Master of Science / Department of Biology / Keith B. Gido / Recognizing habitat needs of fishes across space and time is increasingly important for managing altered stream networks, such as in the Colorado River basin. Recent work on warm-water fishes suggest they might benefit from access to tributaries and their confluences. Fish movements or distributions within tributaries relative to distance from mainstem confluences in two streams with different network types (linear versus dendritic) were investigated in the San Juan River basin, USA. Upstream distance from the San Juan River resulted in species declines (Chaco Wash, linear network) or turnover (McElmo Creek, dendritic network). McElmo Creek movement patterns were likely attributed to spring spawning migrations of flannelmouth sucker (Catostomus latipinnis), spawning aggregations of razorback sucker (Xyrauchen texanus), foraging or refuge seeking by Colorado pikeminnow (Ptychocheilus lucius), and monsoon-related movements for channel catfish (Ictalurus punctatus) and razorback sucker. Razorback sucker and Colorado pikeminnow dominated movements at Chaco Wash, suggesting this backwater-like tributary supplied thermal or current refuge, foraging habitat, or both. Within McElmo Creek, a second study explored the importance of confluences by characterizing habitat use and movements of fishes at the junction of McElmo and Yellow Jacket creeks. Native fish dominated the confluence community composition. The reach downstream of the confluence had consistently higher abundances, species richness, and more frequent detections of tagged fishes relative to upstream reaches. Movement behaviors inferred by detection frequency of tagged fish among reaches surrounding the confluence differed among species. Small flannelmouth sucker (< 300 mm) and roundtail chub (Gila robusta) were commonly detected in Yellow Jacket Creek whereas large flannelmouth sucker (> 300 mm), bluehead sucker (C. discobolus), and channel catfish used McElmo Creek reaches. Monsoons increased McElmo Creek discharge which triggered upstream movements of channel catfish and displaced large flannelmouth sucker and bluehead sucker. Monsoons increased movements between McElmo and Yellow Jacket creeks by roundtail chub, small flannelmouth sucker, and black bullhead (Ameiurus melas). Combined, these two field studies emphasized using links between patterns and processes of tributary fish communities. Conservation, rehabilitation, and maintenance of connectivity and habitat heterogeneity at confluence zones likely can be a localized management strategy with expansive ecosystem effects.

Doctor of Philosophy / Division of Biology / Ari M. Jumpponen / Microorganisms are critically important for establishing and maintaining ecosystem properties and processes that fuel and sustain higher-trophic levels. Despite the universal importance of microbes, we know relatively little about the rules and processes that dictate how microbial communities establish and assemble. Largely, we rely on assumptions that microbial community establishment follow similar trajectories as plants, but on a smaller scale. However, these assumptions have been rarely validated and when validation has been attempted, the plant-based theoretical models apply poorly to microbial communities. Here, I utilized genomics-inspired tools to interrogate microbial communities at levels near community saturation to elucidate the rules and patterns of microbial community assembly. I relied on a community filtering model as a framework: potential members of the microbial community are filtered through environmental and/or biotic filters that control which taxa can establish, persist, and coexist. Additionally, I addressed whether two different microbial groups (fungi and bacteria) share similar assembly patterns. Similar dispersal capabilities and mechanisms are thought to result in similar community assembly rules for fungi and bacteria. I queried fungal and bacterial communities along a deglaciated primary successional chronosequence to determine microbial successional dynamics and to determine if fungal and bacterial assemblies are similar or follow trajectories similar to plants. These experiments demonstrate that not only do microbial community assembly dynamics not follow plant-based models of succession, but also that fungal and bacterial community assembly dynamics are distinct. We can no longer assume that because fungi and bacteria share small propagule sizes they follow similar trends. Further, additional studies targeting biotic filters (here, snow algae) suggest strong controls during community assembly, possibly because of fungal predation of the algae or because of fungal utilization of algal exudates. Finally, I examined various technical aspects of sequence-based ecological investigations. These studies aimed to improve microbial community data reliability and analyses.

