Mountains are vital to ecosystems and human society given their influence on global carbon and water cycles. Yet the extent to which topography regulates montane forest carbon uptake and storage remains poorly understood. To address this knowledge gap, we compared forest aboveground carbon loading to topographic metrics describing energy balance and water availability across three headwater catchments of the Boulder Creek Watershed, Colorado, USA. The catchments range from 1800 to 3500 m above mean sea level with 46-102 cm/yr mean annual precipitation and -1.2 degrees to 12.3 degrees C mean annual temperature. In all three catchments, we found mean forest carbon loading consistently increased from ridges (27 +/- 19 Mg C ha) to valley bottoms (60 +/- 28 Mg C ha). Low topographic positions held up to 185 +/- 76 Mg C ha, more than twice the peak value of upper positions. Toe slopes fostered disproportionately high net carbon uptake relative to other topographic positions. Carbon storage was on average 20-40 Mg C ha greater on north to northeast aspects than on south to southwest aspects, a pattern most pronounced in the highest elevation, coldest and wettest catchment. Both the peak and mean aboveground carbon storage of the three catchments, crossing an 11 degrees C range in temperature and doubling of local precipitation, defied the expectation of an optimal elevation-gradient climatic zone for net primary production. These results have important implications for models of forest sensitivity to climate change, as well as to predicted estimates of continental carbon reservoirs.

Current estimates of terrestrial carbon (C) fluxes overlook explicit hydrological controls. In this research project, a spatially explicit hydro-ecological model, BEPS-TerrainLab V2.0 was further developed to improve our understanding of the non-linearities associated with various hydro-ecological processes. A modeling study was conducted in a humid boreal ecosystem in north central Quebec, Canada. The sizes and nature of various ecosystem-C-pools were comprehensively reconstructed under a climate change and disturbance scenario prior to simulation in order to ensure realistic biogeochemical modeling. Further, several ecosystem processes were simulated and validated using field measurements for two years. A sensitivity analysis was also performed. After gaining confidence in the model’s ability to simulate various hydrologically controlled ecophysiological and biogeochemical processes and having understood that topographically driven sub-surface baseflow is the main process determining the soil moisture regime in humid boreal ecosystem, its influence on ecophysiological and biogeochemical processes were investigated. Three modeling scenarios were designed that represent strategies that are currently used in ecological models to represent hydrological controls. These scenarios were: 1) Explicit, where realistic lateral water routing was considered 2) Implicit, where calculations were based on a bucket-modeling approach 3) NoFlow, where the lateral sub-surface flow was turned off in the model. In general, the Implicit scenario overestimated GPP, ET and NEP, as opposed to the Explicit scenario. The NoFlow scenario underestimated GPP and ET but overestimated NEP. The key processes controlling the differences were due to the combined effects of variations in plant physiology, photosynthesis, heterotrophic respiration, autotrophic respiration and nitrogen mineralization; all of which occurred simultaneously in different directions, at different rates, affecting the spatio-temporal distribution of terrestrial C-sources or sinks (NEP). From these results it was clear that lateral water flow does play a significant role in the net terrestrial C distribution and it was discovered that non-explicit forms of hydrological representations underestimate the sizes of terrestrial C-sources rather than C-sinks. The scientific implication of this work demonstrates that regional or global scale terrestrial C estimates could have significant errors if proper hydrological constraints are not considered for modeling ecological processes due to large topographic variations of the Earth’s surface.

