Freshwater mussels have played an important role in the history of human culture and also in ecosystem functioning. But during the past several decades, the abundance and diversity of mussel species has declined all over the world. To address the urgent need to maintain and restore populations of endangered freshwater mussels, quantitative population dynamics modeling is needed to evaluate population status and guide the management of endangered freshwater mussels. One endangered mussel species, the oyster mussel (Epioblasma capsaeformis), was selected to study its population dynamics for my research. The analysis was based on two datasets, length frequency data from annual surveys conducted at three sites in Clinch River: Wallen Bend (Clinch River Mile 192) from 2004-2010, Frost Ford (CRM 182) from 2005 to 2010 and Swan Island (CRM 172) from 2005 to 2010, and age-length data based on shell thin-sections. Three hypothetical scenarios were assumed in model estimations: (1) constant natural mortality; (2) one constant natural mortality rate for young mussels and another one for adult mussels; (3) age-specific natural mortality. A Bayesian approach was used to analyze the age-structured models and a Bayesian model averaging approach was applied to average the results by weighting each model using the deviance information criterion (DIC). A risk assessment was conducted to evaluate alternative restoration strategies for E. capsaeformis. The results indicated that releasing adult mussels was the quickest way to increase mussel population size and increasing survival and fertility of young mussels was a suitable way to restore mussel populations in the long term. The population of E. capsaeformis at Frost Ford had a lower risk of decline compared with the populations at Wallen Bend and Swan Island. Passive integrated transponder (PIT) tags were applied in my fieldwork to monitor the translocation efficiency of E. capsaeformis and Actinonaias pectorosa at Cleveland Islands (CRM 270.8). Hierarchical Bayesian models were developed to address the individual variability and sex-related differences in growth. In model selection, the model considering individual variability and sex-related differences (if a species has sexual dimorphism) yielded the lowest DIC value. The results from the best model showed that the mean asymptotic length and mean growth rate of female E. capsaeformis were 45.34 mm and 0.279, which were higher than values estimated for males (42.09 mm and 0.216). The mean asymptotic length and mean growth rate for A. pectorosa were 104.2 mm and 0.063, respectively. To test for the existence of individual and sex-related variability in survival and recapture rates, Bayesian models were developed to address the variability in the analysis of the mark-recapture data of E. capsaeformis and A. pectorosa. DIC was used to compare different models. The median survival rates of male E. capsaeformis, female E. capsaeformis and A. pectorosa were high (>87%, >74% and >91%), indicating that the habitat at Cleveland Islands was suitable for these two mussel species within this survey duration. In addition, the median recapture rates for E. capsaeformis and A. pectorosa were >93% and >96%, indicating that the PIT tag technique provided an efficient monitoring approach. According to model comparison results, the non-hierarchical model or the model with sex--related differences (if a species is sexually dimorphic) in survival rate was suggested for analyzing mark-recapture data when sample sizes are small. / Master of Science

The periods of gravidity and glochidial release, required fish hosts, and age and growth characteristics of Fusconaia cuneolus were studied in the upper Clinch River, Virginia during 1986 and 1987. This summer brooder is gravid from mid-May through late July, releasing most glochidia in midJune. Gills of gravid females assume the color of enclosed conglutinates, and hues change from pink to peach as embryos mature to glochidia. As judged by diel sampling of stream drift, glochidia of the fine-rayed pigtoe are most abundant in the water column in early morning. A total of 1,619 fish representing 39 species were electroshocked and examined from the Clinch River at Slant for glochidial attachment. Prevalence of infestation of amblemine glochidia was highest on species of the Cyprinidae (27 to 46%), and six species were identified as possible fish hosts for the glochidia of the fine-rayed pigtoe. Amblemine glochidia were absent on non-cyprinid fish species. Glochidia exhibited host specificity in laboratory experiments, metamorphosing on the following seven minnow and one sculpin species: fathead minnow (Pimephales promelas), river chub (Nocomis micropogon), stoneroller (Campostoma anomalum), telescope shiner (Notropis telescopus), Tennessee shiner (Notropis leuciodus), white shiner (Notropis a/beo/us), whitetail shiner (Notropis galacturus), and mottled sculpin (Cottus bairdi). Tested species of sunfishes, catfishes, and darters did not serve as hosts. Age and growth characteristics were obtained by thin-sectioning shells collected in muskrat middens at Slant and Pendleton Island, Virginia. As predicted by the von Bertalanffy equation, the fine-rayed pigtoe achieves a maximum length of roughly 90 mm and age of 35 yr in the Clinch River. Annual growth in length averaged 5 mm/yr through age 10, decreasing to a rate of roughly 2 mm/yr thereafter. Based upon an age-length key, most individuals (50%) were of intermediate ages (13 to 16 yr). Specimens less than 10 yr old were uncommon, comprising only 22.4% of the deme at Slant. No individuals less than 6 yr old were found at Slant or Pendleton Island. Based on cohort structure at these sites, the fine-rayed pigtoe population appears to be declining in the Clinch River. A monitoring program is suggested. / Master of Science

