Tidal freshwater wetlands provide valuable ecosystem services such as nutrient cycling, coastal flood control, and reproductive habitat for early life stages of economically and ecologically important fishes. The nutrient rich environments that tidal freshwater wetlands occur in support high levels of primary productivity of phytoplankton and vegetation that provide essential reproductive habitat for anadromous clupeid fishes (Alosa spp., Brevoortia tyrannus) in Chesapeake Bay tributaries. Seasonal retention of clupeid eggs and larvae within tidal freshwater habitats may enhance early growth, survival, and year class strength. The primary goal of the present study was to characterize the relative importance of tidal freshwater wetlands to the early life history stages of anadromous, migratory, and resident clupeid fishes in the lower James River. From 2014-2018, conical tow nets were used to collect ichthyoplankton samples from representative locations within the tidal freshwater marsh-creek complex of Curles Neck Creek, Virginia during the period of February through May. Boat electrofishing was used to sample juvenile target clupeids within the marsh-creek complex during March through November 2014-2018. A strong positive correlation between larval and juvenile Blueback Herring and Atlantic Menhaden, supported the hypothesis of seasonal in-system retention. This was the first published study to describe spatial and temporal patterns of occurrence of clupeid fishes with varying life history strategies in a tidal freshwater marsh and highlighted the importance of tidal freshwater marshes as nursery habitat.

Understanding, quantifying, and comparing the temporal distribution of anadromous fish spawning migrations is an important yet vexing problem for fisheries research, management, and conservation. Central to this problem is the lack of a representative and comprehensive standardized suite of quantitative metrics to characterize the complex, multidimensional temporal distribution of migrating anadromous fish. Without this, it is not possible to develop effective sampling regimes, extrapolate counts to accurate population estimates, understand the basic ecology and behavior of anadromous fish, or make the comparisons through time and across river systems that are fundamental to sustainable conservation. In this thesis, I define, calculate, and compare 17 metrics that characterize the temporal distribution of migrating adult river herring [two closely related clupeids, the alewife (Alosa pseudoharengus) and blueback herring (Alosa aestivalis)] as they return to spawn. These metrics are based on fish counts from three southeastern Massachusetts river systems that were obtained through a low-cost video monitoring system.

Nature-like fish passes have been designed with the intent to re-connect river corridors and provide passage for all species occurring in a system. Nature-like fish pass designs have been constructed in Europe and elsewhere with some success, but performance of these designs has not been evaluated for North American species. Re-establishing passage for adult anadromous clupeids to their spawning areas is critical considering their recent dramatic population declines. Two nature-like fish pass designs in New England were evaluated for passage of anadromous adult alewives (Alosa pseudoharengus) using passive integrated transponder (PIT) telemetry and showed differing results. At Town Brook in Plymouth, Massachusetts the 32 m long perturbation boulder rock ramp with a 1:24 slope passed 94% of attempting fish with most ascending in under 22 minutes. At East River in Guilford, Connecticut the 48 m long steppool bypass design with a 1:14 slope passed only 40% of attempting fish with a median transit time of 75 minutes. Two technical fishway designs at the field sites were also evaluated and showed contrasting performance. At Town Brook a 14 m long 1:7 slope pool and weir fishway exhibited attraction and passage deficiencies. At East River two 3.05 m long steeppass fishways both passed the majority of attempting fish but one steeppass fishway may have had poor attraction efficiency. At both sites tagged fish passed rapidly downstream through the fish passes after spawning. Nature like fish pass designs are suitable for the passage of alewife but further evaluations are required to more precisely identify the influence of vertical drop per pool and specific local hydraulics on behaviors and passage performance for this species.