<p>Topography, vegetation, nutrient dynamics, soil features and hydroclimatic forcing are inherently coupled, with feedbacks occurring over a wide range of temporal and spatial scales. Vegetation growth may be limited by soil moisture, nutrient or solar radiation availability, and in turn influences both soil moisture and nutrient balances at a point. These dynamics are further complicated in a complex terrain, through a series of spatial interactions. A number of experiments has characterized the feedbacks between soil moisture and vegetation dynamics, but a theoretical framework linking short-term leaf-level to interannual plot-scale dynamics has not been fully developed yet. Such theory is needed for optimal management of water resources in natural ecosystems and for agricultural, municipal and industrial uses. Also, it complements the current knowledge on ecosystem response to the predicted climate change.</p><p>In this dissertation, the response of vegetation dynamics to unpredictable environmental fluctuations at multiple space-time scales is explored in a modeling framework from sub-daily to interannual time scales. At the hourly time scale, a simultaneous analysis of photosynthesis, transpiration and soil moisture dynamics is carried out to explore the impact of water stress on different photosynthesis processes at the leaf level, and the overall plant activity. Daily soil moisture and vegetation dynamics are then scaled up to the growing season using a stochastic model accounting for daily to interannual hydroclimatic variability. Such stochastic framework is employed to explore the impact of rainfall patterns and different irrigation schemes on crop productivity, along with their implications in terms of sustainability and profitability. To scale up from point to landscape, a probabilistic representation of local landscape features (i.e., slope and aspect) is developed, and applied to assess the effects of topography on solar radiation. Finally, a minimalistic ecosystem model, including soil moisture, vegetation and nutrient dynamics at the year time scale, is outlined; when coupled to the proposed probabilistic topographic description, the latter model can serve to assess the relevance of spatial interactions and to single out the main biophysical controls responsible for ecohydrological variability at the landscape scale.</p> / Dissertation

Current estimates of terrestrial carbon (C) fluxes overlook explicit hydrological controls. In this research project, a spatially explicit hydro-ecological model, BEPS-TerrainLab V2.0 was further developed to improve our understanding of the non-linearities associated with various hydro-ecological processes. A modeling study was conducted in a humid boreal ecosystem in north central Quebec, Canada. The sizes and nature of various ecosystem-C-pools were comprehensively reconstructed under a climate change and disturbance scenario prior to simulation in order to ensure realistic biogeochemical modeling. Further, several ecosystem processes were simulated and validated using field measurements for two years. A sensitivity analysis was also performed. After gaining confidence in the model’s ability to simulate various hydrologically controlled ecophysiological and biogeochemical processes and having understood that topographically driven sub-surface baseflow is the main process determining the soil moisture regime in humid boreal ecosystem, its influence on ecophysiological and biogeochemical processes were investigated. Three modeling scenarios were designed that represent strategies that are currently used in ecological models to represent hydrological controls. These scenarios were: 1) Explicit, where realistic lateral water routing was considered 2) Implicit, where calculations were based on a bucket-modeling approach 3) NoFlow, where the lateral sub-surface flow was turned off in the model. In general, the Implicit scenario overestimated GPP, ET and NEP, as opposed to the Explicit scenario. The NoFlow scenario underestimated GPP and ET but overestimated NEP. The key processes controlling the differences were due to the combined effects of variations in plant physiology, photosynthesis, heterotrophic respiration, autotrophic respiration and nitrogen mineralization; all of which occurred simultaneously in different directions, at different rates, affecting the spatio-temporal distribution of terrestrial C-sources or sinks (NEP). From these results it was clear that lateral water flow does play a significant role in the net terrestrial C distribution and it was discovered that non-explicit forms of hydrological representations underestimate the sizes of terrestrial C-sources rather than C-sinks. The scientific implication of this work demonstrates that regional or global scale terrestrial C estimates could have significant errors if proper hydrological constraints are not considered for modeling ecological processes due to large topographic variations of the Earth’s surface.