Three spined, mussel species occur in the United States along the Atlantic slope; James spinymussel (Pleurobema collina), Tar spinymussel (Elliptio steinstansana), and Altamaha spinymussel (E. spinosa). The James spinymussel was listed as endangered in 1988, and was until recently considered to be endemic to the James River basin (Clarke and Neves 1984; USFWS 1990). Biologists with the North Carolina Department of Transportation (NCDOT) discovered spinymussel populations in the Dan and Mayo rivers in NC in 2000 and 2001, respectively. The U.S. Fish & Wildlife Service (USFWS) tentatively identified this species as Pleurobema collina. My project proposed by the Virginia Cooperative Fish and Wildlife Research Unit to the USFWS and the Virginia Transportation Research Council, determined where P. collina resides in VA and what the extent of its range is within the state. An informal preliminary survey design for P. collina was used during the summer of 2002 and simple random sampling was deployed in 2003-2004 surveys to provide a good basis for comparison to gauge the efficiency of the informal sampling design. In 2002, a total of 116 person-hours were spent surveying 39 localities on the Mayo, Dan, and Smith rivers. A total of 96 P. collina was observed in the South Fork of the Mayo River, Patrick and Henry counties, VA. A documented range of 24 rkm was established in the South Fork Mayo River. During the summers of 2003 and 2004, a total of 228 person-hours were spent surveying 38 equal-area river reaches (10, 000 m2) on the mainstems of the Dan, Smith, South Mayo, and Banister rivers. No specimens of P. collina (live or relic shells) were detected. A simple random sampling approach was designed to be easy, relatively quick and cost effective, applicable to most rivers, and to provide actual numbers for comparison. Negative results were only reported after 6 person-hours of searching within each randomly selected, equal-area river reach had been expended. P. collina was declared absent from the VA random sites surveyed in 2003-2004 with a confidence of ~90%. A genetic characterization of four extant populations of P. collina was conducted to assess its taxonomic affinity and to resolve conservation issues related to recovery planning and management actions. The populations were examined for phenotypic variation, and were characterized phylogenetically using DNA sequences. A comprehensive analysis was performed for both separate and combined mitochondrial (357 bp of cytochrome-b, 916 bp of ND-1) and nuclear (502 bp of ITS-1) DNA sequences. Based on comprehensive molecular, morphological, and life history data, populations of P. collina sampled from the Dan River sub-drainage do not warrant separate species designation from P. collina sampled from the James River drainage. / Master of Science

Due to the increasing need to provide refugia for freshwater mussels impacted by anthropogenic activities and exotic species, facilities should be identified and protocols developed for holding mussels in captivity. White Sulphur Springs National Fish Hatchery (WSSNFH), White Sulphur Springs, WV, has held freshwater mussels for nearly eight years, and has the potential to become an important refugium and propagation facility for conservation of mussels in the Ohio River Basin and elsewhere. The goal of this study was to determine the feasibility of holding adult freshwater mussels in long-term captivity at WSSNFH by evaluating survival, energy reserves, and gametogenesis of captive mussels in a recirculating pond system. I relocated three mussel species in the summer of 2001 and 10 mussel species in the summer of 2002 to a recirculating pond system (reservoir and raceway) at the hatchery. Water quality parameters of pH, alkalinity, hardness, temperature, and dissolved oxygen; and algal concentrations were measured periodically from summer 2001 to summer 2003. Annual survival rates of 10 species were estimated (August 2002 to August 2003) using the program MARK. Glycogen, protein, and lipid concentrations in mantle tissue of three captive species (Actinonaias ligamentina, Cyclonaias tuberculata, and Tritogonia verrucosa) were compared to those of wild mussels in the New River. Gametogenic activity and synchrony in A. ligamentina and C. tuberculata were compared between captive and wild mussels. Water quality parameters, with the exception of temperature, were within