abstract: Nitrogen is an essential, often limiting, element for biological growth that can act as a pollutant if present in excess. Nitrogen is primarily transported by water from uplands to streams and eventually to recipient lakes, estuaries, and wetlands, but can be modulated by biological uptake and transformation along these flowpaths. As a result, nitrogen can accumulate in aquatic ecosystems if supply is high or if biological retention is low. Dryland and urban ecosystems offer interesting contrasts in water supply, which limits transport and biological activity in drylands, and nitrogen supply that increases with human activity. In my dissertation, I ask: What is the relative balance among nitrogen retention, removal, and transport processes in dryland watersheds, and what is the fate of exported nitrogen? My dissertation research demonstrates that water is a major control on where and when nitrogen is retained and removed versus exported to downstream ecosystems. I used a mass-balance model based on synoptic surveys to study seasonal and spatial patterns in nitrate loading to a dryland stream network. I found that irrigation diversions transport nitrate from agricultural areas to the stream network year-round, even during dry seasons, and are an important driver of nitrate loading. I further explored how seasonal precipitation influences flood nutrient export in an intermittent desert stream by coupling long-term data of flood-water chemistry with stream discharge and precipitation data. I found that higher precipitation prior to a flood fills water storage sites in the catchment, leading to larger floods. In addition, higher antecedent precipitation stimulates biological nitrogen retention in the uplands, leading to lower nitrogen concentration in floods. Finally, I evaluated the consequences of nitrogen export from watersheds on how urban wetlands attenuate nitrate through denitrification that permanently removes nitrogen, and dissimilatory nitrate reduction to ammonium (DNRA) that retains nitrogen in another biologically reactive form. I found that DNRA becomes proportionally more important with low nitrate concentration, thereby retaining nitrogen as ammonium. Collectively, my dissertation research addresses how dryland and urban ecosystems can be integrated into models of watershed nitrogen cycling. / Dissertation/Thesis / Doctoral Dissertation Biology 2019

Headwater streams change in wet length in response to storm events and seasonal moisture conditions. These low-order channels with temporary flow are pervasive across arid and humid environments yet receive little attention in comparison to perennial waterways. This dissertation examines headwater stream length dynamics at multiple spatial and temporal scales across the Appalachians. I mapped wet stream length in four Appalachian physiographic provinces--the Appalachian Plateau, Blue Ridge, New England, and Valley and Ridge--to characterize seasonal expansion and contraction of the wet network at a broad, regional scale. Conversely, most existing field studies of stream length in headwaters are limited to a single study area or geographic setting. Field mappings showed that wet stream length varies widely within the Appalachians; network dynamics correlated with regional geology as well as local site lithology, geologic structure, and the depth, size, and spatial distribution of surficial sediment deposits. I used the field data to create logistic regression models of the wet network in each physiographic province at high and low runoffs. Topographic metrics derived from elevation data were able to explain the discontinuous pattern of headwater streams at different flow conditions with high classification accuracy. Finally, I used flow intermittency sensors in a single Valley and Ridge catchment to record channel wetting and drying at a high temporal resolution. The sensors indicated stream length hysteresis during storms with low antecedent moisture, with a higher wet network proportion on the rising limb than on the falling limb of events. As a result, maximum network extension can precede peak runoff by minutes to hours. Accurate maps of headwater streams and an understanding of wet network dynamics through time are invaluable for applications surrounding watershed management and environmental policy. These findings will contribute to the burgeoning research on temporary streams and are additionally relevant for studies of runoff generation, biogeochemical cycling, and mass fluxes of material from headwaters. / Ph. D. / During a rain storm, we may think of streams increasing in depth, width, and velocity. However, we may not necessarily envision streams also getting longer. Headwaters, which form the upstream extremities of river systems, consist of many temporary streams that expand and contract in length due to storms and changes in seasonal moisture conditions. Headwaters are spatially expansive, comprising a majority of total river length, and serve as a primary control on downstream water quality. Therefore, understanding stream length dynamics can inform policy and land use decisions to effectively conserve and manage headwater regions and protect water sources for human use and consumption. This dissertation examines changes in stream length across four study areas of the Appalachian Mountains. I mapped the wet, or active, stream network multiple times at different flow conditions in each study area. Stream length dynamics varied considerably across the Appalachians and demonstrated the same range of network expansion and contraction as other studies observed in diverse settings around the world. Wet stream length greatly depended on regional and local geology. I then sought to predict the location of wet streams at high and low flows using metrics such as slope and drainage area that I calculated from digital elevation information. Comparisons with the field maps I made showed that simple terrain metrics explained the location, length, and disconnected nature of wet networks in each province with high accuracy. I also observed stream length dynamics during storm events in one watershed using sensors that recorded the presence or absence of water. These observations demonstrated that stream length was often higher for a given flow at the beginning of a storm on the rising limb than on the falling limb when flow was decreasing, particularly if conditions were dry before the storm. The findings of this dissertation contribute to existing knowledge of temporary streams and are relevant for future studies investigating the hydrology, biology, and ecology of headwaters.