Swan Lake watershed, a sub-catchment of the salmonid bearing Colquitz Creek watershed located in the municipality of Saanich, on southern Vancouver Island, British Columbia, Canada, was studied to characterise the linkages between urbanisation and ecological health and resilience. Although rarely applied in watershed ecology, resilience (the ability to absorb disturbances without the loss of ecosystem identity) offers a useful construct in this case study to understand the effects of urban development over the past 150 years, and to outline some principles for integrated, watershed-scale management. Baseline landscape characteristics and processes of historical land-use were determined using paleoecology (pollen analysis) and historical records. Watershed health was assessed using: a Proper Functioning Condition assessment of riparian-wetland and stream channels; vegetation community mapping; vegetation plots; surface flow hydrology; and water quality analyses. Vegetation and lake hydrographs were compared with less disturbed reference ecosystems. Findings are discussed in terms of alternative stable state models and energy dissipation at the site and landscape scale. Analysis of the data revealed that over the past 150 years, forest clearing, agriculture, transportation infrastructure, and non-point source pollution have transformed the landscape and substantially altered the water and energy balance. Impervious surfaces and cleared land (covering 25% and 35% of the watershed, respectively) are inferred to have reduced latent heat dissipation of solar energy, an important landscape-scale process affecting resilience to climate change. Degraded stream channels represent reduced ecosystem services and lost social/economic value. The stream/lake hydrographs revealed a typical, urban flashy profile that exacerbates channel erosion and non-point source pollution, while excessive lake stage drawdown is also evident. Water quality is characterized by historic and ongoing excessive nutrient loading and associated cultural eutrophication, heavy metal pollution, and ecosystem “ageing” due to dissolved solids runoff. At the site level, invasive species, particularly reed canarygrass, dominate Swan Lake wetlands, whereas the pollen record shows abundant woody shrubs and associated species (some of which are now extirpated from the site) and an absence of grass; this helps to establish a rationale for vegetation management. Based on the findings of the above studies and according to a proposed conceptual model with assessment criteria in five categories (water, vegetation, energy, soil and nutrients), Swan Lake watershed has impaired ecological health and is not resilient to disturbances such as extreme climate/weather events. Future watershed management should therefore mimic the hydrological function and energy balance of the pre-development conditions.

Soils contain a large amount of carbon (C) that is a critical regulator of the global C budget. Already small changes in the processes governing soil C cycling have the potential to release considerable amounts of CO2, a greenhouse gas (GHG), adding additional radiative forcing to the atmosphere and hence to changing climate. Increased temperatures will probably create a feedback, causing soils to release more GHGs. Furthermore changes in soil C balance impact soil fertility and soil quality, potentially degrading soils and reducing soils function as important resource. Consequently the assessment of soil C dynamics under present, recent past and future environmental conditions is not only of scientific interest and requires an integrated consideration of main factors and processes governing soil C dynamics. To perform this assessment an eco-hydrological modelling tool was used and extended by a process-based description of coupled soil carbon and nitrogen turnover. The extended model aims at delivering sound information on soil C storage changes beside changes in water quality, quantity and vegetation growth under global change impacts in meso- to macro-scale river basins, exemplary demonstrated for a Central European river basin (the Elbe). As a result this study: ▪ Provides information on joint effects of land-use (land cover and land management) and climate changes on croplands soil C balance in the Elbe river basin (Central Europe) presently and in the future. ▪ Evaluates which processes, and at what level of process detail, have to be considered to perform an integrated simulation of soil C dynamics at the meso- to macro-scale and demonstrates the model’s capability