is known that the fauna plays a key role in composing and structuring the vegetal community, favoring some species and harming others. Thus, the loss or change of animal-plant interactions affects the framework and composition of species. We simultaneously analyzed the antagonistic animal-plant interactions as possible limiting factors in the natural regeneration on tree successional stages of the Anchieta Island: the old fields (OF), the early secondary forest (ESF) and old growth forest (OGF). Therefore, in each habitat we are evaluating post-dispersal seed predation processes: post-dispersal seed predation, soil seed bank, and the herbivory. We found spatial (OGF>ESF>OF) and temporal effects on seed predation, with highest rates on dry season. Quantitatively, soil seed bank did not represent a limiting factor; but qualitatively, it was composed by few woody species. Our data pointed to the absence of a viable soil seed bank in the OF, probably due to an intense runoff. The forested habitats presented similar soil bank. Moreover, the total mortality of saplings was 72.27%, being higher on the unfenced treatment for all species and in all habitats, showing a very strong negative effect of vertebrate herbivores on the vegetal community. Therefore, we are detecting distinct bottlenecks acting simultaneously in the natural regeneration process in all of the successional stages of the Anchieta Island. Our results showed that management actions are required, aiming to both minimize the abiotic effects on the old fields and in the early secondary forest and to favor the mutual interactions and inhibit the activity of antagonic animals in the old growth and early secondary forests, accelerating, this way, the natural regeneration process of the Anchieta Island. Considering simultaneously the obstacles in the regeneration process will help to define restoration and recuperation procedures of degraded areas more effective and affordable.

Ziel des Forschungsprojektes war der Gewinn umfassender verjüngungsökologischer Kenntnisse zu den Pionierbaumarten Sandbirke (Betula pendula Roth), Salweide (Salix caprea L.) und Eberesche (Sorbus aucuparia L.) im Hinblick auf eine natürliche, eingriffsfreie Wiederbewaldung von Schadflächen (z.B. nach Sturmwurf). Die Abschätzung des Besiedlungserfolges von Schadflächen durch Pionierbaumarten ist aufgrund unzureichender verjüngungsökologischer Kenntnisse gegenwärtig noch mit großen Unsicherheiten verbunden. Die Untersuchungen fanden auf fünf 4-12 ha großen Kyrill-Sturmwurfflächen in den Hoch- und Kammlagen (750-900 m ü. NN) des Thüringer Waldes statt. Alle potenziellen Samenbäume der Pionierbaumarten wurden in den angrenzenden, geschlossenen Fichtenbeständen lokalisiert. Als Versuchsdesign wurde in Abhängigkeit der vorgefundenen Samenbaumdichten und -verteilungen ein Kreuz- bzw. Sterntransekt auf den Sturmwurfflächen etabliert. Entlang der Transektlinien wurden alle 20 m Samenfallen installiert. Als Samenfallen kamen für die Sandbirke Netztrichterfallen (0,2 m²), für die Salweide Klebfallen (0,043 m²) und für die endozoochore Ausbreitung durch frugivore Vogelar-ten Kotfallen (0,25 m²) zum Einsatz. Für die Modellierung der Samenausbreitung von Sandbirke und Salweide wurden inverse Modelle bzw. geostatistische Modelle erstellt. Zudem wurden auf einer der Sturmwurfflächen genetische Nachkommenschaftsanalysen bei Sal-weide mittels Kern-DNA-Primer durchgeführt. Die Bodensamenbankuntersuchungen fanden in jeweils drei geschlossenen Birkenbeständen, Fichten-Birken-Beständen, Fichtenbeständen mit einer einzeln eingemischten Birke und reinen Fichtenbeständen im Tharandter Wald und Thüringer Wald statt. Mittels eines 10,2 cm breiten Stechzylinders wurden 10 cm tiefe Bodenproben gewonnen. Die Lagerung der Proben fand im Kaltgewächshaus statt, wo alle 14 d die gekeimten Samen erfasst wurden. Weiterhin wurde ein Eingrabungsexperiment installiert. Dafür wurden Sandbirkensamen, Ebereschensamen und Ebereschenfrüchte in 2 cm, 5 cm und 10 cm tiefen Mineralboden vergraben und in sechsmonatigen Intervallen jeweils eine Keimprobe zum Test der verbliebenen Keimfähigkeit entnommen. Die Auswertung der Bodensamenbankuntersuchungen erfolgte mittels GLM und GLMM. Während der zweijährigen Untersuchung zur zeitlichen und räumlichen Samenausbreitung von Salix caprea auf