With a new focus on flow regulation by the Tennessee Valley Authority (TVA) in reservoir tailwaters, it is now possible to recover many mussel species that once occurred in these reaches. Before flows can be modified to create habitat for freshwater mussels, suitable microhabitat conditions must be defined. In this study, I used multiple approaches to define suitable microhabitats for species in the free-flowing upper Clinch River, Virginia and Tennessee, where reproducing mussel populations persist. During summer low flows in 2003 and 2004, I measured flow and substrate conditions in over 1000 microhabitat patches (0.25 m² quadrat samples) across five river reaches. Flow characteristics and embeddedness were significantly different between microhabitats occupied and unoccupied by the most abundant species (MRPP, p < 0.05). Comparison of simple and multiple logistic regression models with Akaike's Information Criteria (AIC) demonstrated that increasing Fleisswasserstammtisch (FST) hemisphere number (a measure of shear stress), decreasing degree of embeddedness, and increasing mean column velocity best explained species occurrences in a microhabitat patch. Subtle differentiation in habitat use among species was observed; however, most species appeared to be microhabitat generalists. Species were grouped into three habitat guilds using corresponding canonical analysis and cluster analysis: fast-flow specialists (FFS), fast-flow generalists (FFG), and slow-flow tolerant (SFT). I used the same data set to develop and test transferability of Habitat Suitability Criteria (HSC) for three habitat guilds and seven species of adult freshwater mussels. Nonparametric tolerance limits were used to define the range of suitable and optimal habitat during summer low flows. Optimal habitat was defined as those ranges of FST hemisphere number, mean column velocity, and embeddedness occupied by the central 50% of independent observations for a species or guild, whereas suitable habitat was defined by those ranges occupied by the central 90% of observations. The transferability of criteria to other reaches of the Clinch River was assessed using one-sided Chi-square tests. Criteria developed for the fast-flow specialist (FFS) and fast-flow generalist (FFG) guilds, as well as most criteria for species in those guilds, transferred to destination reaches. In contrast, criteria developed for the slow-flow tolerant (SFT) guild and individual constituent species consistently failed to transfer. Criteria for FFS and FFG guilds and their constituent species should be incorporated into flow simulation models such as PHABSIM to gauge the effect of minimum flows on mussel habitat quality and quantity. These criteria could also be used to determine suitable sites for mussel translocations. However, my criteria require further testing in other rivers before they can be transferred beyond the Clinch River. Behavior and physiological responses to laboratory manipulations of flow velocity and substrate particle size were used to elucidate microhabitat preferences of Actinonaias pectorosa, Potamilus alatus, and Ptychobranchus subtentum. These species appeared less stressed in the fastest flow treatment, demonstrating significantly higher oxygen consumption and oxygen-to-nitrogen (O:N) ratios than in slower flow treatments. Only P. alatus demonstrated a preference for substrate particle size, and consistently selected finer particle sizes. Actinonaias pectorosa and P. subtentum demonstrated preference for fast-flow microhabitats by readily burrowing in those conditions, while abandoning slow-flow conditions. The lack of preference for substrate particle size demonstrated by A. pectorosa and P. subtentum supports conclusions of previous studies that substrate particle size is of little or secondary importance for explaining mussel microhabitat use. These results, along with previous studies in the Clinch River, demonstrate that the stable habitats of riffles and runs; characterized by fast flows during summer low flows, low percent bedrock, and low embeddedness, are the most suitable habitats for mussel assemblages. To create and maintain suitable habitat conditions in tailwaters, releases should maintain flow over riffles at a minimum depth of no less than 30 cm in riffles that provide higher shear stress conditions (FST number > 7) and velocities (> 0.70 m/s). Periodic releases that are sufficient to transport silt and sand, but not high enough to transport larger substrate should be adequate to maintain substrates with a low degree of embeddedness. Doing so would create suitable habitat for all mussels, from the most to least specialized. Additionally, HSC developed for FFS and FFG guilds can be used to determine suitable destination sites for translocations of species belonging to these guilds. / Master of Science