desirable ranges for most of the study. Temperatures of > 28° C were observed for several days during summers 2002 and 2003. Algal concentrations averaged 1903 cells ml-1 in the raceway (range: 300 to 4658 cells ml-1), which is comparable to algal concentrations reported for nearby rivers. The overall survival rate for 10 freshwater mussel species held in the raceway for one year was 77%. Villosa vanuxemensis had the highest survival rate (96%), and Lampsilis cardium had the lowest survival rate (31%). Although there were fluctuations in glycogen, protein, and lipid levels over 2 yr, there were no overall differences in energy substrates between captive and wild mussels at the end of the study. Captivity did not appear to have a negative affect on gametogenesis. Captive C. tuberculata spawned within the expected time frame between January and June, but slightly earlier than their wild counterparts in the New River. Due to the infestation of the gonads of both captive and wild A. ligamentina by digenean trematodes, little gametogenesis was observed. However, captive holding did not appear to have an effect on trematode infestation rates. From these results, I conclude that captive holding conditions in the recirculating pond system at WSSNFH were adequate for long-term holding of a wide range of mussel taxa. I recommend that WSSNFH continue to be used as an adult holding facility. Further research should be conducted to determine food and habitat preferences of freshwater mussel species in captivity so that optimal holding conditions can be provided for each species. / Master of Science

Epibiosis is an ecological relationship that has been described as one of the closest possible associations in marine ecosystems. In the space limited rocky intertidal, mussel beds provide important secondary space for barnacles. The epibiotic relationship between mussels and barnacles on the south-east coast of South Africa was considered at different scales, from large-scale, natural patterns of epibiosis on the rocky shore, to fine-scale settlement choices of barnacles and the effects on the condition and growth rates of individual mussels. Mussel and barnacle assemblages were generally stable over a 12-month period. The tracking of individual mussels with and without barnacle epibionts resulted in a significant increase in mortality rate of mussels with epibionts over 12 months (two-way ANOVA, p = 0.028). Barnacles on rocks, as well as on mussels, were also tracked with no significant effect of substratum on mortality of barnacles (two-way ANOVA, p = 0.119). Prevalence and intensity of barnacle infestations was also examined in relation to coastline topography on two co-occurring mussel species, the indigenous Perna perna and invasive Mytilus galloprovincialis. The results were complex, but bay status had significant effects on prevalence and intensity for both mussel species, depending on the time and zone. The effect of bay in relation to time was particularly relevant for M. galloprovincialis (four-way nested ANOVA, Season X Site(Bay): p = 0.0002), where summer prevalence was higher than that of winter in bays, regardless of zone, while in open coast sites, the effect of season was only significant in the mid zone. Patterns of intensity generally showed higher values in summer. Substratum preference by barnacles was investigated by recording settlement, survival and mortality of Chthamalus dentatus barnacles on various treatments. There was a strong preference for the rock-like plastic substratum by primary settlers (pair-wise tests of PERMANOVA: Dead < Rock mimic (p = 0.0001); Replica < Rock mimic (p = 0.019) and Live < Rock mimic (p = 0.0001)). This indicates that barnacles settle on mussel shells only as a secondary choice and that micro-topography is an important variable in barnacle settlement. The effect of barnacle epibiosis on condition index and growth of P. perna and M. galloprovincialis was also examined as a direct indication of the health of mussels subjected to the biological stress of epibiosis. Although not significant (PERMANOVA: P. perna: p(perm) = 0.890; M. galloprovincialis: p(perm) = 0.395), growth for both mussel species was slower for barnacle-infested individuals in summer, which is the main growing season for mussels in the region. Results from condition index calculations, however, showed no negative impacts of epibiotic barnacles (three-way ANCOVA: P. perna: p = 0.372; M. galloprovincialis: p = 0.762). Barnacle epibionts create a new interface between the mussel and its environment and this interaction can affect other members of the community. The possibility of the barnacle epibiont causing increased drag also needs further investigation. Biological processes operating within a wide range of physical stressors drive the interactions on the rocky shore, such as epibiosis. Overall, the results of this study suggest that the epibiotic relationship between mussels and barnacles on the south-east coast of South Africa does not significantly affect the mussel species present and that barnacles only use mussel shells as a secondary choice of substratum.

