Cities can be sources of nitrate to downstream ecosystems resulting in eutrophication, harmful algal blooms, and hypoxia that can have negative impacts on economies and human health. One potential solution to this problem is to increase nitrate removal in cities by providing locations where denitrification¬— a microbial process in which nitrate is reduced to N2 gas permanently removing nitrate from systems— can occur. Accidental urban wetlands– wetlands that results from human activities, but are not designed or managed for any specific outcome¬– are one such feature in the urban landscape that could help mitigate nitrate pollution through denitrification.

The overarching question of this dissertation is: how do hydrology, soil conditions, and plant patches affect patterns of denitrification in accidental urban wetlands? To answer this question, I took a three-pronged approach using a combination of field and greenhouse studies. First, I examined drivers of broad patterns of denitrification in accidental urban wetlands. Second, I used a field study to test if plant traits influence denitrification indirectly by modifying soil resources. Finally, I examined how species richness and interactions between species influence nitrate retention and patterns of denitrification using both a field study and greenhouse experiment.

Hydroperiod of accidental urban wetlands mediated patterns of denitrification in response to monsoon floods and plant patches. Specifically, ephemeral wetlands had patterns of denitrification that were largely unexplained by monsoon floods or plant patches, which are common drivers of patterns of denitrification in non-urban wetlands. Several plant traits including belowground biomass, above- and belowground tissue chemistry and rooting depth influenced denitrification indirectly by changing soil organic matter or soil nitrate. However, several other plant traits also had significant direct relationships with denitrification, (i.e. not through the hypothesized indirect relationships through soil organic matter or soil nitrate). This means these plant traits were affecting another aspect of soil conditions not included in the analysis, highlighting the need to improve our understanding of how plant traits influence denitrification. Finally, increasing species richness did not increase nitrate retention or denitrification, but rather individual species had the greatest effects on nitrate retention and denitrification.

It’s no secret that wetlands have dramatically declined in the arid and semiarid American West, yet the small number of wetlands that persist provide vital ecosystem services. Ciénega is a term that refers to a freshwater arid-land wetland. Today, even in areas where ciénegas are prominent they occupy less than 0.1% of the landscape. This investigation assesses the distribution of vascular plant species within and among ciénegas and address linkages between environmental factors and wetland plant communities. Specifically, I ask: 1) What is the range of variability among ciénegas, with respect to wetland area, soil organic matter, plant species richness, and species composition? 2) How is plant species richness influenced locally by soil moisture, soil salinity, and canopy cover, and regionally by elevation, flow gradient (percent slope), and temporally by season? And 3) Within ciénegas, how do soil moisture, soil salinity, and canopy cover influence plant species community composition? To answer these questions I measured environmental variables and quantified vegetation at six cienegas within the Santa Cruz Watershed in southern Arizona over one spring and two post-monsoon periods. Ciénegas are highly variable with respect to wetland area, soil organic matter, plant species richness, and species composition. Therefore, it is important to conserve the ciénega landscape as opposed to conserving a single ciénega. Plant species richness is influenced negatively by soil moisture, positively by soil salinity, elevation, and flow gradient (percent slope), and is greater during the post-monsoon season. Despite concerns about woody plant encroachment reducing biodiversity, my investigation suggests canopy cover has no significant influence on ciénega species richness. Plant species community composition is structured by water availability at all ciénegas, which is consistent with the key role water availability plays in arid and semiarid regions. Effects of canopy and salinity structuring community composition are site specific. My investigation has laid the groundwork for ciénega conservation by providing baseline information of the ecology of these unique and threatened systems. The high variability of ciénega wetlands and the rare species they harbor combined with the numerous threats against them and their isolated occurrences makes these vanishing communities high priority for conservation.

The Upper Verde River of central Arizona flows through a landscape of complex geology at the meeting of seven biotic communities and three physiographic provinces. This has resulted in notably diverse flora and fauna and a hub of rare and endemic plant species. The river has sustained cultures since pre-history, however current regional water use is predicted to diminish streamflow over the next century. Prior to this project, no floristic inventory had been conducted along any section of the Verde. The purpose of this study was to develop a Flora of the Upper Verde River, with the goals of documenting rare and endemic species, the composition and abundance of wetland plants, and the factors shaping plant diversity in the region.

I made a total of 1856 collections and reviewed past collections to produce a checklist of 729 vascular plant taxa in 403 genera and 98 families. The most species-rich family is the Poaceae, followed by Asteraceae and Fabaceae. The flora includes 159 wetland taxa, 47 endemics, and 26 taxa of conservation concern, eight of which are Federally listed. Several new populations were found in these categories and of rarely-collected taxa including one state record, three county records and several range extensions. I report on the local status of several endemics, wetland taxa with limited distributions, and relict populations of a tepary bean (Phaseolus acutifolius) that were likely transported to the region and cultivated by pre-Columbian cultures. I categorize thirteen distinct plant communities, the most abundant being Pinyon/Juniper Woodland, Chihuahuan/Apacherian Scrub, and Riparian Deciduous Forest.