fünf Sturmwurfflächen konnten ein schwächeres und ein stärkeres Samenjahr nachgewiesen werden. Des Weiteren erstreckte sich der Samenflugzeitraum im Frühjahr in Abhängigkeit von den klimatischen Bedingungen über 12 Wochen in 2015 und 6 Wochen in 2016. Die höchsten Samenmengen von 23-156 Samen je Falle wurden jeweils unter den Kronen von Salweiden-Samenbäumen nachgewiesen. Ab einer Entfernung von 350 m zum Samenbaum bis zur untersuchten Distanz von 870 m wurden, unabhängig von der Distanz, der Hangneigung, der Anzahl der Samen-bäume und der Windrichtung (Anisotropie), im Durchschnitt 0,6-2,1 Samen je Falle er-fasst. Die genetischen Analysen zur Nachkommenschaft ergaben, dass 29 % der untersuch-ten Verjüngungspflanzen von einem der 20 lokalisierten Elternbäume in der bewaldeten, 500 m breiten Suchzone abstammten. Die Ausbreitungsdistanzen der nachweislich am erfolgreichsten verjüngten Samenbäume betrugen dabei 550-800 m. Insgesamt zeigte die Salweidenverjüngung eine höhere Allel-Variation, als die 20 Elternbäume, was auf einen externen Genfluss und lange Samen- und Pollenausbreitungsdistanzen hinweist. Im Zuge des zweijährigen Untersuchungszeitraums zur Samenausbreitung von Betula pendula auf zwei Kyrill-Sturmwurfflächen konnten ein Mastjahr und ein Zwischenjahr nachgewiesen werden. Die Ergebnisse der inversen Modellierung mittels isotroper Mo-delle ergaben dabei flächenunabhängig Produktionsmengen für einen Samenbaum von 20 cm im Bhd von 300.000-366.000 Samen je Baum im Zwi-schenjahr 2015 und 1.430.000-1.530.000 Samen je Baum im Mastjahr 2016. Mittels räumlicher Modellierung der Samenausbreitung konnte keine Anisotropie belegt werden. Unabhängig von den beprobten Flächen und Un-tersuchungsjahren, belegen die Modellschätzungen allesamt eine isotrope Ausbreitung der Samen. Die mittleren Ausbreitungsdistanzen (MDD) beliefen sich dabei hangaufwärts auf 86-97 m und hangabwärts auf 367-380 m. Maximal ab-gelagerte Samendichten von 2.015 n m-² im Zwischenjahr und 9.557 n m-² im Mastjahr fanden sich bis 40-50 m Entfernung zum Samenbaum. Die Untersuchungen der endozoochoren Samenausbreitung auf fünf Sturmwurfflächen weisen auf eine bevorzuge Nutzung der Vogelarten von Rast- und Sitzgelegenheiten (Strukturelemente) auf Freiflächen zum Absetzen von Kot hin (2,7 Kothaufen je m²). Unter Freiflächenbedingungen - ohne Strukturelemente - ergaben sich im Mittel 0,4 Kothaufen je m². Die höchsten mittleren Kotdichten wurden unter aufra-genden Totästen (20 n m-²), umgeklappten Wurzeltellern (4,6 n m-²) und Hochstubben (3,9 n m-²) nachgewiesen. Schwach dimensionierte Verjüngungspflanzen der Sandbirke, Eberesche und Fichte, und Strukturelemente unter einem Meter Höhe wurden dagegen weitgehend für das Absetzen von Kot gemieden. Das Vermögen zum Aufbau einer Bodensamenbank durch Betula pendula und Sorbus aucuparia unterschied sich deutlich. 56-100 % der eingegrabenen Sandbirkensamen wa-ren auch nach 2,5 Jahren keimfähig, wohingegen lediglich 3-16 % der eingegrabenen Ebereschensamen ohne Fruchthülle und 0-19 % der eingegrabenen Ebereschensamen mit Fruchthülle vital waren. Die Auswertung mittels GLM prognostizierte einen kom-pletten Verlust der Keimfähigkeit nach 12 Jahren, 4,5 Jahren und 3 Jahren der Sandbirkensamen, sowie der Ebereschensamen mit und ohne Fruchthülle. Ein Einfluss der Lagerungstiefe war nur für Sandbirke nachweisbar. Die Untersuchungen der Bodensamenbank von Birke in Fichtenbeständen mit unter-schiedlichen Birkensamenbaumanteilen ergab einen straffen Zusammenhang zwischen der Anzahl von Samenquellen und den nachgewiesenen Samendichten im Boden. In den Birkenbeständen fanden sich stets die höchsten Dichten von 489-1.142 Birkensamen je m². Die Analyse unterschiedlicher Bodenschichten zeigte zudem signifikant abneh-mende Birkensamendichten mit zunehmender Bodentiefe. Die Ergebnisse der Untersuchungen zeigen, dass die Fruktifikation von Betula pendula, Salix caprea und Sorbus aucuparia durch klimatische Verhältnisse beeinflusst wird, weshalb die drei Pionierbaumarten nicht alljährlich hohe Samenmengen produzieren (Mastjahre und Zwischenjahre). Zum Ausgleich von Produktionsdefiziten in den Zwischenjahren unter-scheiden sich die Pionierbaumarten in ihrer Strategie. Dies gilt es bei der Umsetzung des Konzeptes einer natürlichen, eingriffsfreien Wiederbewaldung von Schadflächen nach