The evolution of metabolism regulation is an intertwined process, where different strategies are constantly being developed towards a cognitive ability to perceive and respond to an environment. Organisms depend on an orchestration of a complex set of chemical reactions: maintaining homeostasis with a changing environment, while simultaneously sending material and energetic resources to where they are needed. The success of an organism requires efficient metabolic regulation, highlighting the connection between evolution, population dynamics and the underlying biochemistry. In this work, I represent organisms as coupled information-processing networks, that is, gene-regulatory networks receiving signals from the environment and acting on chemical reactions, eventually affecting material flows. I discuss the mechanisms through which metabolism control is improved during evolution and how the nonlinearities of competition influence this solution-searching process. The propagation of the populations through the resulting landscapes generally point to the role of the rhythm of cell division as an essential phenotypic feature driving evolution. Subsequently, as it naturally follows, different representations of organisms as oscillators are constructed to indicate more precisely how the interplay between competition, maturation timing and cell-division synchronisation affects the expected evolutionary outcomes, not always leading to the \"survival of the fastest\".

In der Dissertation werden Fragen zur Populationsbiologie des neozonalen Nordamerikanischen Waschbären (Procyon lotor) behandelt. Die knapp sechsjährigen Freilanduntersuchungen fanden im Rahmen eines umfangreichen Waschbärenforschungsprojektes (www.projekt-waschbaer.de) in einem naturnahen Tieflandbuchenwald im Müritz-Nationalpark (Mecklenburg-Vorpommern) statt. Das Nationalparkgebiet wird nachweislich seit Ende der 1970er Jahre vom Waschbären besiedelt und stellt aufgrund seines Gewässerreichtums und seiner alten Laubbaumbestände einen idealen Lebensraum für Waschbären dar. Die Dissertation schließt die populationsbiologischen Arbeiten des Gesamtprojektes ab und stellt die Ergebnisse in fünf separaten Themenschwerpunkten vor (I. Raumverhalten, II. Sozialverhalten, III. Reproduktionsbiologie, IV. Populationsstruktur, V. Populationsdynamik). Übergeordnetes Ziel der Arbeit war die Erhebung valider populationsbiologischer Daten, um eine grundlegende ökologische Charakterisierung des Waschbären unter dem Aspekt des Natur- und Artenschutzes vornehmen zu können. Dazu wurden zwischen 2006 und 2011 in einem 1.114 ha großen Fallennetz im Serrahner Teilgebiet des Nationalparks an 53 verschiedenen Fallenstandorten 145 verschiedene Waschbären (62 ♀♀, 83 ♂♂) insgesamt 489 Mal gefangen, genetisch beprobt, vermessen und individuell markiert. 51 adulte Waschbären (23 ♀♀, 28 ♂♂) und 18 Jungtiere (10 ♀♀, 8 ♂♂) wurden darüber hinaus mit einem UKW-Halsbandsender ausgestattet und im Rahmen der telemetrischen Arbeiten insgesamt 31.202 Mal geortet (≙ im Mittel 452 Lokalisationen pro Tier). Im Kernuntersuchungsgebiet wurde an 36 beköderten Standorten ein Fotofallenmonitoring durchgeführt. Bei einer Überwachungsdauer von 5.365 Fotofallennächten entstanden dabei 18.721 Aufnahmen von 183 verschiedenen Waschbären. 82 % aller Waschbärenbilder zeigten individuell markierte Individuen. Alle 145 gefangenen Waschbären wurden im Rahmen eines separaten Teilprojektes mit hochvariablen Mikrosatelliten erfolgreich genotypisiert, so dass die Verwandtschaftsverhältnisse sowie der individuelle Reproduktionserfolg der Untersuchungstiere bekannt sind. Für die Analysen zur Populationsstruktur wurden unter anderem von 120 verendet aufgefunden Waschbären (Totfunden) aus dem unmittelbaren Umfeld des Nationalparks klassische morphometrische und phänotypische Merkmale sowie die Mortalitätsursachen erfasst. / This study considers questions concerning the spatial and social behaviour, reproduction, population structure and dynamics of the alien North American raccoon (Procyon lotor) in Germany. The investigations took place within the framework of a comprehensive raccoon research project (www.projekt-waschbaer.de) over a period of nearly six years in a close-to-nature lowland beech forest in the Müritz National Park (Mecklenburg-West Pomerania). The National Park has been verifiably colonized by raccoons since the end of the 1970s, and due to its abundance of water and its old deciduous tree population it represents an ideal habitat for this mammal. Between 2006 and 2011, 145 individual raccoons (62 ♀♀ and 83 ♂♂) were captured, genetically sampled, measured and individually tagged. Sampling took place within a 1,114 hectare area of the National Park, at 53 trap sites and with 489 trappings. 51 adult raccoons (23 ♀♀, 28 ♂♂) and 18 juveniles (10 ♀♀, 8 ♂♂) were also fitted with radio collars and located a total of 31,202 times as part of the telemetric survey (=452 localisations per individual). Camera trap monitoring was carried out at 36 baited locations of the main investigation area (1,628 ha): 18,721 camera trap pictures were taken of 183 different raccoons over a monitorring period of 5,365 nights. 82 % of all the raccoon pictures showed individually tagged ani-mals. All 145 of the trapped raccoons were successfully genotyped as part of a subproject with highly polymorphic microsatellites. Both the familial relationships and the individual reproductive success of the subject animals could be determined with the genotyping results. For the analyses of the population structure, classic morphometric and phenotypical characteristics, as well as the cause of mortality of 120 raccoon carcasses in the immediate vicinity of the National Park, were recorded.

