Spatial and temporal heterogeneity of water and nutrient pools is closely associated with the existence of different plant communities in hydrologically-controlled ecosystems such as the Everglades. These various communities differ in their flooding and fire tolerances as well as nutrient requirements. Upland plant communities are of particular ecological significance since they have high biodiversity and provide habitat to terrestrial fauna, yet comprise less than 10 % of the total area. Restoration and maintenance of such communities requires an understanding of their water and nutrient requirements. Chapter 2 compares water source utilization in hammocks and pine rocklands on the Miami Rock Ridge using stable isotopes of water. Hammocks do not flood, while adjacent pinelands may flood between 2-3 months. In the wet season, hammocks were found to use phosphorus (P) rich soilwater, a local pool of water and nutrients while pineland plants primarily relied upon groundwater, the regional pool. Access to a rich pool of P in the oligotrophic Everglades was associated with higher community-level foliar P concentration in hammocks. However in the dry season, hammocks utilized groundwater, which suggests sensitivity to extended droughts. Chapter 3 compares the hammock (upland or head) and swamp forests (lowland or tail) on tree islands in the Shark River Slough. Uplands were associated with P-rich soilwater uptake in the wet season, with regional water uptake in the dry season. Accordingly, tree island heads are rich in foliar P and thereby P-hotspots in the Everglades. Foliar nutrient concentrations can thus indicate limiting nutrient availability in the Everglades. Chapter 4 looks at how leaf phenology patterns are tied to water and nutrient pools. Leaf fall in ridge hammocks is associated with high foliar carbon isotope values over the dry season, which is not the case for tree island hammocks. However, in some species, high levels of foliar nitrogen are also associated with high foliar C13 values indicating stomatal limitation of photosynthesis. Growing season for most hammock species is the wet season coinciding with high availability of P, as reflected in high foliar P in this season. Linking water sources to foliar nutrients elucidates roles of water and nutrient pools in leading to different plant communities within an ecosystem.