to simulate these processes compared to observations. ▪ Proposes a process description relating soil C pools and turnover properties to readily measurable quantities. This reduces the number of model parameters, enhances the comparability of model results to observations, and delivers same performance simulating long-term soil C dynamics as other models. ▪ Presents an extensive assessment of the parameter and input data uncertainty and their importance both temporally and spatially on modelling soil C dynamics. For the basin scale assessments it is estimated that croplands in the Elbe basin currently act as a net source of carbon (net annual C flux of 11 g C m-2 yr-1, 1.57 106 tons CO2 yr-1 entire croplands on average). Although this highly depends on the amount of harvest by-products remaining on the field. Future anticipated climate change and observed climate change in the basin already accelerates soil C loss and increases source strengths (additional 3.2 g C m-2 yr-1, 0.48 106 tons CO2 yr-1 entire croplands). But anticipated changes of agro-economic conditions, translating to altered crop share distributions, display stronger effects on soil C storage than climate change. Depending on future use of land expected to fall out of agricultural use in the future (~ 30 % of croplands area as “surplus” land), the basin either considerably looses soil C and the net annual C flux to the atmosphere increases (surplus used as black fallow) or the basin converts to a net sink of C (sequestering 0.44 106 tons CO2 yr-1 under extensified use as ley-arable) or reacts with decrease in source strength when using bioenergy crops. Bioenergy crops additionally offer a considerable potential for fossil fuel substitution (~37 PJ, 1015 J per year), whereas the basin wide use of harvest by-products for energy generation has to be seen critically although offering an annual energy potential of approximately 125 PJ. Harvest by-products play a central role in soil C reproduction and a percentage between 50 and 80 % should remain on the fields in order to maintain soil quality and fertility. The established modelling tool allows quantifying climate, land use and major land management impacts on soil C balance. New is that the SOM turnover description is embedded in an eco-hydrological river basin model, allowing an integrated consideration of water quantity, water quality, vegetation growth, agricultural productivity and soil carbon changes under different environmental conditions. The methodology and assessment presented here demonstrates the potential for integrated assessment of soil C dynamics alongside with other ecosystem services under global change impacts and provides information on the potentials of soils for climate change mitigation (soil C sequestration) and on their soil fertility status. / Böden speichern große Mengen Kohlenstoff (C) und beeinflussen wesentlich den globalen C Haushalt. Schon geringe Änderungen der Steuergrößen des Bodenkohlenstoffs können dazu führen, dass beträchtliche Mengen CO2, ein Treibhausgas, in die Atmosphäre gelangen und zur globalen Erwärmung und dem Klimawandel beitragen. Der globale Temperaturanstieg verursacht dabei höchstwahrscheinlich eine Rückwirkung auf den Bodenkohlenstoffhaushalt mit einem einhergehenden erhöhten CO2 Fluss der Böden in die Atmosphäre. Weiterhin wirken sich Änderungen im Bodenkohlenstoffhaushalt auf die Bodenfruchtbarkeit und Bodenqualität aus, wobei eine Minderung der Bodenkohlenstoffvorräte wichtige Funtionen des Bodens beeinträchtigt und folglich den Boden als wichtige Ressource nachhaltig beinflusst. Demzufolge ist die Quantifizierung der Bodenkohlenstoffdynamik unter heutigen und zukünftigen Bedingungen von hohem Interesse und erfordert eine integrierte Betrachtung der wesentlichen Faktoren und Prozesse. Zur Quantifizierung wurde ein ökohydrologisches Flusseinzugsgebietsmodell erweitert. Ziel des erweiterten Modells ist es fundierte Informationen zu Veränderungen des Bodenkohlenstoffhaushaltes, neben Veränderungen der Wasserqualität, der Wasserverfügbarkeit und des Vegetationswachstums unter Globalem Wandel in meso- bis makroskaligen Flusseinzugsgebieten bereitzustellen. Dies wird am Beispiel eines zentraleuropäischen Flusseinzugsgebietes (der Elbe) demonstriert. Zusammenfassend ergibt diese Arbeit: ▪ eine Quantifizierung der heutigen und zukünftigen Auswirkungen des Klimawandels sowie von Änderungen der Landnutzung (Bodenbedeckung und Bodenbearbeitung) auf den Bodenkohlenstoffhaushalt agrarisch genutzter Räume im Einzugsgebiet der Elbe. ▪ eine Beurteilung