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Das Schutzziel in der Ökotoxikologie ist die Population. Un­ter­suchungen zur Wirkung von subletalen Konzentrationen einer Umweltchemikalie auf Populationsebene, zum Beispiel mit künst­lichen Fließgewässersystemen (Mikrokosmen) können aussagekräftigere Beiträge zur ökotoxikologischen Bewertung einer Umweltchemikalie liefern. Außerdem können bei solchen Untersuchungen mögliche indirekte Effekte erfaßt werden. Über die Reaktion von Fließgewässer-Biozönosen gegenüber Um­welt­chemikalien ist relativ wenig bekannt. Die überwiegende Zahl der Untersuchungen zur Abschätzung des Ge­fähr­dungs­po­tentials von Umweltchemikalien wurde mit Testsystemen für Lebensgemeinschaften in stehenden Gewässern untersucht. Die Übertragbarkeit der Ergebnisse dieser Tests auf Fließgewässer-Lebensgemeinschaften ist meist nicht gegeben. Daher ist es notwendig Testsysteme zu etablieren, mit denen die Wirkung von Umweltchemikalien auf Fließgewässer-Le­bens­ge­mein­schaf­ten untersucht werden kann. In einem Gewächshaus wurden fünf Fließrinnen etabliert, mit denen die physikalisch-chemischen Bedingungen in einem Bach simuliert werden können. Im Ge­gensatz zu Untersuchungen einer komplexen Le­bens­ge­mein­schaft mit hoher Variabilität, wie sie sich beispielsweise durch das Einbringen von na­tür­lich­em Sediment aus Fließgewässern einstellt, wurde in diesen Experimenten die Wirkung von Che­mikalien auf eine einfache Lebensgemeinschaft untersucht. Die Le­bens­ge­mein­schaft in den Fließrinnen bestand deshalb aus wenigen, aus­ge­wählten Arten. Untersucht wurden die Kon­zen­tra­tionen 0,6, 6, 60 und 600 µg/l (Terbutryn) und 0,05, 0,5, 5 und 50 µg/l (Fenoxycarb). Gegenstand der vorliegenden Arbeit waren die Untersuchungen mit Gammarus fossarum. In einem akuten Toxizitätstest wurde die LC50 von Terbutryn für adulte und juvenile Gammariden ermittelt. In den Fließ­rin­nen­ex­perimenten mit Terbutryn und Fenoxycarb wurden po­pu­la­tions­relevante Pa­ra­meter der Gammaridenpopulationen untersucht. Ob und in welchem Um­fang sich Effekte, die mit den Stan­dard­tests gemessen wurden, auf bestimmte öko­toxi­ko­lo­gi­sche End­punkte der Population auswirken, kann nicht immer unmittelbar abgeleitet werden. Eine Möglichkeit wäre die aufwendige Durchführung von Po­pu­la­tionsexperimenten mit einfachen oder komplexeren Mo­dell­öko­systemen über eine lange Zeit. Eine andere Möglichkeit ist die Nutzung mathematischer Modelle zur Beschreibung der Populationsdynamik. Das begleitend zu den Untersuchungen entwickelte individuenbasierte Re­pro­duk­tions­modell GamMod bildet die Populationsdynamik einer ab­ge­schlos­senen Population von Gammarus fossarum in künstlichen Fließgewässersystemen ab. Es wird die Struktur und Dy­na­mik des realen Systems (Populationsdynamik) unter Ein­beziehung der Kenntnisse des Reproduktionszyklus modelliert. Mo­dell­szenarien sollen Aus­sagen über den Einfluß der Än­der­ung einer Variablen bezüglich der Populationsdynamik liefern.