The Leon River, a small-medium river in central Texas, is highly impacted by multiple impoundments, enrichment from agricultural runoff, and decreased dissolved oxygen levels. This degraded river contains sixteen unionid species, two of which are both endemic to the region and candidates for the federal endangered species listing (Quadrula houstonensis and Truncilla macrodon). While there is a short historical record for this river basin and a recent modern survey completed in 2011, zooarchaeological data adds evidence for conservation efforts by increasing the time depth of data available and providing another conservation baseline. Zooarchaeological data for the Leon River is available from the two Late Holocene archaeological sites: 41HM61 and the Belton Lake Assemblages. Data generated from these assemblages describe the prehistoric freshwater mussel community of the Leon River in terms of taxonomic composition and structure. By comparing this zooarchaeological data to the data generated by the longitudinal modern survey of the Leon River, long term changes within the freshwater mussel community can be detected. A conceptual model is constructed to evaluate how robusticity, identifiability, and life history ecology affect unionid taxonomic abundances in zooarchaeological data. This conceptual model functions as an interpretive tool for zooarchaeologists to evaluate forms of equifinality in zooarchaeological assemblages. This thesis determines differences between the late Holocene and modern freshwater community of the Leon River, explores how different alternative mechanisms influence zooarchaeological data, and exemplifies of how zooarchaeological data can be used for conservation biology.

In this thesis, the researcher has attempted to analyze the full complexities of employing the co-management system in conservation of brown mussels (Perna perna) through the use of the case study areas, namely: Sokhulu in KwaZulu-Natal (KZN) and Coffee Bay in the Eastern Cape (EC) Provinces of South Africa. This study is a comparative approach of 2 coastal areas, Sokhulu, in the northern part of KZN and Coffee Bay near the Wild Coast. Communities currently involved in brown mussel resource management in Sokhulu are part of the joint resource management team. The project, started in 1995 by Dr. Harris of KZN Nature Conservation Services (KZN NCS), has proven to be successful in its mission of ensuring wise use of the brown mussel resources. The Coffee Bay project, which was started in 2000, has no local resource management structures yet in place. The research information was gathered with the assistance of household questionnaire; open-ended interviews; literature review; visual review, and commentaries. Experiences drawn from Sokhulu mussel project in KZN have indicated a need for the establishment of a mussel co-management structure at Coffee Bay for the conservation of the resource, and the well-being of the community. Some recommendations for implementation of a co-management system at Coffee Bay have been made. The recommendations are in line with the efficient conservation strategies and utilization of brown mussel resources and blend the needs of the community of Coffee Bay, sound resource use principles provided by the resource managers, and practical management requirements. The study does not present statistically significant conclusions hence it has been highlighted that there is a need for follow-up studies with respect to co-management concept if one is to understand more fully the processes involved. / Thesis (M.Env.Dev.)-University of Natal, Pietermaritzburg, 2002.