Doctor of Philosophy / Department of Geography / Melinda Daniels / Geomorphic properties of streams are linked to ecosystem function through processes related to storage, transport, and other drivers regulating biogeochemical conditions. Disturbances altering the physical template of a stream are associated with cascading impacts on ecosystem function. However, few disturbances are studied at long time scales and so the legacy of such events and the implications for ecosystem structure and function are not well understood. This research investigates the role of historic tie-driving, a channel disturbance legacy, in shaping present-day stream channel conditions in the Rocky Mountain region and the associated implications for organic matter dynamics. Using a combination of geomorphic and riparian surveys, organic matter and vegetation sampling, and modeling, I show that components of mountain stream ecosystems have recovered from tie-driving at varying rates. First, I addressed how tie-driving has altered channel morphology and wood loading. Tie-driven streams are narrower, shallower, less rough, and have less wood than non-driven reference reaches. In a second study, I focused on differences in carbon storage within the stream and riparian area between tie-driven and non-driven streams. Carbon stored on the landscape represents a long-term component of the terrestrial carbon cycle and some, but not all, components have been impacted by tie-driving. Large instream wood, coarse downed wood, and fine downed wood were identified as carbon storage components that were significantly smaller in tie-driven stream-riparian corridors. Finally, I modeled whole stream ecosystem metabolism and tested whether abiotic drivers influenced variations in rates of gross primary productivity (GPP), ecosystem respiration (ER), and net ecosystem productivity (NEP). Results from this work suggest that rates of GPP were significantly different between tie-driven and non-driven streams and were partially explained by variations in light related to canopy structure. However, variations in ER and NEP were not significantly different between tie-driven and non-driven sites. Taken as a whole, this work shows that ecosystems bear the imprint of historic disturbances but individual ecosystem components recover at differing rates. Additionally, integrating stream hydro-geomorphic and ecological dynamics is an effective approach to understanding the impact of channel disturbances in shaping ecosystem function at a variety of spatial and temporal scales.

Master of Science / Division of Biology / Martha E. Mather / In contrast to well documented adverse impacts of large dams, little is known about how smaller low-head dams affect fish biodiversity. Over 2,000,000 low-head dams fragment United States streams and rivers and can alter biodiversity. The spatial impacts of low-head dams on geomorphology and ecology are largely untested despite how numerous they are. A select review of how intact low-head dams affect fish species identified four methodological inconsistencies that impede our ability to generalize about the ecological impacts of low-head dams on fish biodiversity. We tested the effect of low-head dams on fish biodiversity (1) upstream vs. downstream at dams and (2) downstream of dammed vs. undammed sites. Fish assemblages for both approaches were evaluated using three summary metrics and habitat guilds based on species occurrence in pools, riffles, and runs. Downstream of dams vs. undammed sites, we tested if (a) spatial extent of dam disturbance, (b) reference site choice, and (c) site variability altered fish biodiversity at dams. Based on information from geomorphic literature, we quantified the spatial extent of low-head dam impacts using width, depth, and substrate. Sites up- and downstream of dams had different fish assemblages regardless of the measure of fish biodiversity. Richness, abundance and Shannon’s index were significantly lower upstream compared to downstream of dams. In addition, only three of seven habitat guilds were present upstream of dams. Methodological decisions about spatial extent, and reference choice affected observed fish assemblage responses between dammed and undammed sites. For example, species richness was significantly different when comparing transects within the spatial extent of dam impact but not when transects outside the dam footprint were included. Site variability did not significantly influence fish response. These small but ubiquitous disturbances may have large ecological impacts because of their potential cumulative effects. Therefore, low-head dams need to be examined using a contextual riverscape approach. How low-head dam studies are designed has important ecological insights for scientific generalizations and methodological consequences for interpretations about low-head dam effects. My research provides a template on which to build this approach that will benefit both ecology and conservation.