Sturmwurf durch die Naturverjüngung von Pionierbaumarten zu beachten. Die einzige der drei Pionierbaumarten, die dem allgemeinen Bild einer Pionierbaumart ent-spricht, ist die Salweide. Ihr Besiedlungserfolg ist allein von den aktuellen, alljährlich variierenden, aber dennoch stets hohen Samenproduktionsmengen und den enorm weiten Aus-breitungsdistanzen (>800 m) abhängig. Hinsichtlich der Samenausbreitung haben die Him-melsrichtung, die Position der Samenbäume und die Anzahl vorhandener Samenquellen ab einer Distanz von 50 m zur Schadfläche keinen bedeutenden Einfluss auf die abgelagerten Samenmengen mehr. Die auf 86-380 m limitierte Samenausbreitung von Sandbirke wurde dagegen stark vom Geländerelief (Hangneigung), der Position der Samenbäume (Tal, Kuppe, Hanglage) und der Distanz der Samenbäume zur Sturmwurffläche beeinflusst. Zum Ausgleich der limitierten Samenausbreitung und deutlich reduzierten Samenmengen im Zwischenjahr ist Sandbirke jedoch zum Aufbau einer short-term persistenten Bodensamenbank befähigt. Den gesamten Verjüngungszyklus betrachtend entspricht die Eberesche eher einer Schluss-waldbaumart. Unter ungünstigen klimatischen Bedingungen kommt es häufig zum kompletten Ausfall der Samenproduktion. Ihr enormes Wiederbewaldungspotential von Sturmwurfflächen speist sich hauptsächlich aus dem Aufbau einer Sämlingsbank und weni-ger durch den aktuellen Samenregen oder der short-term persistenten Bodensamenbank. Die limitierte Samenausbreitung von Sandbirke und Eberesche macht eine „räumliche Optimierung“ der Samenbaumpositionen durch die Forstwirtschaft erforderlich. Aufgrund der allgegenwärtigen Omnipräsenz von Weidensamen ist dies für Salweide nicht zwingend not-wendig. Das detailreiche Wissen zur Verjüngungsökologie der untersuchten Pionierbaumar-ten ermöglicht eine gezielte waldbauliche Steuerung im Sinne des Vorhalts und der Pflege von Pionierbaumarten im Wirtschaftswald. Dies ist gegenwärtig und zukünftig vor allem von besonderer Bedeutung, da aufgrund der zu erwartenden Zunahme von Schadereignissen und deren Unvorhersehbarkeit die Fähigkeit der Wälder zur natürlichen Wiederbewaldung von Schadflächen durch Pionierbaumarten zunehmend an Interesse gewinnen wird.:Table of abbreviations III Summary IV Zusammenfassung VIII Chapter 1 – General introduction 1 1.1 Introduction 2 1.1.1 Importance and relevance of the study 2 1.1.2 Research interest - regeneration ecology 5 1.3 Aims, scope and hypotheses 9 1.4 Study outline 12 1.5 References 13 Chapter 2 – Seed dispersal capacity of Salix caprea L. assessed by seed trapping and parentage analysis 25 2.1 Abstract 26 2.2 Introduction 26 2.3 Materials and methods 29 2.3.1 Study area 29 2.3.2 Experimental design 31 2.3.3 Genetic analysis 32 2.3.4 Seed trap data analysis 33 2.3.5 Geostatistical models 34 2.4 Results 36 2.4.1 Temporal patterns of seed dispersal 37 2.4.2 Dispersal distance and spatial patterns of seed dispersal 37 2.4.3 Genetic parentage analysis 40 2.5 Discussion 42 2.5.1 Seed production and temporal patterns of seed dispersal 42 2.5.2 Dispersal distance and spatial patterns of seed dispersal 43 2.5.3 Genetic parentage analysis 45 2.6 Conclusions for silvicultural practice 46 2.7 References 48 Chapter 3 – Restrictions on natural regeneration of storm-felled spruce sites by silver birch (Betula pendula Roth) through limita-tions in fructification and seed dispersal 57 3.1 Abstract 58 3.2 Introduction 58 3.3 Materials and methods 60 3.3.1 Study area 60 3.3.2 Experimental design 63 3.3.3 Data analysis 64 3.3.4 Seed dispersal model 64 3.3.5 Simulations for practical management decisions 66 3.4 Results 66 3.4.1 Seed production 66 3.4.2 Seed dispersal and spatial patterns 68 3.5 Discussion 71 3.5.1 Seed production 71 3.5.2 Directionality 72 3.5.3 Spatial patterns and seed dispersal distances 72 3.6 Seed dispersal scenarios for silvicultural management decisions 74 3.6.1 Seed dispersal scenarios 74 3.6.2 Conclusions for silvicultural management decisions 76 3.7 References 78 Chapter 4 – The impact of structural elements on storm-felled sites on endozoochorous seed dispersal by birds – a case study 85 4.1 Abstract 86 4.2 Introduction 86 4.3 Materials and methods 88 4.3.1 Study area 88 4.3.2 Experimental design 90 4.3.3 Data analysis 91 4.4 Results 92 4.5 Discussion 95 4.6 Conclusions for silvicultural practice 97 4.7 References 98 Chapter 5 – Soil seed banks of pioneer tree species in European tempe-rate forests: a review 104 5.1 Abstract 105 5.2 Introduction 105 5.3 Methods of literature search 107 5.4 Species-specific reproductive ecology determining the potential of soil seed banks 110 5.5 Characterization and classification of soil seed banks 112 5.5.1 Soil seed bank of Betula spp. 114 5.5.2 Soil seed bank of Alnus glutinosa (L.) GAERTN. 