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The general aim of my dissertation was to investigate the role of plant interactions in driving population dynamics. Both theoretical and empirical approaches were employed. All my studies were conducted on the basis of metabolic scaling theory (MST), because the complex, spatially and temporally varying structures and dynamics of ecological systems are considered to be largely consequences of biological metabolism. However, MST did not consider the important role of plant interactions and was found to be invalid in some environmental conditions. Integrating the effects of plant interactions and environmental conditions into MST may be essential for reconciling MST with observed variations in nature. Such integration will improve the development of theory, and will help us to understand the relationship between individual level process and system level dynamics. As a first step, I derived a general ontogenetic growth model for plants which is based on energy conservation and physiological processes of individual plant. Taking the mechanistic growth model as basis, I developed three individual-based models (IBMs) to investigate different topics related to plant population dynamics: 1. I investigated the role of different modes of competition in altering the prediction of MST on plant self-thinning trajectories. A spatially-explicit individual-based zone-of-influence (ZOI) model was developed to investigate the hypothesis that MST may be compatible with the observed variation in plant self-thinning trajectories if different modes of competition and different resource availabilities are considered. The simulation results supported my hypothesis that (i) symmetric competition (e.g. belowground competition) will lead to significantly shallower self-thinning trajectories than asymmetric competition as predicted by MST; and (ii) individual-level metabolic processes can predict population-level patterns when surviving plants are barely affected by local competition, which is more likely to be in the case of asymmetric competition. 2. Recent studies implied that not only plant interactions but also the plastic biomass allocation to roots or shoots of plants may affect mass-density relationship. To investigate the relative roles of competition and plastic biomass allocation in altering the mass-density relationship of plant population, a two-layer ZOI model was used which considers allometric biomass allocation to shoots or roots and represents both above- and belowground competition simultaneously via independent ZOIs. In addition, I also performed greenhouse experiment to evaluate the model predictions. Both theoretical model and experiment demonstrated that: plants are able to adjust their biomass allocation in response to environmental factors, and such adaptive behaviours of individual plants, however, can alter the relative importance of above- or belowground competition, thereby affecting plant mass-density relationships at the population level. Invalid predictions of MST are likely to occur where competition occurs belowground (symmetric) rather than aboveground (asymmetric). 3. I introduced the new concept of modes of facilitation, i.e. symmetric versus asymmetric facilitation, and developed an individual-based model to explore how the interplay between different modes of competition and facilitation changes spatial pattern formation in plant populations. The study shows that facilitation by itself can play an important role in promoting plant aggregation independent of other ecological factors (e.g. seed dispersal, recruitment, and environmental heterogeneity). In the last part of my study, I went from population level to community level and explored the possibility of combining MST and unified neutral theory of biodiversity (UNT). The analysis of extensive data confirms that most plant populations examined are nearly neutral in the sense of demographic trade-offs, which can mostly be explained by a simple allometric scaling rule based on MST. This demographic equivalence regarding birth-death trade-offs between different species and functional groups is consistent with the assumptions of neutral theory but allows functional differences between species. My initial study reconciles the debate about whether niche or neutral mechanisms structure natural communities: the real question should be when and why one of these factors dominates. A synthesis of existing theories will strengthen future ecology in theory and application. All the studies presented in my dissertation showed that the approaches of individual-based and pattern-oriented modelling are promising to achieve the synthesis.