Mussels are important ecological engineers on intertidal rocks where they create habitat that contributes substantially to overall biodiversity. They provide secondary substratum for other free-living, infaunal or epifaunal organisms, and increase the surface area for settlement by densely packing together into complex multilayered beds. The introduction of the alien invasive mussel Mytilus galloprovincialis has extended the upper limit of mussels on the south coast of South Africa, potentially increasing habitat for associated fauna. The aim of this study was to describe the structure of mussel beds, the general biodiversity associated with multi- and monolayered mussel beds of indigenous Perna perna and alien M. galloprovincialis, and to determine the relationship between mussels and epibiotic barnacles. This was done to determine the community structure of associated macrofauna and the role of mussels as biological facilitators. Samples were collected in Plettenberg Bay, South Africa, where M. galloprovincialis dominates the high mussel zone and P. perna the low zone. Three 15 X 15 cm quadrats were scraped off the rock in the high and low zones, and in the mid zone where the two mussel species co-exist. The samples were collected on 3 occasions. In the laboratory mussel-size was measured and sediment trapped within the samples was separated through 75 μm, 1 mm and 5 mm mesh. The macrofauna was sorted from the 1 mm and 5 mm sieves and identified to species level where possible. The epibiotic relationship between mussels and barnacles was assessed by measuring the prevalence and intensity of barnacle infestation and the condition index of infested mussels. Multivariate analysis was used on the mean abundance data of the species for each treatment (Hierarchical clustering, multidimensional scaling, analysis of similarity and similarity of percentages) and ANOVA was used for most of the statistical analyses. Overall, the results showed that tidal height influences the species composition and abundance of associated fauna. While mussel bed layering influenced the accumulation of sediments; it had no significant effect on the associated fauna. Time of collection also had a strong effect. While there was an overlap of species among samples from January, May and March, the principal species contributing to similarity among the March samples were not found in the other two months. The outcomes of this study showed that low shore mussel beds not only supported a higher abundance and diversity of species, but were also the most structurally complex. Although the condition index of mussels did not correlate to the percentage cover of barnacle epibionts, it was also evident that low shore mussels had the highest prevalence. The levels of barnacle infestation (intensity) for each mussel species were highest where it was common and lowest where it was least abundant. This is viewed as a natural artefact of the distribution patterns of P. perna and M. galloprovincialis across the shore. Mussels are more efficient as facilitators on the low mussel zone than the high mussel zone possibly because they provide habitats that are more effective in protecting the associated macrofauna from the effects of competition and predation, than they are at eliminating the effects of physical stress on the high shore. Although mussels create less stressful habitats and protect organisms from the physical stress of the high shore, there are clear limitations in their ability to provide ideal habitats. The biological associations in an ecosystem can be made weak or strong depending on the external abiotic factors and the adaptability of the affected organisms.

To realise the potential for mussel culture in British Columbia, mariculture research must identify specific environments and suitable locations which promote maximum growth in mussels. The present study investigates the possible advantages, through nutritional enrichment, of salmon farms as sites for mussel culture. Mussels were cultured at different distances around two salmon farms on the east coast of Vancouver Island (Departure Bay and Genoa Bay). Three parameters of mussel growth: condition index, carbohydrate content, and crude protein content were monitored at 3-6 wk intervals from September 1988 to August 1989. Distinct seasonal differences in growth were observed, but distance from the farm did not substantially influence mussel growth. Adult mortality and larval settlement were similarly unaffected. Contrary to prediction, the farms did not increase available food for mussels. Measures of seston and chlorophyll concentration, made concurrently with the mussel collections, indicated that neither a direct contribution of nutrients in the form of feed and fish faeces, nor an indirect contribution of waste ammonia to augment phytoplankton production, occurred. This was despite currents flowing, at least part of the time, in such a direction as to transport potential nutrients from the farms to the mussels. / Science, Faculty of / Zoology, Department of / Graduate