This research tested the ability of fish morphology to predict membership of fishes in habitat guilds, their swimming performance, and habitat preference. Further, it considered methods for choosing a surrogate species to identify habitat of target species. Morphological discriminant functions were developed using morphological traits of fishes from one river to identify membership in two habitat guild systems (mesohabitat and microhabitat). Functions were then used to test factors influencing classification success of holdout tests and validated using fishes of a second river. Morphology was only partly successful (50%) at predicting membership in habitat guilds. Morphology identified species by shape, i.e., classifying test species into guilds with members of their genus, but not habitat use, because morphology and habitat were not strongly linked through function. By improving guild definition, relationships between morphology and habitat (Froude number) were identified for all fish groups examined (darters, benthic minnows, pelagic minnows, and suckers). Relationships were not transferable among groups. Further, morphology of eight minnows was linked to swimming performance, a key task for using habitat, in lab measurements of critical swimming speeds. In turn, swimming performance was related to habitat (Froude number). Morphology will be most successful at predicting habitat use of fishes when (1) more, discrete guilds are used, (2) guilds are identified within families, (3) variation in lifestyles (benthic vs. pelagic) is considered, and (4) key tasks related to using habitat are strongly associated with morphology. Finally, I examined a phylogenetic approach to identifying useable habitat. Closely related surrogate species were not more accurate in identifying habitat of target species than surrogates chosen by other methods. When a target species used only one mesohabitat, the highest overlap in habitat use occurred with other fishes of the same family using that mesohabitat (within a physiographic province). For target species using several mesohabitat types, surrogates from the next highest taxonomic unit, e.g., genus or subgenus, provided the most accurate information. Ecomorphology offers a mechanistic and defensible method for identifying habitat preferences of fishes and should be more widely considered as a tool for establishing habitat relationships of stream fishes. / Ph. D.

The diversity of fish species found in warmwater stream systems provides a perplexing challenge when selecting species for Instream Flow Incremental Methodology (IFIM) studies. An often-suggested approach has been to use habitat guilds to incorporate the diversity found in these systems. My goal is to determine the feasibility of developing habitat suitability criteria (HSC) for the entire fish assemblage in the North and South Fork Shenandoah River, Virginia, using habitat guilds. I examined the strengths and weaknesses of direct underwater observation via snorkeling and throwable anode electrofishing to sample fish habitat use (e.g., depth, velocity, distance to cover, dominant and subdominant substrate, cover, and embeddedness) indicates that using the data collected from both techniques may produce better criteria than using just one of the two sampling techniques. To develop habitat suitability criteria using habitat guilds I placed each species a priori into a guild based on a hypothesized guild structure. Transitional life stages with significantly different habitat use were placed separately into the guild structure. The four guilds (riffle, fast generalist, pool-run, and pool-cover) were found to be significantly different from each other using the data collected for the species assigned to the guilds. Criteria were then developed for representative species from each guild and the entire guilds. Criteria developed for depth, velocity, Froude number, cover, distance to cover, substrate, and embeddedness were used to estimate a habitat response function (i.e., the relations between usable habitat and stream flow) for a representative species from each guild, the guild itself, and for a second species from each guild for comparisons. Both the representative species and guild criteria showed similar habitat response functions for the riffle guild, fast generalist guild, and pool-run guild. However, neither set of criteria performed well for the pool-cover guild. For guilds, other than pool-cover, either the guild or the representative species approach may be a viable option to developing habitat suitability criteria. The transferability tests were performed to determine if criteria developed in the North Fork Shenandoah River, Virginia would transfer to the South Fork Shenandoah River, Virginia. Only criteria for the margined madtom (Noturus insignis) and the juvenile smallmouth bass (Micropterus dolomieu) transferred for both suitable and optimal habitat. Criteria for mottled sculpin (Cottus bairdi), Cyprinella sp. (spotfin and satinfin shiners), river chub (Nocomis micropogon), adult and juvenile redbreast sunfish (Lepomis auritus), and adult smallmouth bass did not transfer. Only the pool-cover guild criteria transferred for both suitable and optimal habitat, while riffle guild, fast generalist guild, and pool-run guild criteria did not transfer. I recommend the use of site-specific criteria for the South Fork Shenandoah or different variable combinations. / Master of Science