Folgende Themenkomplexe wurden behandelt: Biogeographie, Pflanzendiversität, Verbreitungsmuster von Lebensformen und Arten in Mikrohabitaten sowie entlang ökologischer Gradienten, Epiphyt-Porophyt-Beziehungen, Vergesellschaftung, Sekundärsukzession und die Auswahl von Indikatorarten. Zum ersten Mal wird bei einer derartigen Untersuchung die sekundäre Sukzession von Kryptogamengesellschaften in tropischen Bergregenwäldern voll berücksichtigt. Die Untersuchung wurde im Einzugsgebiet des Río Savegre auf der pazifikseitigen Abdachung der westlichen Cordillera de Talamanca (Costa Rica) durchgeführt.Kapitel 1 gibt eine generelle Einführung in Tropische Bergregenwälder und die Bedeutung von Moosen und Makroflechten als wichtige Komponenten dieser Ökosysteme. Der Leser wird in das Untersuchungsgebiet, sein Klima, seine Geologie und seine Vegetation eingeführt.Im Kapitel 2 werden die pflanzengeographischen Muster der Moose montaner Eichenwälder der Cordillera de Talamanca dargestellt und diskutiert.Kapitel 3 diskutiert Diversität, Mikrohabitatdifferenzierung und Lebensformenspektren der Moose in einem hochmontanen Eichenwald Costa Ricas. Es wurde versucht, die Moosflora aller Mikrohabitate eines Eichenwaldes, inklusive der Strauch- und Bodenschicht zu erfassen. Kapitel 4 und 5 beinhalten die Beschreibung epiphytischer Kryptogamengesellschaften (Moose und Makroflechten) in Primär- und Sekundärwäldern. Mit Hilfe einer aus dem Alpinismus abgeleiteten Klettertechnik wurden Bäume von der Stammbasis bis in die Zweige der äußeren Krone beprobt. Die Vegetationsaufnahmen repräsentieren die Variation der epiphytischen Vegetation. Prozentuale Deckungsgrade der rindenbewohnenden Moose und Makroflechten wurden geschätzt und mit Hilfe einer Detrended Correspondence Analysis (DCA)' und nicht-metrischer Multi-response Permutation Procedure (MRPP)' verglichen. Rindenproben der untersuchten Bäume wurden mit deionisiertem Wasser extrahiert und der pH-Wert bestimmt.Kapitel 4 behandelt die epiphytischen Kryptogamengemeinschaften auf den dominierenden Baumarten (Quercus copeyensis und Q. costaricensis) primärer, hochmontaner Eichenwälder Costa Ricas. Kapitel 5 vergleicht Artenreichtum, Vergesellschaftung und Ökologie der epiphytischen Kryptogamen in primären und sekundären hochmontanen Eichenwäldern Costa Ricas. Quercus copeyensis-Bäume der oberen Baumschicht von Primär- und Sekundärwäldern wurden beprobt mit dem Ziel einen Einblick in Regenerationspotential, -muster und -prozesse der Epiphytensukzession nach anthropogener Störung zu erhalten.

It appears to be unanimous that alien species in island environments tend to cause considerably more negative than positive impacts. To assess the potential level of threat aliens may pose to the native environment, understanding a species’ population structure and dynamics is of ultimate importance. Assessing both impacts and consequences of management interventions to alien species is likewise only possible through the comprehension of its population structure and dynamics. This can be achieved by estimating the number of individuals in the study site, as well as other population parameters through time, applying population models such as capture-recapture to the collected datasets. Nonetheless, alien species that have low capture rates, such as small mammals, might present a considerable obstacle for conservation, as available capturerecapture models need a relatively large dataset to precisely and accurately estimate population parameters. To improve accuracy and precision of estimates that use sparse datasets, the present study developed an integrated concurrent marking-observation capture-recapture model (C-MOM). The model proposed here, contrary to the commonly available mark-recapture and mark-resight models, allows for two different datasets (i.e. a capture-recapture and a population count) to be integrated, as well as for marking and observation (recapture) data to be collected simultaneously. While few models can integrate different datasets, no model is known to allow for concomitant capture-markobservation activities. To assess the performance of the C-MOM when estimating population parameters for sparse datasets, a virtual ecology study was carried out. The population dynamics of a small rodent, the rock cavy (Kerodon rupestris), as well as capture-recapture and population count datasets, were simulated under different scenarios. The sampled datasets were then analyzed by the C-MOM, and by two other established statistical models: a classical mark recapture (CMR) (based on the Jolly-Seber model), and a zero-truncated Poisson log-normal mixed effects (ZPNE), the only integrated mark-resight model that allows for recapture sampling with replacement. Estimates of population parameters provided by the three models were then compared in terms of bias, precision and accuracy. C-MOM and ZPNE models were afterwards applied to real data collected on a rock cavy colony in the island of Fernando de Noronha. The estimated parameters were used to extrapolate the number of individuals in the rock cavy colony to the whole population in the island. Subsequently, these results were used to develop a risk assessment