Four primary factors influence floristic diversity of the Upper Verde region: 1) a location at the junction of three physiographic and floristic provinces—represented by co-occurrence of species with affinities to the Sonoran, Intermountain and Madrean regions, 2) geologic diversity—as distinct groups of species are associated with particular geologic types, 3) topographic and habitat complexity—allowing species adapted to disparate environments to co-occur, and 4) human introductions—since over 15% of the flora is composed of introduced species from Eurasia and several taxa were introduced to the region and cultivated by pre-Columbian cultures.

Stream flow permanence plays a critical role in determining floristic composition, abundance, and diversity in the Sonoran Desert, but questions remain about the effects of stream flow permanence on butterfly composition, abundance, and diversity. Understanding the effects of flow permanence on butterflies and relevant subsets of butterflies (such as butterflies whose host plants are present) and comparing them to these same effects on plants and relevant subsets of plants (such as butterfly nectar plants and larval host plants) provided insight into pollinator and riparian conservation and restoration.

I surveyed four Sonoran desert stream sites, and found significant relationships between flow permanence and plant and butterfly species richness and abundance, as well as strong relationships between plant and butterfly abundance and between plant and butterfly species richness. Most notably, my results pointed to hosted butterflies as a break-out category of butterflies which may more clearly delineate ecological relationships between butterfly and plant abundance and diversity along Sonoran Desert streams; this can inform conservation decisions. Managing for hosted (resident) butterflies will necessarily entail managing for the presence of surface water, nectar forage, varying levels of canopy cover, and plant, nectar plant, and host plant diversity since the relationships between hosted butterfly species richness and/or abundance and all of these variables were significant, both statistically and ecologically.

In riparian ecosystems, reptiles and amphibians are good indicators of environmental conditions. Herpetofauna have been linked to specific microhabitat characteristics, microclimates, and water resources in riparian forests. My objective was to relate herpetofauna abundance to changes in riparian habitat along the Virgin River caused by the Tamarix biological control agent, Diorhabda carinulata, and riparian restoration.

During 2013 and 2014, vegetation and herpetofauna were monitored at 21 riparian locations along the Virgin River via trapping and visual encounter surveys. Study sites were divided into four stand types based on density and percent cover of dominant trees (Tamarix, Prosopis, Populus, and Salix) and presence of restoration activities: Tam, Tam-Pros, Tam-Pop/Sal, and Restored Tam-Pop/Sal. Restoration activities consisted of mechanical removal of non-native trees, transplanting native trees, and introduction of water flow. All sites were affected by biological control. I predicted that herpetofauna abundance would vary between stand types and that herpetofauna abundance would be greatest in Restored Tam-Pop/Sal sites due to increased habitat openness and variation following restoration efforts.

Results from trapping indicated that Restored Tam-Pop/Sal sites had three times more total lizard and eight times more Sceloporus uniformis captures than other stand types. Anaxyrus woodhousii abundance was greatest in Tam-Pop/Sal and Restored Tam-Pop/Sal sites. Visual encounter surveys indicated that herpetofauna abundance was greatest in the Restored Tam-Pop/Sal site compared to the adjacent Unrestored Tam-Pop/Sal site. Habitat variables were reduced to six components using a principle component analysis and significant differences were detected among stand types. Restored Tam-Pop/Sal sites were most similar to Tam-Pop/Sal sites. S. uniformis were positively associated with large woody debris and high densities of Populus, Salix, and large diameter Prosopis.

Restored Tam-Pop/Sal sites likely supported higher abundances of herpetofauna, as these areas exhibited greater habitat heterogeneity. Restoration activities created a mosaic habitat by reducing canopy cover and increasing native tree density and surface water. Natural resource managers should consider implementing additional restoration efforts following biological control when attempting to restore riparian areas dominated by Tamarix and other non-native trees.

The explicit role of soil organisms in shaping soil health, rates of pedogenesis, and resistance to erosion has only just recently begun to be explored in the last century. However, much of the research regarding soil biota and soil processes is centered on maintaining soil fertility (e.g., plant nutrient availability) and soil structure in mesic- and agro- ecosystems. Despite the empirical and theoretical strides made in soil ecology over the last few decades, questions regarding ecosystem function and soil processes remain, especially for arid areas. Arid areas have unique ecosystem biogeochemistry, decomposition processes, and soil microbial responses to moisture inputs that deviate from predictions derived using data generated in more mesic systems. For example, current paradigm predicts that soil microbes will respond positively to increasing moisture inputs in a water-limited environment, yet data collected in arid regions are not congruent with this hypothesis. The influence of abiotic factors on litter decomposition rates (e.g., photodegradation), litter quality and availability, soil moisture pulse size, and resulting feedbacks on detrital food web structure must be explicitly considered for advancing our understanding of arid land ecology. However, empirical data coupling arid belowground food webs and ecosystem processes are lacking. My dissertation explores the resource controls (soil organic matter and soil moisture) on food web network structure, size, and presence/absence of expected belowground trophic groups across a variety of sites in Arizona.