116 5.5.3 Soil seed banks of Salix spp. and Populus tremula L. 117 5.5.4 Soil seed bank of Sorbus aucuparia L. 118 5.6 Conclusions 119 5.7 References 120 Chapter 6 – Do birch and rowan establish soil seed banks sufficient to compensate for a lack of seed rain after forest disturbance? 134 6.1 Abstract 135 6.2 Introduction 135 6.3 Materials and methods 137 6.3.1 Study areas 137 6.3.2 Data collection 139 6.3.3. Statistical analysis 140 6.4 Results 141 6.4.1 Study A - Artificial seed burial experiment 141 6.4.2 Study B - Soil core sampling in the forest 145 6.5 Discussion 147 6.5.1 Study A - Artificial seed burial experiment 147 6.5.2 Study B - Soil core sampling in the forest 149 6.6 Conclusions 151 6.7 References 152 Chapter 7 – General discussion 160 7.1 Discussion of important aspects of regeneration ability 161 7.1.1 Fructification and seed production in Salix caprea, Betula pendula and Sorbus aucuparia 165 7.1.2 Ecological processes within the regeneration cycle of Salix caprea 167 7.1.3 Ecological processes within the regeneration cycle of Betula pendula 168 7.1.4 Ecological processes within the regeneration cycle of Sorbus aucuparia 169 7.2. Conclusions for silviculture and management recommendations 172 7.3 References 175 Table of appendix i / The aim of the study was to obtain comprehensive knowledge of the regeneration ecology of the pioneer tree species silver birch (Betula pendula Roth), goat willow (Salix caprea L.) and rowan (Sorbus aucuparia L.). The findings should contribute to better management of the natural regeneration of disturbed sites (e.g., windthrown sites) by pioneer tree species. Insufficient knowledge of the regeneration ecology of pioneer tree species renders forest managers’ abilities to assess the success of regeneration of windthrown sites uncertain. The study took place in the years 2015 and 2016. The study sites were located on the slopes and mountain tops (plateaus) of the Thuringian Forest (715-900 m a.s.l.), on five windthrown open areas (4-13 ha) created by the storm ‘Kyrill’ in January 2007. All seed trees of pioneer tree species were mapped within the forested search zone around each site. This zone extended 200 m for silver birch and rowan and 500 m for goat willow. Following the mapping of these seed trees and an analysis of their spatial distribution, seed traps were placed along two or four crossing line transects, with intervals of 20 m between traps. The traps were funnel shaped net seed traps for silver birch (0.2 m²), seed traps with a sticky, non-drying glue for goat willow (0.043 m²) and dropping traps for seeds dispersed endozoochorously by frugivorous birds (0.25 m²). A phenomenological model and model-based geostatistics were used to investigate silver birch and goat willow seed dispersal. For goat willow a parentage analysis was performed at one of the study sites using nuclear-DNA-primers. The soil seed bank study was carried out in three birch stands, spruce stands with admixed birch, spruce stands with one isolated birch tree and pure spruce stands in the Tharandter Forest and in the Thuringian Forest. Soil core samples with a diameter of 10.2 cm were taken from the litter layer and the mineral soil to a depth of 10 cm. The soil samples were placed in a greenhouse and seed germination was checked every 14 days. An artificial seed burial experiment was also carried out. Silver birch seeds, rowan seeds and rowan fruits were buried in mineral soil at depths of 2 cm, 5 cm and 10 cm. At intervals of 6 months sample sets were removed from the soil and the germination capacity checked. The analysis of the soil seed banks was based on generalized linear mixed models (GLMM) and generalized linear models (GLM). The 2-year study of the temporal and spatial dispersal of seeds of Salix caprea on five Kyrill-felled areas involved one year with lower seed production and one with more bountiful