for the species by modelling historical and management scenarios, simulating both the establishment of the species in the island, and the consequences of different management interventions applied to it. The virtual ecology study showed that, in comparison to the CMR and the ZPNE, the C-MOM presented improved accuracy without overestimating the precision of population parameter’s estimates. The last also presented reduced amplitude of the calculated credible interval at 95% when applied to real data in comparison to the ZPNE. While the extrapolation of C-MOM estimates suggests that the rock cavy population in Fernando de Noronha is 6,652 ± 1,587, ZPNE estimates are of 5,854 ± 3,269 individuals. In the risk assessment, historical simulation models demonstrated that even though different combinations of uncertainty in reproductive parameters of the rock cavy might be possible for the species, these did not interfere significantly in either establishment or spread of the rock cavy population in the island. Moreover, historical yearly mortality has most likely been under 30%. Regarding the species’ management simulations, the most effective management interventions to achieve population extinction were spaying and neutering of both sexes, although harvest effort presented the highest influence on this populations’ extirpation. Nonetheless, the relative influence of female and both sexes’ based interventions did not differ significantly regarding the frequency of extinction of stochastic replicates’. Moreover, none of the management interventions guaranteed the population extinction within the time span and harvest effort proposed for the management program. Neutering of both sexes was most inversely influential on time to extinction of this population, followed by removal of both sexes. Briefly, the C-MOM has proven to be a resourceful and precise model to estimate population parameters when low capture rates result in sparse datasets. Moreover, the rock cavy is well established in the island and likely at carrying capacity. In general, the risk assessment showed that the management interventions in the time span and harvest effort simulated in the present study were ineffective to extinguish the rock cavy population in Fernando de Noronha. Considering this, as well as the importance of investigating other vital factors to decide in favour of or contrary to the management of this species, it is recommended that both an impact assessment of the rock cavy and a cost-effectiveness analysis of the management interventions should be performed to complement the current study.:Acknowledgement III Abstract IV Zusammenfassung VI Resumen IX Table of Contents XII List of Tables and Figures XIV List of Abbreviations XIX 1. Introduction 1 1.1. Invasive alien species and their consequences 1 1.2. Population dynamics analysis 2 Capture-recapture models 3 Observation models 4 Integrated population models 5 Software 7 Model analysis 8 1.3. Fernando de Noronha and the rock cavy 10 1.4. Objectives 12 Overall Objectives 12 Specific Objectives 13 2. Study Framework 15 3. Methods 19 3.1. Study area 19 3.2. Study case species 21 3.3. Research Steps 24 RESEARCH STEP I: Comparing the C-MOM to established models – does this concurrent marking-observation model produces accurate estimates of population parameters for sparse datasets? 24 RESEARCH STEP II: C-MOM application to a real case study 40 RESEARCH STEP III: The rock cavy population in Fernando de Noronha 45 RESEARCH STEP IV: The colonization and eradication of the rock cavy in Fernando de Noronha 47 4. Results 63 4.1. RESEARCH STEP I: Comparing the C-MOM to established models – does this concurrent marking-observation model produces accurate estimates of population parameters for sparse datasets? 63 4.2. RESEARCH STEP II: C-MOM application to a real case study 72 4.3. RESEARCH STEP III: The rock cavy population in Fernando de Noronha 73 4.4. RESEARCH STEP IV: The colonization and eradication of the rock cavy in Fernando de Noronha 74 Sensitivity analysis 74 Simulation experiments 80 5. Discussion 83 5.1. Bias, precision and accuracy of population dynamic models for sparse datasets 85 Simulated data 85 Study case 90 5.2. Advantages and disadvantages of the C-MOM approach 93 5.3. Development and applications of the integrated models and the C-MOM 96 5.4. The reversed use of the PVA software Vortex to simulate AS and IAS populations’ extinction 97 5.5. Status of the rock cavy population in the island of Fernando de Noronha 100 The colonization of the rock cavy in Fernando de Noronha 101 Management of the rock cavy in Fernando de Noronha 104 Study case limitations and future researches 112 6. Conclusion 116 References 118 Appendices 124 APPENDIX I – Assessment of biological invasions 124 APPENDIX II – Population dynamics simulation and dataset sampling 125 APPENDIX III – CMR and C-MOM model codes in R 134 APPENDIX IV – ZPNE model code in R 138 APPENDIX V – C-MOM model used for real datasets 143 APPENDIX VI – Rock cavy colony sizes and number of individuals in Fernando de Noronha 145 APPENDIX VII – Parameter’s ranking of C-MOM, CMR and ZPNE models 148 APPENDIX VIII – Bias, precision and accuracy table 149