Novel resource inputs represent an increasingly common phenomenon in ecological systems as global change alters environmental factors and species distributions. In semiarid riparian areas, hydric pioneer tree species are being replaced by drought-tolerant species as water availability decreases. Additionally, introduced omnivorous crayfish, which feed upon primary producers, allochthonous detritus, and benthic invertebrates, can impact communities at multiple levels through both direct and indirect effects. In arid and semiarid systems of the American Southwest, crayfish may be especially important as detrital processors due to the lack of specialized detritivores. I tested the impact of virile crayfish (Orconectes virilis) on benthic invertebrates and detrital resources across a gradient of riparian vegetation drought-tolerance using field cages with leaf litter bags in the San Pedro River in Southeastern Arizona. Virile crayfish increased breakdown rate of drought-tolerant saltcedar (Tamarix ramosissima), but did not impact breakdown of Fremont cottonwood (Populus fremontii), Gooding's willow (Salix goodingii), or seepwillow (Baccharis salicifolia). The density and composition of the invertebrate community colonizing leaf litter bags were both heavily influenced by litter species but not directly by crayfish presence. As drought-tolerant species become more abundant in riparian zones, their litter will become a larger component of the organic matter budget of desert streams. By increasing breakdown rates of saltcedar, crayfish shift the composition of leaf litter in streams, which in turn may affect the composition and biomass of colonizing invertebrate communities. More research is needed to determine the full extent to which these alterations change community composition over time.

Over the past century in the southwestern United States human actions have altered hydrological processes that shape riparian ecosystems. One change, release of treated wastewater into waterways, has created perennial base flows and increased nutrient availability in ephemeral or intermittent channels. While there are benefits to utilizing treated wastewater for environmental flows, there are numerous unresolved ecohydrological issues regarding the efficacy of effluent to sustain groundwater-dependent riparian ecosystems. This research examined how nutrient-rich effluent, released into waterways with varying depths to groundwater, influences riparian plant community development. Statewide analysis of spatial and temporal patterns of effluent generation and release revealed that hydrogeomorphic setting significantly influences downstream riparian response. Approximately 70% of effluent released is into deep groundwater systems, which produced the lowest riparian development. A greenhouse study assessed how varying concentrations of nitrogen and phosphorus, emulating levels in effluent, influenced plant community response. With increasing nitrogen concentrations, vegetation emerging from riparian seed banks had greater biomass, reduced species richness, and greater abundance of nitrophilic species. The effluent-dominated Santa Cruz River in southern Arizona, with a shallow groundwater upper reach and deep groundwater lower reach, served as a study river while the San Pedro River provided a control. Analysis revealed that woody species richness and composition were similar between the two systems. Hydric pioneers (Populus fremontii, Salix gooddingii) were dominant at perennial sites on both rivers. Nitrophilic species (Conium maculatum, Polygonum lapathifolium) dominated herbaceous plant communities and plant heights were greatest in effluent-dominated reaches. Riparian vegetation declined with increasing downstream distance in the upper Santa Cruz, while patterns in the lower Santa Cruz were confounded by additional downstream agricultural input and a channelized floodplain. There were distinct longitudinal and lateral shifts toward more xeric species with increasing downstream distance and increasing lateral distance from the low-flow channel. Patterns in the upper and lower Santa Cruz reaches indicate that water availability drives riparian vegetation outcomes below treatment facilities. Ultimately, this research informs decision processes and increases adaptive capacity for water resources policy and management through the integration of ecological data in decision frameworks regarding the release of effluent for environmental flows.

Climate change has the potential to affect vegetation via changes in temperature and precipitation. In the semi-arid southwestern United States, heightened temperatures will likely lead to accelerated groundwater pumping to meet human needs, and altered storm patterns may lead to changes in flood regimes. All of these hydrologic changes have the potential to alter riparian vegetation. This research, consisting of two papers, examines relationships between hydrology and riparian vegetation along the Verde River in central Arizona, from applied and theoretical perspectives. One paper investigates how dominance of tree and shrub species and cover of certain functional groups change along hydrologic gradients. The other paper uses the Verde River flora along with that river's flood and moisture gradients to answer the question of whether functional groups can be defined universally. Drying of the Verde River would lead to a shift from cottonwood-willow streamside forest to more drought adapted desert willow or saltcedar, a decline in streamside marsh species, and decreased species richness. Effects drying will have on one dominant forest tree, velvet ash, is unclear. Increase in the frequency of large floods would potentially increase forest density and decrease average tree age and diameter. Correlations between functional traits of Verde River plants and hydrologic gradients are consistent with "leaf economics," or the axis of resource capture, use, and release, as the primary strategic trade-off for plants. This corresponds to the competitor-stress tolerator gradient in Grime's life history strategy theory. Plant height was also a strong indicator of hydrologic condition, though it is not clear from the literature if plant height is independent enough of leaf characteristics on a global scale to be considered a second axis. Though the ecohydrologic relationships are approached from different perspectives, the results of the two papers are consistent if interpreted together. The species that are currently dominant in the near-channel Verde River floodplain are tall, broad-leaf trees, and the species that are predicted to become more dominant in the case of the river drying are shorter trees or shrubs with smaller leaves. These results have implications for river and water management, as well as theoretical ecology.