Master of Science / Department of Biology / David C. Hartnett / The effects of bison activities on forb diversity and cover have been well-established, but less is known about how forb life history is altered by bison-mediated habitat changes. This study had three main objectives: 1) to evaluate the hypothesis that release from aboveground competition with grasses may contribute to the increased cover and diversity of forbs in prairie grazed by bison relative to ungrazed prairie, 2) to determine whether differences in forb reproductive effort between grazed and ungrazed habitats were size-dependent, and 3) to look for evidence of a trade-off between allocation to vegetative and sexual reproduction. The growth, biomass allocation, and sexual reproduction of six common unpalatable perennial species were measured and compared between bison-grazed and ungrazed tallgrass prairie burned at 2-year intervals: Ambrosia psilostachya, Artemisia ludoviciana, Baptisia australis, Psoralidium tenuiflorum, Solidago canadensis, and Vernonia baldwinii. Vegetative reproduction was also measured for B. australis¸ S. canadensis, and V. baldwinii. Light availability, canopy density and height, and percent cover of neighboring plants were measured in each studied individual’s immediate neighborhood and compared between habitats to establish the possibility of differing aboveground competition. Aboveground competition may be lower in bison-grazed habitats, as evidenced by differences in habitat characteristics and plant performance found in this study. In bison-present habitats, sexual reproduction was elevated for all six species and average plant size was greater for all species except A. psilostachya. Vegetative reproduction was not clearly different between habitats for all three species examined. Sexual reproduction increased with size for all species, and the relationship differed significantly between habitats for all species except A. psilostachya. Allocation to vegetative reproduction was not generally related to aboveground biomass, nor was there a clear trade-off between allocation to vegetative and sexual reproduction. The results of this study provide evidence that release from aboveground competition with grasses promotes the growth and sexual reproduction of the studied species of forb, and that differences in sexual reproduction are not entirely size-dependent. Patterns in allocation to vegetative reproduction were less clear and were not clearly tied to sexual reproductive allocation.

Doctor of Philosophy / Biology / Walter K. Dodds / Ari M. Jumpponen / Due to agricultural practices and urbanization, tallgrass prairie ecosystems have become threatened as < 5% of its historical coverage exists today. The small remainder of praire that does exist is further threatened by the encroachment of woody plant species. Woody plant encroachment may not only alter prairie ecosystem function, but also prairie microbial communities responsible for these functional processes. Further, prairies are high disturbance ecosystems, especially prairie streams which are hydrologically harsh. They support communities that frequently undergo succession due to recurring flood and drought conditions, yet little is known about the response of microbial communities to these disturbances. In my dissertation, I first address the degree of woody vegetation expansion in riparian corridors (parallel to streams) in watersheds with variable fire frequency and grazing. I found that the rate of riparian woody expansion declines with higher fire intervals and is not affected by grazing, but even annual burns may not prevent woody plant expansion in riparian zones from occurring. Second, I quantified the effect of using restorations of riparian corridors, through removal of woody plants, on physical, chemical, and microbial community (bacteria and fungi) dynamics across stream to upslope soils. Removal restoration causes a decrease in NH₄⁺ and soil water content, and causes streams and upslope soils to become similar in fungal community richness unlike forested landscapes. Bacterial communities were minimally impacted by removals, but were highly structured among stream to upslope soils due to multiple environmental gradients (i.e., pH, NO₃⁻, soil moisture). Lastly, I examined the successional development of biofilm-associated microbial communities in a prairie stream from both a functional and structural perspective. I found that biofilm microbes exhibited strong successional trajectories, with communities developing towards net autotrophy and therefore becoming reliant upon in-stream derived carbon. Further, bacterial communities displayed spatial differences, but much stronger temporal patterns in community composition were detected. These studies highlight how woody plant encroachment may influence stream ecosystems in addition to spatiotemporal trends in microbial community assembly.