Lake ecosystems across the globe have responded to climate warming of recent decades. However, correctly attributing observed changes to altered climatic conditions is complicated by multiple anthropogenic influences on lakes. This thesis contributes to a better understanding of climate impacts on freshwater phytoplankton, which forms the basis of the food chain and decisively influences water quality. The analyses were, for the most part, based on a long-term data set of physical, chemical and biological variables of a shallow, polymictic lake in north-eastern Germany (Müggelsee), which was subject to a simultaneous change in climate and trophic state during the past three decades. Data analysis included constructing a dynamic simulation model, implementing a genetic algorithm to parameterize models, and applying statistical techniques of classification tree and time-series analysis. Model results indicated that climatic factors and trophic state interactively determine the timing of the phytoplankton spring bloom (phenology) in shallow lakes. Under equally mild spring conditions, the phytoplankton spring bloom collapsed earlier under high than under low nutrient availability, due to a switch from a bottom-up driven to a top-down driven collapse. A novel approach to model phenology proved useful to assess the timings of population peaks in an artificially forced zooplankton-phytoplankton system. Mimicking climate warming by lengthening the growing period advanced algal blooms and consequently also peaks in zooplankton abundance. Investigating the reasons for the contrasting development of cyanobacteria during two recent summer heat wave events revealed that anomalously hot weather did not always, as often hypothesized, promote cyanobacteria in the nutrient-rich lake studied. The seasonal timing and duration of heat waves determined whether critical thresholds of thermal stratification, decisive for cyanobacterial bloom formation, were crossed. In addition, the temporal patterns of heat wave events influenced the summer abundance of some zooplankton species, which as predators may serve as a buffer by suppressing phytoplankton bloom formation. This thesis adds to the growing body of evidence that lake ecosystems have strongly responded to climatic changes of recent decades. It reaches beyond many previous studies of climate impacts on lakes by focusing on underlying mechanisms and explicitly considering multiple environmental changes. Key findings show that climate impacts are more severe in nutrient-rich than in nutrient-poor lakes. Hence, to develop lake management plans for the future, limnologists need to seek a comprehensive, mechanistic understanding of overlapping effects of the multi-faceted human footprint on aquatic ecosystems. / Weltweit haben Seeökosysteme auf den Klimawandel der letzten Jahrzehnte reagiert. Beobachtete Veränderungen eindeutig dem Klimawandel zuzuordnen, wird jedoch häufig dadurch erschwert, dass Seen gleichzeitig vielfachen anthropogenen Einflüssen ausgesetzt sind. Diese Arbeit trägt zu einem besseren Verständnis des Klimaeinflusses auf Algen bei, die am Anfang der Nahrungskette stehen und maßgeblich die Wasserqualität eines Sees beeinflussen können. Zum größten Teil stützt sich die Arbeit auf eine dreißigjährige Datenreihe eines unregelmäßig geschichteten Flachsees im Nordosten von Deutschland (Müggelsee), in dem sowohl steigende Wassertemperaturen als auch sinkende Nährstoffeinträge zu verzeichnen waren. Bei der Datenanalyse wurde ein neu erstelltes dynamisches Simulationsmodell, genetische Algorithmen zur Parametrisierung von Modellen, und statistische Methoden der Klassifizierung und Zeitreihenanalyse genutzt. Ergebnisse dieser Arbeit zeigen, dass nicht nur klimatische Faktoren sondern auch die Nährstoffverfügbarkeit im See den Zeitpunkt der Algenfrühjahrsblüte (Phänologie) beeinflussen. Durch eine Veränderung der Mechanismen, die zum Kollaps der Blüte führen, trat diese trotz ähnlich milder Winterbedingungen bei hoher Nährstoffverfügbarkeit früher auf als bei niedriger. Ein neuentwickelter Ansatz zur Modellierung von Phänologie erwies sich als geeignet, um vorherzusagen, wann Algen und ihre Räuber in einem künstlich periodisch angetriebenen Laborsystem ihre Populationshöhepunkte erreichten. Eine Verlängerung der Wachstumsperiode führte dazu, dass diese früher auftraten. Die Untersuchung, warum sich Blaualgen im betrachteten See während jüngster Hitzewellenereignisse überraschend unterschiedlich entwickelt hatten, ergab, dass ungewöhnlich warmes Wetter nicht wie häufig vermutet generell förderlich für ihre Entwicklung ist. Der Zeitpunkt und die Dauer der Hitzewellen waren entscheidend dafür, ob für Blaualgen kritische Schwellenwerte der thermischen Schichtung im See überschritten wurden. Zudem zeigte sich, dass saisonale Erwärmungsmuster einen bedeutenden Einfluss auf Räuber nahmen, die das Auftreten von Algenblüten verhindern können. Diese Arbeit reiht sich in eine wachsende Anzahl von Studien ein, die zeigen, dass Seeökosysteme bereits stark auf die Klimaveränderungen der letzen Jahrzehnte reagiert haben. Mit ihrem Fokus auf Mechanismen und der expliziten Berücksichtigung simultaner anthropogener Einflüsse geht diese Arbeit gleichzeitig über viele bisherige Studien hinaus, die sich auf reine Beobachtung und die Betrachtung klimatischer Faktoren beschränkten. Kernergebnisse deuten daraufhin, dass Klimafolgen in nährstoffreichen Seen stärker ausfallen als in nährstoffarmen Seen. Nur mit einem umfassenden, mechanistischen Verständnis des vielfältigen anthropogenen Einflusses wird eine hohe Wasserqualität in Seen auch in Zukunft aufrechtzuerhalten sein.

<p>Recent changes in the timing of annual events are a sign that climate change is already impacting ecosystems. Carbon sequestration by forests increases with longer growing seasons. Biodiversity can be affected by mis-timing of events through shading interactions and frost damage. Projecting forests' ability to provide these ecosystem services in the future requires an understanding of trees' phenological responses to a new climate. I begin by proposing a first order definition of an `optimal' phenological response to warming: that the mean temperature following budburst should remain essentially constant. Analogously, the temperature preceding senescence can serve the same role. </p><p>To understand which environmental cues will drive future changes in phenology, I assimilate clues from observational and experimental literature. For budburst in woody plants, spring warmth, over-winter chilling and light drive nearly all behavior, but species' responses vary widely. Species using chilling or light as safety mechanisms against budburst during mid-winter thaws are thought to be less able to phenologically track a warming climate. However, I show that even species cued solely by spring warmth are likely to under-track temperature changes. Fall cues are more idiosyncratic, and a plant's driver of senescence is likely to vary from year to year. </p><p>Models are a tempting method to untangle species budburst cues and forecast phenology under warmer climate scenarios. I tested two models' ability to recover parameters used to simulate budburst data. The simpler model was cued only by spring warmth while the complex one modulated warmth requirements with chilling exposure. For the simple model, parameters could be recovered consistently from some, but not all, regions of parameter space. The complex model's parameters were largely unrecoverable. To understand the consequences of parameter uncertainty, I applied both models to an 18 year phenological record of 13 deciduous tree species. While a few species fell into identifiable regions of the simple model's parameter space, most did not, and projected budburst dates had wide parameter-derived uncertainty intervals. These bands were wider still under a 5°C warming scenario. Even greater uncertainty resulted from the complex model.</p><p>To better understand plants' potential for growing season extension I subjected seedlings to warmer climates in a series of open-topped chambers in sites at each end of the eastern deciduous biome. Soil and air were heated to 3 or 5°C above ambient, or left unheated. For nearly all species, warming hastened budburst and germination and delayed senescence. However, these events failed to track temperature changes, happening at warmer temperatures in hotter chambers. Individual species showed a remarkable variability of all events' dates within treatments, and even within chambers. Because phenological traits are heritable, this offers a potential for evolutionary response to climate change.</p><p>This research has shown that while individual trees extend their growing seasons under warmer temperatures, they typically under-respond to the magnitude of warming, suggesting forests' capacity for increased carbon sequestration may reach a limit. However, within populations, trees vary substantially in their phenological responses, forming a possibility for evolutionarily adaptation to changing cues.</p> / Dissertation