welche Prozesse, und zu welchem Prozessdetail, zur integrierten Simulation der Bodenkohlenstoffdynamik in der meso- bis makroskala zu berücksichtigen sind. Weiterhin wird die Eignung der Modellerweiterung zur Simulation dieser Prozesse unter der Zuhilfenahme von Messwerten dargelegt. ▪ darauf begründet wird eine Prozessbeschreibung vorgeschlagen die die Eigenschaften der Bodenkohlenstoffspeicher und deren Umsetzungsrate mit in der betrachteten Skala zur Verfügung stehenden Messdaten und Geoinformationen verbindet. Die vorgeschlagene Prozessbeschreibung kann als robust hinsichtlich der Parametrisierung angesehen werden, da sie mit vergleichsweise wenigen Modelparametern eine ähnliche Güte wie andere Bodenkohlenstoffmodelle ergibt. ▪ eine umfassende Betrachtung der Modell- und Eingangsdatenunsicherheiten von Modellergebnissen in ihrer räumlichen und zeitlichen Ausprägung. Das in dieser Arbeit vorgestellte Modellsystem erlaubt eine Quantifizierung der Auswirkungen des Klima- und Landnutzungswandels auf den Bodenkohlenstoffhaushalt. Neu dabei ist, dass neben Auswirkungen auf den Bodenkohlenstoffhaushalt auch Auswirkungen auf Wasserverfügbarkeit, Wasserqualität, Vegetationswachstum und landwirtschaftlicher Produktivität erfasst werden können. Die im Rahmen dieser Arbeit dargelegten Ergebnisse erlauben eine integrierte Betrachtung der Auswirkungen des Globalen Wandels auf wichtige Ökosystemfunktionen in meso- bis makro-skaligen Flusseinzugsgebieten. Weiterhin können hier gewonnene Informationen zur Potentialabschätzung der Böden zur Linderung des Klimawandels (durch C Festlegung) und zum Erhalt ihrer Fruchtbarkeit genutzt werden.

Tropical forests are storehouses of more thanhalf of the world‘s biodiversity and play a key role in global carbon, water and energy cycles. However, as a consequence of rapid anthropogenic climate change, biodiversity and climate functions of these forests are under a threat. Climate is changing not only in mean state but its variability is increasing, with extreme events such as droughts, heat waves and storms also rising. Water is fundamental to plants‘ existence, and in the tropics, is a key determinant of plant species‘richness, composition, growth and survival. There is thus an increasing interest in understanding how changing rainfall may cause functional changes in forests or change their species composition. Therefore, the overarching goal of thisdissertation was to understand the impact of water variability on tropical forest tree growth and vulnerability to drought.Forest tree growth along spatial and temporal rainfall gradientsObservational studies that measure whole forest tree growth along spatial or temporal gradients of rainfall are the most common way of formulating forest growth response curves to water availability, when manipulative experiments are cost-prohibitive or impractical (fire or large mammal disturbance). In the tropics, since very few species show anatomically distinct tree rings, estimating tree growth from trunk diameter is the standard practice to obtain growth patterns across species. However, this method—of equating woody growth to diameter change--is susceptible to bias from water-induced stem flexing. In the absence of bias correction, temporal variability in growth is likely to be overestimated and incorrectly attributed to fluctuations in resource availability, especially in forests with high seasonal and inter-annual variability in water. This problem has been largely ignored in the absence of any corrective measure and due to under-appreciation of the magnitude of error. While diameter re-censuses in permanent sampling plots (PSPs) have been most commonly done at 3-5 year scale (using a graduate tape), increasingly they are done at seasonal and annual scales (using band dendrometers) to closely match variation in rainfall, the scales at which hydrostatic bias may be greater in magnitude relative to woody growth. Besides, along a spatial rainfall gradient, inter-annual variability in water may vary, causing systematic differences in the hydrostatic bias for forests along the gradient. Therefore, one broad objective of this thesis was to evaluate the problem of hydrostatic bias in whole forest growth-rainfall relationship at annual and supra-annual scales, for temporal as well as spatial rainfall gradients and propose and test a novel corrective solution.Further, it also examines if growth-diameter relationship vary along the spatial gradient, which it may arise due to differences in light environments and/or disturbance history and species composition. The