Doctor of Philosophy / Department of Biology / Ari M. Jumpponen / Dark septate endophytes (DSE) are darkly pigmented microfungal ascomycetes commonly observed in the healthy plant roots. Studying the functional roles of DSE is challenging as fundamental information about their identity, nutritional requirements, host range or host preference are lacking. Objective 1: root colonizing fungi were isolated from Konza plants roots and DSE fungi were identified by testing Koch’s postulates using leek plants. Periconia macrospinosa and Microdochium sp., were identified as DSE as they produced microsclerotia and chlamydospores in the root cortex. Select DSE were tested for their enzymatic capabilities and ability to utilize nitrogen sources: fungi tested positive for amylase, cellulase, polyphenol oxidases and gelatinase. Periconia isolates utilized organic and inorganic nitrogen suggesting facultative biotrophic and saprotrophic habits. Objective 2: a Microdochium isolate and three Periconia isolates were screened on 16 plant species (six native grasses and forbs, four crops) in a resynthesis system to test host range. DSE colonized all plant species, albeit to varying degrees. Host biomass and nutritional levels to DSE colonization varied within and among host species confirming the broad host range. Based on % responsiveness to DSE colonization, a metric similar to ‘mycorrhizal dependency’, grasses responded positively, while forbs and crops responded negatively. To test this observed ‘host preference’ under natural conditions, Konza roots from seven grass and nine forb species were surveyed for DSE colonization. Grasses hosted 50% greater DSE than forbs, supporting the broad host range and host preference of DSE fungi. Objective 3: three conspecific Arabidopsis ecotypes, Col-0, Cvi-0 and Kin-1 were inoculated with 25 P. macrospinosa isolates in resynthesis system. The three ecotypes responded differently to inoculation: Col-0 and Cvi-0 responded negatively, while Kin-1 response was neutral. Despite the negative or neutral response, each ecotype responded positively to one or two isolates. The outcomes were along the mutualism-parasitism continuum precluding an unambiguous assignment to any particular life-style. This study shows that the outcomes along this continuum are dictated by host and fungal genotypes. However, the more important question about their function remains. Additional studies with Arabidopsis microarrays are likely to provide unique insights into the potential roles of DSE.

Doctor of Philosophy / Department of Biology / Samantha Wisely / Habitat fragmentation and destruction are the most ubiquitous and serious environmental threats confronting the long-term survival of plant and animal species worldwide. However, some native or exotic species can take advantages of these alterations and expand their range, placing endemic species at risk of extinction by changing the composition of biotic communities and altering ecosystem. Capybara (Hydrochoerus hydrochaeris) are a widely distributed rodent throughout most of South and Central America, but restricted to areas of standing water. As the Gran Chaco ecosystem of Paraguay has been converted from dry tropical forest to pastureland, I hypothesized that this habitat alteration created potential for invasion by capybara into newly fragmented areas. I used ecological niche modeling to generate hypotheses about how the distribution of capybara has been affected by land use change, and tested those hypotheses with phylogeographic analyses. To understand the mechanisms that have allowed the invasion, I investigated home range, habitat use and thermoregulation of capybara via radiotelemetry in a deforested area in which capybara had recently invaded. Genetic analyses confirm a rapid range expansion scenario with evidence of secondary contact between two distinct phylogroups which had previously been disjunct. Modeling results indicated that conversion of forest to pastureland allowed the expansion to occur. Capybara selected water significantly more than it was available to them, and avoided shrub forest. I found a significant positive correlation between body temperature and distance from water, and a significant negative correlation between distance from water and Chaco ambient temperature. Capybara proximity to water appeared to be tightly linked to body thermoregulation. These results suggest that although capybara have expanded into the Chaco forest as it is converted to pastureland, the presence of permanent water sources in those pastures are the mechanism that allow capybara to persist in this habitat. This is the first study to characterize capybara in a xeric habitat without a year round water source, and scarce natural grasslands. My results show how anthropogenic habitat modification has allowed capybara to thrive. Understanding how capybara invade and utilize the deforested Central Dry Chaco will provide valuable information for the future management of the species and the Chaco ecosystem.