Global and local climate patterns may affect barren-ground caribou (Rangifer tarandus groenlandicus) populations. I predicted global climate changes to be correlated with periods of population decline, and local changes to be more pronounced on the habitat of caribou with a declining population. In chapter 1, the Arctic Oscillation (AO), changes in normalized difference vegetation index and phenology were used as measures of global and local climate. In chapter 2 environmental variables and caribou presence points were used to build Maxent habitat models. There was no consistent correlation with the positive AO phase and periods of population decline, or phenology trends and the habitat of caribou with a declining population. Maxent models underestimated the amount of suitable habitat spatially and failed to model suitable habitat temporally. This thesis is the first to look at a range of density-independent variables over a long time period and model suitable habitat for multiple herds.

The projected increase of shrubs across the Arctic is expected to alter patterns of snow cover, which may affect the phenology and survival of arthropods such as spiders. In this study, we simulated shrub encroachment on a series of tundra plots and examined the effects on the spider assemblages during the following growing season. Our simulated shrub treatment did not affect the abundance or composition of spider communities over the season; however, adults from the dominant genus Pardosa (Lycosidae) had significantly higher body mass on treatment plots. This difference in mass was observed following snow melt and persisted until halfway through the growing season. Given the importance of spiders as arthropod predators and as food sources for breeding birds, such a change in summer body mass could represent a shift in spiders’ functional contributions to Arctic ecosystems.

The projected increase of shrubs across the Arctic is expected to alter patterns of snow cover, which may affect the phenology and survival of arthropods such as spiders. In this study, we simulated shrub encroachment on a series of tundra plots and examined the effects on the spider assemblages during the following growing season. Our simulated shrub treatment did not affect the abundance or composition of spider communities over the season; however, adults from the dominant genus Pardosa (Lycosidae) had significantly higher body mass on treatment plots. This difference in mass was observed following snow melt and persisted until halfway through the growing season. Given the importance of spiders as arthropod predators and as food sources for breeding birds, such a change in summer body mass could represent a shift in spiders’ functional contributions to Arctic ecosystems.

Warming induced changes in Arctic vegetation have to date been studied through observational and experimental field studies, leaving significant uncertainty about the representativeness of selected field sites as well as how these field scale findings scale up to the entire pan-Arctic. The purposes of this thesis were therefore to 1) analyze remotely-sensed/modeled temperature, Normalized Difference Vegeta- tion Indices (NDVI) and plant Net Primary Productivity (NPP) to assess coarse- scale changes (1982–2006) in vegetation; and 2) compare field, remote sensing and model outputs to estimate limitations, challenges and disagreements between data formats. The following data sources were used: • Advanced Very High Resolution Radiometer Polar Pathfinder Extended (APP- x, temperature & albedo) • Moderate Resolution Imaging Spectroradiometer (MODIS, Normalized Dif- ference Vegetation Index (NDVI) & Enhanced Vegetation Index (EVI) ) • Landsat Enhanced Thematic Mapper (Landsat ETM, NDVI) • Global Inventory Modeling and Mapping Studies (GIMMS, NDVI) • Global Productivity Efficiency Model (GloPEM, Net Primary Productivity (NPP)) Over the pan-Arctic (1982-2007), increases in temperature, total annual NPP and maximum annual NDVI were observed. Increases in NDVI and NPP were found to be closely related to increases in temperature according to non-parametric Sen’ slope and Mann Kendall tau tests. Variations in phenology were largely non- significant but related to increases in growing season temperature. Snow melt onset and spring onset correspond closely. MODIS, Landsat and GIMMS NDVI data sets agree well, and MODIS EVI and NDVI are very similar for spring and summer at Fosheim Peninsula. GloPEM NPP and field estimates of NPP are poorly correlated, whereas GIMMS NDVI and GloPEM NPP are well correlated, indicating a need for better calibration of model NPP to field data. In summary, increases in pan-Arctic biological productivity indicators were ob- served, and were found to be closely related to recent circumpolar warming. How- ever, these changes appear to be focused in regions from which recent field studies have found significant ecological changes (Alaska), and coarse resolution remote sensing estimates of ecological changes have been less marked in other regions. Dis- crepancies between results from model, field data and remote sensing, as well as central questions remaining about the impact of increases in productivity on soil- vegetation-atmosphere feedbacks, indicate a clear need for continued research into warming induced changes in pan-Arctic vegetation.