missing link of Eco-hydrology Differential responses of tree species in terms of growth and survival to variation in water that they can access, the proximate cause is likely shaped through their life-history strategies, the ultimate cause. However, we neither know the depths at which the diverse tree species in a forest draw water from and its dynamics, nor variation in water at those depths vis-à-vis rainfall patterns—for lack of appropriate methods. This has been a key missing link in understanding how water shapes trees‘ life-history strategies, their demographic trade-offs and co-existence, and also our predictive ability to determine species-specific responses to changing rainfall patterns, especially droughts. Since droughts are highly stochastic events and trees‘ responses to their drought ―experiences‖ may be revealed at decadal scales, long-term evaluations are key. Therefore, the second broad objective of this thesis was to develop a framework to determine trees’ water uptake depths, variation in water availability at those depths and trees’ demographic responses over multiple decades. From this, to understand how belowground hydrology shapes drought-vulnerability, demographic trade-offs and coexistence of forest tree species. This thesis titled—Eco-Hydrology of a Seasonally Dry Tropical Forest: Tree Growth, Belowground Water Dynamics and Drought-Vulnerability—is organized as follows: Chapter 1 lays down an introduction to the thesis, followed by a description of the study site and datasets used in the thesis in Chapter 2. This thesis uses a variety of methods and multiple datasets, all of which are from the protected Seasonally Dry Tropical Forests of the Western Ghats in southern India in the Mudumalai and Bandipur National Parks. It is then followed by three data chapters: Chapter 3 describes the seasonal fluctuations in a five year long (1980-1985) tree diameter time series (using dendrometers) of a Seasonally Dry Tropical Forest in Bandipur National Park to illustrate the issue of hydrostatic stem-flexing. It investigates the possibility that band dendrometers may themselves underestimate stem shrinkage at diurnal or seasonal scale. It also evaluates if there could be a best season and time of the day for undertaking forest diameter censuses that can minimize hydrostatic bias. Chapter 4(published in Forest Ecology and Management)measures the hydrostatic bias in a sample of trees in a 50 ha PSP of a Seasonally Dry Tropical Forest in Mudumalai National Park, and proposes a novel way to correct this bias at the whole community level in the 20 year long 4-year interval growth time series. Chapter 5 (in review with Environmental Research Letters) investigates and presents two new confounding factors in growth-rainfall relationships along a spatial rainfall gradient: hydrostatic bias and size-dependency in growth rates. For this it evaluates forest tree growth estimates in seven 1-ha PSPs (~800 trees, 3-year annual time series 9using dendrometers) along a 1000 mm rainfall gradient spanning a mesic savanna-moist forest transition in Mudumalai National Park. Using the period for which seasonal diameter time series was available (2 yrs), it evaluates if the extent of seasonal fluctuations systematically vary along the gradient—most likely due to hydrostatic stem flexing. It also describes the presence of an anomalous size-diameter relationship in the mesic savanna from a large plots (50 ha PSP, diameter records using graduated tape). These observations are then used to draw insights for ―space for time‖ substitution modeling. Chapter 6 (in prep for Nature Plants) analyses belowground water environments of trees over two decades (1992-2012), a period that includes a prolonged and intense drought, in the 50 ha PSP of a Seasonally Dry Tropical Forest in Mudumalai. It uses a locally parametarised dynamic hydrological model in which site rainfall is also a forcing variable. It then develops a novel dynamic growth model and inversely estimates water uptake depths for adult trees of all common species (include ~9000 trees) in the PSP from their above-ground growth patterns over two decades vis-à-vis belowground water availability at multiple depths. It then examines if species‘ water uptake depth obtained thus is a predictor of their drought-driven mortality. Finally, this is used to evaluate the hydrological niche partitioning tree species operate under and how that drives their water uptake strategies, demographic trade-offs, and drought-vulnerability. Summarizes the thesis and suggests future directions