seed production. The duration of the spring seed rain was about 12 weeks in 2015, and only 6 weeks in 2016 because of contrasting weather conditions. The highest seed numbers of 23-156 n per trap occurred close to the base of the seed trees. Beyond 350 m from the seed trees, up to the maximum distance in the study of 870 m, the average numbers of seeds per trap (0.6-2.1 seeds) were independent of the dispersal distance, inclination, the number of seed sources and the dispersal direction (anisotropy). Parentage analyses showed that 29% of the saplings stemmed from one of the 20 parent trees within the 500 m search zone extending from the edge of the open area. The seed dispersal distances of the most successful seed parents were between 550-800 m. The saplings revealed a higher allelic variation than the 20 parent trees, indicating external gene flow and long seed and pollen dispersal distances. During the 2-year study of Betula pendula seed dispersal on two Kyrill-felled areas there was a mast year and a non-mast year. Independent of the site, the seed production rate of a silver birch seed tree with a mean diameter at breast height (dbh) of 20 cm predicted by isotropic inverse models was approximately 300,000-366,000 seeds in 2015 and 1,430,000-1,530,000 seeds per tree in the mast year 2016. Directionality (anisotropic inverse modelling) of seed dispersal around an individual seed tree could not be confirmed. The model results revealed the isotropic model (no directionality) to be an appropriate approach for all sites and years. The mean dispersal distances (MDD) were 86 m and 97 m (uphill) and 367 m and 380 m (downhill). The maximum seed numbers occurred within 40-50 m of a seed tree, amounting to 2,015 n m-² in the non-mast year and 9,557 n m-² in the mast year. The study of endozoochorous seed dispersal on the five sites felled by the storm Kyrill showed a preference of frugivorous birds for perches and resting places (structural elements) from which to defecate onto open areas (2.7 droppings per m²). On completely open areas – with no structural elements – an average of 0.4 droppings per m² was recorded. The highest mean bird dropping density was observed under towering dead branches (20 n m-²), upturned root plates (4.6 n m-²) and high stumps (3.9 n m-²). Young, small diameter silver birch, rowan and spruce trees, and structural elements less than 1 m in height generally, were avoided by frugivorous birds as a place from which to defecate. The abilities of Betula pendula and Sorbus aucuparia to form a soil seed bank differed. Between 56-100 % of the buried silver birch seeds were still viable after 2.5 years, whereas only 3-16 % of the rowan seeds buried without pulp and 0-19 % of the rowan seeds within pulp were viable. The maximum durations of storage in the soil predicted for silver birch seeds and rowan seeds with and without pulp by GLM were 12 years, 4.5 years and 3 years. An influence of the storage depth was found for silver birch seeds only. The investigation of the soil seed banks of birch in three birch stands and nine spruce forests with different numbers of admixed birch seed trees showed a strong correlation between the number of seed sources and the seed density in the soil. The birch stands contained the highest mean densities of viable birch seeds in soil, between 489-1,142 n m-². The analysis of the different soil layers showed significantly declining birch seed densities with increasing soil depth across all sites. The results of the study showed that the fructification of Betula pendula, Salix caprea and Sorbus aucuparia is influenced by weather conditions, with the three pioneer tree species failing to produce high numbers of seeds every year (mast and non-mast years). The three species differed in their strategies to compensate for low seed production in non-mast years. This must be considered when implementing a concept for the reforestation of disturbed sites based on natural regeneration by pioneer tree species. Goat willow was the only one of the three specie studied with characteristics corresponding to the general assumptions made about pioneer tree species. The regeneration success of goat willow is dependent upon the variable but generally high annual seed production and long seed dispersal distances (> 800 m). The azimuth direction, position