Warming induced changes in Arctic vegetation have to date been studied through observational and experimental field studies, leaving significant uncertainty about the representativeness of selected field sites as well as how these field scale findings scale up to the entire pan-Arctic. The purposes of this thesis were therefore to 1) analyze remotely-sensed/modeled temperature, Normalized Difference Vegeta- tion Indices (NDVI) and plant Net Primary Productivity (NPP) to assess coarse- scale changes (1982–2006) in vegetation; and 2) compare field, remote sensing and model outputs to estimate limitations, challenges and disagreements between data formats. The following data sources were used: • Advanced Very High Resolution Radiometer Polar Pathfinder Extended (APP- x, temperature & albedo) • Moderate Resolution Imaging Spectroradiometer (MODIS, Normalized Dif- ference Vegetation Index (NDVI) & Enhanced Vegetation Index (EVI) ) • Landsat Enhanced Thematic Mapper (Landsat ETM, NDVI) • Global Inventory Modeling and Mapping Studies (GIMMS, NDVI) • Global Productivity Efficiency Model (GloPEM, Net Primary Productivity (NPP)) Over the pan-Arctic (1982-2007), increases in temperature, total annual NPP and maximum annual NDVI were observed. Increases in NDVI and NPP were found to be closely related to increases in temperature according to non-parametric Sen’ slope and Mann Kendall tau tests. Variations in phenology were largely non- significant but related to increases in growing season temperature. Snow melt onset and spring onset correspond closely. MODIS, Landsat and GIMMS NDVI data sets agree well, and MODIS EVI and NDVI are very similar for spring and summer at Fosheim Peninsula. GloPEM NPP and field estimates of NPP are poorly correlated, whereas GIMMS NDVI and GloPEM NPP are well correlated, indicating a need for better calibration of model NPP to field data. In summary, increases in pan-Arctic biological productivity indicators were ob- served, and were found to be closely related to recent circumpolar warming. How- ever, these changes appear to be focused in regions from which recent field studies have found significant ecological changes (Alaska), and coarse resolution remote sensing estimates of ecological changes have been less marked in other regions. Dis- crepancies between results from model, field data and remote sensing, as well as central questions remaining about the impact of increases in productivity on soil- vegetation-atmosphere feedbacks, indicate a clear need for continued research into warming induced changes in pan-Arctic vegetation.

<p>Under the increasing threat of climate change, it is imperative to understand the impact that environmental phenomena have on the demography and behavior of natural populations. In the last few decades an ever increasing body of research has documented dramatic changes in mortality rates and breeding phenology for a large number of species. A number of these have been attributed to the current trends in climate change, which have been particularly conspicuous in bird populations. However, datasets associated to these natural populations as well as to the environmental variables that affect their biology tend to be partial and incomplete. Thus, ecological research faces the urgent need to tackle these questions while at the same time develop inferential models that can handle the complex structure of these datasets and their associated uncertainty. Therefore, my dissertation research has focused on two main objectives: 1) to understand the relationship that demographic rates and breeding phenology of a colony of seabirds has with the environment in the context of climate change; and 2) to use and develop models that can encompass the complex structure of these natural systems, while also extending the process not only to inference but to building predictions. I divided this work in three research projects; for the first one I developed a hierarchical Bayesian model for age-specific survival for long lived species with capture-recapture data that allows the use of incomplete data (i.e. left-truncated and right-censored), and builds predictions of years of birth and death for all individuals while also drawing inference on the survivorship function. I compared this method to more traditional ones and address their limitations and advantages. My second research chapter makes use of this method to determine the age-specific survivorship of the Dry Tortugas sooty tern population, and explores the effect of changes in sea surface temperature on their cohort mortality rates. Finally, my third research chapter addresses the dramatic shift in breeding season experienced by the Dry Tortugas sooty tern colony, the most unprecedented shift reported for any bird species. I explore the role of climatic and weather variables as triggering mechanisms.</p> / Dissertation

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