Master of Science / Department of Biology / Craig Paukert / The North American river otter (Lontra canadensis) was extirpated throughout much of its range but is now recovering in many areas. Consequently, there is a need to determine river otter occupancy and habitat associations. We conducted sign surveys from January to April 2008 and 2009 in eastern Kansas to assess how local- and landscape-scale habitat affects river otter occupancy and how survey methods and habitat affect the detectability of river otter sign. Multiple observers surveyed 3-9 400-m stretches of stream and reservoir shorelines for 110 randomly-selected sites and measured local-scale (within a 100 m buffer of site) habitat variables (e.g., stream order, sinuosity, proportion of land cover types) and landscape-scale (Hydrological Unit Code 14 watershed) habitat variables (e.g., road density, shoreline diversity, proportion of land cover types). We then modeled occupancy and detection probability as a function of these covariates using Program PRESENCE. The overall probability of occupancy accounting for detection probability was 0.329. The best-fitting model indicated river otter occupancy increased with the proportion of woodland cover and decreased with the proportion of cropland and grassland cover at the local scale. The best-fitting model also indicated occupancy increased with decreased shoreline diversity, waterbody density, and stream density at the landscape scale, possibly because of the influence of large reservoirs in the watershed. Occupancy was not affected by land cover or human disturbance at the landscape scale, perhaps due to our relatively homogeneous study area or because river otters are habitat generalists. Detection probability for 400-m surveys was highest in mud substrates (p = 0.600) and lowest in snow (p = 0.180) and litter substrates (p = 0.267). Detection probability for scat was more than double that for tracks, and detection probabilities were 17-64% lower for novice observers than experienced observers. Detection probability also increased with survey length. Sign surveys are a useful technique for monitoring many species, including river otters, and accounting for detection probability will improve estimation of occupancy. Furthermore, understanding the ecological factors and the scale important to river otter occurrence will be useful in identifying areas for restoration and management efforts.

Master of Science / Department of Biology / Keith B. Gido / The upper Gila River basin in southwest New Mexico, USA is one of the few unimpounded drainage basins in North America and is a stronghold for the unique and endemic fishes west of the Continental Divide. Multiple non-indigenous fishes have been introduced to the Gila River and are a potential threat to native fishes, yet very little is known of the trophic ecology of the native and nonnative fishes. We used diet and stable isotopes collected from native and nonnative fishes to identify their trophic relationships and evaluate potential interactions in the upper Gila River basin during June-July, 2007 and 2008. Diet and stable isotope data indicated aquatic invertebrates were the primary food for both native and nonnative fishes. Native large-bodied fishes were mainly algivore/detritivores and native small-bodied fishes were primarily insectivores. Small-bodied nonnative fishes fed on detritus and aquatic invertebrates. Nonnative predators preyed on small-bodied fishes and predaceous aquatic invertebrates and had higher trophic positions than all native fishes. Although nonnative predators did not rely exclusively on native fishes as prey, their presence extended community food-chain lengths, and the combined predation on juvenile native fishes by multiple apex predators may threaten persistence of native fishes. The lack of concise evidence for negative effects suggested that impacts of nonnative predators were more subtle and confirmed the underlying complexity of a relatively simple community The extensive database on feeding relations of Gila River fishes allowed us to further understand how energy moves through ecosystems. Specifically, the goal of chapter two was to characterize variation in fish-community food web structure within and among study reaches on the Gila River using [superscript]13C and [superscript]15N stable isotopes. We hypothesized that food web structure would reflect variation in fish community structure, resource availability and environmental conditions across habitats. Food web structure in isotope bi-plot space was estimated using community-wide measures of trophic structure, mean trophic position, and food-chain length. Permutational multivariate analysis of variance indicated that indices of food web structure were more variable among than within reaches and this pattern was primarily associated with variation in trophicl area occupied by taxa in isotope bi-plot space and mean trophic position of those taxa. Variation in food web structure was significantly associated with fish species richness across macrohabitats but was weakly associated with abiotic reach-scale factors. Variation in food web structure was concordant with variation in fish community composition and suggested that factors influencing the distribution of fishes also influence food web structure.