and number of seed trees have no meaningful influence on seed numbers at a distance of more than 50 m from the seed source. The limited mean seed dispersal distances of 86-380 m determined for silver birch were influenced by site inclination, the seed tree location (valley, slope or plateau) and the distance between the seed tree and the windthrown site. Silver birch seed shadow is also influenced by the number of seed sources. To compensate for the limited dispersal distances and the significantly lower seed production in non-mast years, silver birch is able to build up a short-term persistent soil seed bank. The regeneration cycle of rowan is more reminiscent of that of a shade-tolerant tree species. Unfavorable weather conditions often result in a complete failure to produce seeds. The enormous regeneration potential of rowan on disturbed sites stems primarily from a seedling bank, which is built up over years. The seed rain in any given year and its short-term persistent soil seed bank are of secondary importance. Forest management targeting a ‘spatial optimization’ of silver birch and rowan seed trees is necessary to ensure successful natural regeneration given the limited seed dispersal. The omnipresence of goat willow seeds renders specific spatial management measures for its establishment unnecessary. Detailed knowledge of the regeneration ecology of the studied pioneer tree species makes possible an approach to silviculture that is targeted to the conservation and revitalization of pioneer tree species in managed forests. The expected increase in the frequency of disturbances, and their unpredictability, means that the ability of forests to naturally regenerate using pioneer tree species is likely to grow in importance.:Table of abbreviations III Summary IV Zusammenfassung VIII Chapter 1 – General introduction 1 1.1 Introduction 2 1.1.1 Importance and relevance of the study 2 1.1.2 Research interest - regeneration ecology 5 1.3 Aims, scope and hypotheses 9 1.4 Study outline 12 1.5 References 13 Chapter 2 – Seed dispersal capacity of Salix caprea L. assessed by seed trapping and parentage analysis 25 2.1 Abstract 26 2.2 Introduction 26 2.3 Materials and methods 29 2.3.1 Study area 29 2.3.2 Experimental design 31 2.3.3 Genetic analysis 32 2.3.4 Seed trap data analysis 33 2.3.5 Geostatistical models 34 2.4 Results 36 2.4.1 Temporal patterns of seed dispersal 37 2.4.2 Dispersal distance and spatial patterns of seed dispersal 37 2.4.3 Genetic parentage analysis 40 2.5 Discussion 42 2.5.1 Seed production and temporal patterns of seed dispersal 42 2.5.2 Dispersal distance and spatial patterns of seed dispersal 43 2.5.3 Genetic parentage analysis 45 2.6 Conclusions for silvicultural practice 46 2.7 References 48 Chapter 3 – Restrictions on natural regeneration of storm-felled spruce sites by silver birch (Betula pendula Roth) through limita-tions in fructification and seed dispersal 57 3.1 Abstract 58 3.2 Introduction 58 3.3 Materials and methods 60 3.3.1 Study area 60 3.3.2 Experimental design 63 3.3.3 Data analysis 64 3.3.4 Seed dispersal model 64 3.3.5 Simulations for practical management decisions 66 3.4 Results 66 3.4.1 Seed production 66 3.4.2 Seed dispersal and spatial patterns 68 3.5 Discussion 71 3.5.1 Seed production 71 3.5.2 Directionality 72 3.5.3 Spatial patterns and seed dispersal distances 72 3.6 Seed dispersal scenarios for silvicultural management decisions 74 3.6.1 Seed dispersal scenarios 74 3.6.2 Conclusions for silvicultural management decisions 76 3.7 References 78 Chapter 4 – The impact of structural elements on storm-felled sites on endozoochorous seed dispersal by birds – a case study 85 4.1 Abstract 86 4.2 Introduction 86 4.3 Materials and methods 88 4.3.1 Study area 88 4.3.2 Experimental design 90 4.3.3 Data analysis 91 4.4 Results 92 4.5 Discussion 95 4.6 Conclusions for silvicultural practice 97 4.7 References 98 Chapter 5 – Soil seed banks of pioneer tree species in European tempe-rate forests: a review 104 5.1 Abstract 105 5.2 Introduction 105 5.3 Methods of literature search 107 5.4 Species-specific reproductive ecology determining the potential of soil seed banks 110 5.5 Characterization and classification of soil seed banks 112 5.5.1 Soil seed bank of Betula spp. 114 5.5.2 Soil seed bank of Alnus glutinosa (L.) GAERTN. 