Temporary streams and their dynamics have often been largely overseen in hydrological research and there is relatively little knowledge about how the occurrence of flow in these streams varies temporally and spatially. Temporary streams are important from a hydro- logical perspective because they affect water quantity and quality in downstream peren- nial reaches, and from an ecological perspective because they provide habitat to unique species. In order to gain knowledge about these important streams, this maser thesis was conducted, within the Msc program in Water and Environmental Engineering at Uppsala University and the Swedish University of Agricultural Sciences, in collaboration with the Hydrology and Climate group at the University of Zurich. In this study, the temporal and spatial variation of the temporary streams in a small pre-Alpine catchment in Switzerland were investigated, both in terms of the presence of flowing water and stream chemistry. The 20 ha Studibach catchment is typical for the pre-Alpine area, with frequent precipi- tation. The streams in the lower part of the Studibach catchment were mapped in the field during September 2020. The temporal and spatial variations of the presence of flow and stream chemistry within the stream network was investigated in September and October 2020 during varying weather conditions. During ten field campaigns the flow state of the streams was classified and the Electrical Conductivity (EC) of the streams was mea- sured approximately every 20 meter. The findings from the field campaigns were related to topographic indices, in particular the Topographic Wetness Index (TWI) and Upslope Accumulated Area (A), in order to see how topography influenced the presence of stream- flow and stream EC. The results show a high temporal and spatial variation in both stream chemistry and streamflow. The active network length expanded by a factor of two in re- sponse to precipitation events. The stream EC also had a large spatial variation, and the streams in the southeast part of the catchment had a higher EC than the other streams. This spatial variation is expected to reflect the large variability in groundwater EC within the catchment. The spatial variation of the streamflow demonstrated a difference between the north-middle and the south part of the catchment, where the south part responded quicker to events and drained and retracted faster after the event. The findings also indicate that topographic indices can predict the occurrence of flow in the stream network, with sites with higher topographic index values having a higher probability of flowing water in the stream. Topography also influences the stream chemistry. The variation in stream chem- istry was smaller for sites with higher values for the topographic indices, something that can be explained by the Representative Elementary Area (REA) concept, because sites with higher topographic index values are located further downstream and water at these locations is a mixture of the smaller streams that feed these streams. / Temporära bäckar och dess dynamik har länge varit förbisedda inom hydrologisk forskn- ing, och en djupgående kunskap rörande temporära och rumsliga variationer saknas. Tem- porära bäckar är viktiga utifrån ett hydrologisk perspektiv eftersom de påverkar både kvantitet och kvalitet på vattnet nedströms, och från ett ekologiskt perspektiv eftersom de bidrar med habitat till unika arter. Detta examensarbete har genomförts för att öka kunskapen kring dynamiken i dessa temporära nätverk. Examensarbetet genomfördes inom Civilingenjörsprogrammet i Miljö och Vattenteknik vid Uppsala Universitet och Sveriges Lantbruksuniversitet, i ett samarbete med Hydrologi- och Klimatgruppen vid University of Zurich. Studien har undersökt temporära och rumsliga variationer i ett tem- porärt bäcknätverk med avseende på flöden och kemin i vattnet, i ett mindre pre-alpint avrinningsområde i centrala Schweiz. Bäckarna i den nedre delen av avrinningsområdet Studibach karterades i fält för hand med karta och kompass under september 2020. Avrin- ningsområdet är på 20 ha och räknas som typiskt för ett pre-Alpint område, med frekvent nederbörd. Tio fältkampanjer genomfördes där temporära och rumsliga variationer un- dersöktes genom klassificering av flöden och mätningar av Elektrisk Konduktivitet (EC) i bäckarna ungefär var 20e meter, under september och oktober 2020 i varierande väder- förhållanden. Resultaten från fältkampanjerna relaterades till de topografiska indexen Topographic Wetness Index (TWI) och Upslope Accumulated Area (A) för att undersöka hur topografin påverkar flöden och bäckkemin. Studien kom fram till att bäckarna i den nedre delen av Studibach visar både en temporär och en rumslig variation för både flöde och bäckkemi. De aktiva bäckarna i nätverket visade på en expansion med en faktor två som svar på nederbörd. En rumslig variation för flödet påträffades även mellan den södra och nord-centrala delen av nätverket där den södra svarade snabbare mot event och även drogs ihop snabbare. Kemin i bäckvattnet visade på en stor rumslig variation, med högt EC i den sydöstra delen av avrinningsområdet, vilket förmodas bero på den stora rumsliga variationen av EC i grundvattnet. Resultaten visar även på att topografiska index kan till viss del förutspå flöden i bäckarna, där platser med högre topografiska index har högre sannolikhet att det flödar i bäcken. Topografin påverkar även bäckkemin. Variationen i bäckkemin var mindre för platser med högre topografiska index, vilket kan förklaras med Representative Elementary Area (REA) konceptet, eftersom platser med högre to- pogragiska index värden återfinns längre nedströms och vattnet på dessa platser är en blandning av de mindre bäckarna som tillför vattnet till de större.