Sötvattensmusslor av ordningen unionoida är en av världens mest hotade organismgrupper och tjockskalig målarmussla (Unio crassus) klassas som starkt hotad enligt IUCN:s lista över rödlistade arter. Bevarandearbeten har tidigare utförts där tjockskalig målarmussla har återintroducerats till Klingavälsån och Fyleån i södra Sverige. Vid återintroduktion är utgångspunkten genetiskt släktskap, men när den informationen inte finns väljer man att återintroducera den population som är geografiskt närmast. Syftet med studien var att undersöka ifall geografisk avstånd mellan de fyra vattendragen Bråån, Tommarpsån, Klingavälsån och Fyleån påverkar juvenilöverlevnad. Syftet var även att ta reda på vilken av Bråånmusslor och Tommarpsånmusslor som är bäst lämpad för återintroduktion i Fyleån och Klingavälsån. Jag hade följande hypoteser, (1) Bråånmusslor har högst överlevnad i Bråån, följt av Klingavälsån, Fyleån och Tommarpsån och (2) Tommarpsånmusslor har högst överlevnad i Tommarpsån följt av, Fyleån, Klingavälsån och Bråån. Befruktade musslor fångades in från Bråån och Tommarpsån i maj 2016. Musslorna släppte i från sig glochidielarver som sedan användes till att infektera fiskar. Stensimpor (Cottus gobio) elfiskades från Tommarpsån och elritsor (Phoxinus phoxinus) elfiskades från Bråån, därefter infekterades stensimpor med glochidielarver från Tommarpsån och elritsor med glochidielarver från Bråån. Juvenila musslor släppte från fiskarna 25-35 dagar senare och matades under hela sommaren, därefter sattes musslorna ut i boxar i de fyra vattendragen. Överlevnaden för musslorna mättes och dokumenterades vid två tillfällen. Fyra Kruskal-Wallis tester genomfördes där ”överlevnad” jämfördes med ”vattendrag” för Bråånmusslor och Tommarpsånmusslor. Resultatet stämde inte överens med hypotesen för vare sig Bråånmusslor eller Tommarpsånmusslor, förutom att Bråånmusslor hade lägst överlevnad i det mest avlägsna vattendraget vid andra mättillfället. Överlevnaden för Bråånmusslor skilde sig mellan flera av vattendragen men för Tommarpsånmusslor var det ingen skillnad i överlevnad. Bråånmusslor verkade mer lämpliga för återintroduktion i Fyleån jämfört med Klingavälsån och Tommarpsånmusslor verkade inte sämre lämpade för återintroduktion till vare sig Fyleån eller Klingavälsån. Sammanfattningsvis verkade överlevnaden inte generellt bero på geografisk närhet. Troligtvis har faktorer som lokaladaptation, vattenkvalité och sjukdomar även en påverkan på överlevnaden hos juvenila musslor. / Freshwater mussels are one of the most endangered groups of organisms worldwide, with the thick-shelled river mussel (Unio crassus) being classified as critically endangered according to the IUCN:s red list of threatened species. Conservation efforts have been undertaken in southern Sweden, where the thick-shelled river mussel has been reintroduced to Klingavälsån and Fyleån. Genetic relatedness is typically used as the starting point when reintroducing mussels, but if this information is not available, the population closest geographically is chosen. The objective of this study was to investigate whether the distance between four watercourses, namely Bråån, Tommarpsån, Klingavälsån and Fyleån affects juvenile survival and to determine which population of mussels from Bråån and Tommarpsån is best suited for reintroduction to Fyleån and Klingavälsån. The hypotheses are that (1) Bråån mussels have the highest growth rate in Bråån, followed by Klingavälsån, Fyleån and Tommarpsån, and (2) Tommarpsån mussels have the highest growth rate in Tommarpsån, followed by Fyleån, Klingavälsån and Bråån. Fertilized mussels were collected from Bråån and Tommarpsån in May 2016, and the mussels released glochidia larvae, which were used to infect fish. European bullheads (Cottus gobio) were electrofished from Tommarpsån and European minnows (Phoxinus phoxinus) were electrofished from Bråån, and then the European bullheads were infected with glochidia larvae from Tommarpsån and European minnows were infected with glochidia larvae from Bråån. Juvenile mussels let go of the fish 25 – 35 days later and were fed throughout the summer, after which the mussels were placed in boxes in the four watercourses. Survival of the mussels was measured and documented on two occasions. Four Kruskal-Wallis tests were performed where “survival” was compared with “watercourse” for Bråån mussels and for Tommarpsån mussels. The results did not match the hypothesis for either Bråån mussels or Tommarpsån mussels, except for the fact that Bråån mussels had the lowest survival rate in the most remote stream at the second measurement occasion. The survival of the Bråån mussels differed between several of the streams, but for the Tommarpsån mussels there were no significant difference in survival. The Bråån mussels seemed more suitable for reintroduction into Fyleån compared to Klingavälsån, and Tommarpsån mussels did not seem less suitable for reintroduction to either Fyleån or Klingavälsån. In summary, survival did not appear to generally depend on geographic proximity. Factors such as local adaptation, water quality and diseases likely also have an impact on the survival of juvenile mussels.