116 5.5.3 Soil seed banks of Salix spp. and Populus tremula L. 117 5.5.4 Soil seed bank of Sorbus aucuparia L. 118 5.6 Conclusions 119 5.7 References 120 Chapter 6 – Do birch and rowan establish soil seed banks sufficient to compensate for a lack of seed rain after forest disturbance? 134 6.1 Abstract 135 6.2 Introduction 135 6.3 Materials and methods 137 6.3.1 Study areas 137 6.3.2 Data collection 139 6.3.3. Statistical analysis 140 6.4 Results 141 6.4.1 Study A - Artificial seed burial experiment 141 6.4.2 Study B - Soil core sampling in the forest 145 6.5 Discussion 147 6.5.1 Study A - Artificial seed burial experiment 147 6.5.2 Study B - Soil core sampling in the forest 149 6.6 Conclusions 151 6.7 References 152 Chapter 7 – General discussion 160 7.1 Discussion of important aspects of regeneration ability 161 7.1.1 Fructification and seed production in Salix caprea, Betula pendula and Sorbus aucuparia 165 7.1.2 Ecological processes within the regeneration cycle of Salix caprea 167 7.1.3 Ecological processes within the regeneration cycle of Betula pendula 168 7.1.4 Ecological processes within the regeneration cycle of Sorbus aucuparia 169 7.2. Conclusions for silviculture and management recommendations 172 7.3 References 175 Table of appendix i

Les changements globaux, notamment les changements d'usage des terres, modifient profondément le fonctionnement des écosystèmes et la biodiversité et, ont déjà impacté de nombreux services écosystémiques. La disparition de ces écosystèmes souligne la nécessité de préserver les zones intactes, et la restauration des zones détruites ou perturbées peut permettre de venir en appui aux efforts de conservation et minimiser les dommages. Ce travail a pour objet d’étude les campos rupestres, des pelouses néotropicales d’altitude, faisant partie du Cerrado (savane brésilienne), qui recèlent une importante biodiversité et qui, comme bien d'autres écosystèmes de montagne, fournissent de précieux services écosystémiques tels que la filtration de l’eau. Ils ont été, et sont encore, grandement affectés par les activités humaines (les travaux de génie civil, les carrières ou les mines). Le premier objectif de cette thèse était de décrire l'écosystème de référence, afin de définir clairement un objectif de restauration et mesurer les progrès et le succès de la restauration. Nous avons montré que les campos rupestres sont composés d'au moins deux communautés végétales distinctes (une avec un substrat caillouteux et l’autre avec un substrat sableux), chacune ayant une composition en espèces et une structure particulières ainsi qu’une grande biodiversité. La phénologie reproductive varie au sein des communautés herbacées: la majorité des espèces fleurissent et fructifient pendant la saison des pluies, d'autres se reproduisent en revanche durant la saison sèche. Tout au long de nos 2 années de suivis phénologiques, certaines espèces dominantes, notamment des Poaceae, n'ont pas été observées en fleur impliquant une dispersion limitée de ces espèces vers les zones dégradées. Les communautés végétales de campos rupestres ne sont pas résilientes aux fortes perturbations: plusieurs années après, presque aucune des espèces cibles n’ont été trouvées en zones dégradées, les sols ont complètement été modifiés et les banques de graines ne se sont recomposées qu’avec des espèces rudérales non désirées. Selon le modèle des filtres, une communauté résulte d’un pool régional d’espèce sélectionné par un ensemble de filtres : de dispersion, abiotique et biotique. Les interventions de restauration mises en place avaient pour but d’agir sur ces différents filtres afin de diriger la dynamique des communautés végétales. Nous avons mis en place trois protocoles de restauration in-situ (le transfert de foin, la translocation d’espèce et la translocation de plaque de végétation) pour restaurer les deux types de communautés de campos rupestres identifiées. Le transfert de foin n’a pas permis la restauration des communautés végétales de campos rupestres en raison de l’importante altération des sols et, surtout, à cause de la mauvaise qualité des graines. En effet, nos études de germination ont montré que, alors que certaines espèces de Xyridaceae et Velloziaceae germent très bien, certaines espèces dominantes de Poaceae, de Cyperaceae ou d’Asteraceae ont des graines soit vides, soit non viables, soit dormantes ; le semis se révèle alors peu efficace. Nous n’avons pas mis en évidence d’effet positif du feu sur la germination des espèces de campos rupestres. La translocation d'espèces s’avère un succès pour une seule espèce, Paspalum erianthum. Pour les autres, les dommages causés au niveau des racines lors de la translocation limitent probablement leur survie. Enfin la translocation de plaques de végétation s’avère être la méthode la plus efficace permettant à de nombreuses espèces d’être réintroduites en zones dégradées. En raison de la faible résilience des campos rupestres dans lesquels les plaques de végétation ont été prélevées, cette méthode ne peut être envisagée que pour sauver des habitats dans le cas extrême où leur destruction est inévitable. Face à la difficulté de restaurer les campos rupestres, leur protection et leur conservation doit être une priorité / Global environmental changes, especially land-use changes, have profound effects on both ecosystem functioning and biodiversity, having already altered many ecosystem services. These losses emphasize the need to preserve what remains; however when conservation programs are not sufficient, restoring areas that have been destroyed or disturbed can improve conservation efforts and mitigate damages. This work focuses on campos rupestres, Neotropical grasslands found at altitudes, which are part of the Cerrado (Brazilian savannas). They host a great biodiversity with a high level of endemism and, like other mountain ecosystems, provide valuable ecosystem services, such as water purification and recreational services. They have been and still are being impacted by human activities, such as civil engineering construction, quarrying or mining. The first objective of this thesis was to describe the reference ecosystem in order to aim for a clear restoration target and to monitor progress and success. We show that campos rupestres are composed of at least two distinct plant communities (i.e. sandy and stony grasslands), each having a specific composition and structure, hosting a great biodiversity. Several phenological patterns occur among the herbaceous communities: the majority of species flowers and fruits appear during the rainy season but other patterns can be observed. During our 2-year survey, some dominant species belonging to Poaceae, among others, were not observed reproducing, which implies limited chances to disperse on degraded areas. Campo rupestre vegetation is not resilient following a strong disturbance: several years after the disturbance, almost no native species are encountered on the degraded areas, soils are completely altered and seed bank recomposes only with non-target ruderal species. According to the filter model, a local community is a subset of the regional species pool determined by a set of dispersal, abiotic and biotic filters. Acting on the different filters to influence the plant community was the core of our restoration interventions. We then applied three in-situ restoration protocols (hay transfer, species translocation and turf translocation) to restore both kinds of grassland. Hay transfer does not allow the restoration of campo rupestre vegetation because of soil alteration and mainly because of poor seed quality. Indeed, germination studies show that, while some Xyridaceae and Velloziaceae have a high germinability, some dominant Poaceae, Cyperaceae or Asteraceae species have embryoless, unviable or dormant seeds, which makes seeding less efficient. There is no evidence that fire-related cues enhance germination in campos rupestres. Species translocation is successful for only one species, Paspalum erianthum; for the others, root damages probably impede survival. Finally, turf translocation is the most successful method, since numerous species are re-introduced on degraded areas. However due to the low resilience of pristine campos rupestres where turfs are taken from, turf translocation can only be considered in the case of habitat rescue, in circumstances when complete habitat destruction is otherwise unavoidable. Face to the difficulty to restore these peculiar grasslands, the protection and the conservation of campos rupestres must be made a high priority

No description available.

No description available.

No description available.