River herring, collectively alewives (Alosa pseudoharengus) and blueback herring (A. aestivalis), have experienced substantial population declines over the past five decades due in large part to overfishing, combined with other sources of mortality, and disrupted access to critical freshwater spawning habitats. Anadromous river herring populations are currently assessed by counting adults in rivers during upstream spawning migrations, but no field-based assessment methods exist for estimating juvenile densities in freshwater nursery habitats. Counts of 4-year-old migrating adults are variable and prevent understanding about how mortality acts on different life stages prior to returning to spawn (e.g., juveniles and immature adults in lakes, rivers, estuaries, and oceans). This in turn makes it challenging to infer a link between adult counts and juvenile recruitment and to develop effective management policy. I used a pelagic purse seine to investigate juvenile river herring densities, growth, and mortality across 16 New England lakes. First, I evaluated the effectiveness and sampling precision of a pelagic purse seine for capturing juvenile river herring in lakes, since this sampling gear has not been systematically tested. Sampling at night in June or July resulted in highest catches. Precision, as measured by the coefficient of variation, was lowest in July (0.23) compared to June (0.32), August (0.38), and September (0.61). Simulation results indicated that the effort required to produce precise density estimates is largely dependent on lake size with small lakes (<50 >ha) requiring up to 10 purse seine hauls and large lakes (>50 ha) requiring 15–20 hauls. These results suggested that juvenile recruitment densities can be effectively measured using a purse seine at night in June or July with 10–20 hauls. Using juvenile fishes captured during purse seining in June–September 2015, I calculated growth and mortality rates from sagittal otoliths. Density, growth, and mortality were highly variable among lakes, and mixed-effects regression models explained 11%–76% of the variance in these estimates. Juvenile densities ranged over an order of magnitude and were inversely related to dissolved organic carbon. Juvenile growth rates were higher in productive systems (i.e., low secchi depth, high nutrients) and were strongly density-dependent, leading to much larger fish at age in productive lakes with low densities of river herring compared to high density lakes. Water temperature explained 56%–85% of the variation in juvenile growth rates during the first 30 days of life. Mortality was positively related to total phosphorous levels and inversely related to hatch date, with earlier hatching cohorts experiencing higher mortality. These results indicate the importance of water quality and juvenile densities in nursery habitats for determining juvenile growth and survival. This study encourages future assessments of juvenile river herring in freshwater and contributes to an understanding of factors explaining juvenile recruitment that can guide more effective and comprehensive management of river herring.

Conserving and restoring anadromous fish populations is an important research and management priority. For conservation to be effective, researchers must understand the behavior of the fish they seek to restore. Telemetry has allowed researchers to understand the upstream migrations of these fish in freshwater, how migration patterns vary, and if there is a relationship between behavior and environmental variables. In the northeastern United States, alewife (Alosa pseudoharengus), one of two species collectively referred to as river herring, has historically been an important component of coastal rivers. However, populations of these fish have experienced recent declines, and a commonly used method to restore river herring is stocking. In this thesis, I summarize research that seeks to understand anadromous alewife behavior with the goal of providing insights that will help manage and conserve this species and the coastal systems in which they live. My thesis has a primary research chapter (Chapter 1), a second ancillary research chapter (Chapter 2), and four appendices that summarize related information as part of the funding proposal. First, to examine if fish origin (native or stocked) and fish release location (upstream or downstream) affected the pre-spawning movements of fecund alewives, I undertook a reciprocal experiment. In Chapter 1, for fish of both origins and release locations, I examined how long fish were in the river, where they spent their time, and how much and how fast they moved. For this, I gastrically tagged alewives with Lotek Nanotags NTC-6-1 radio tags and monitored movements in the lower 30 km of the Ipswich River (northeastern Massachusetts) using an array of 9 Lotek SRX_400 receivers. Based on these movement trajectories I concluded that in 2007, origin affected the total time fish spend in the river and release location affected where they spend their time. Downstream movements of upstream migrating fish have typically been viewed as a behavioral assay of adverse tag effects. For this reason, alosine telemetry studies rarely release tagged fish upstream of the capture site. However, fisheries managers often release fish upstream near spawning grounds during stocking. In Chapter 2, I re-evaluated whether downstream movements of upstream stocked fish were consistent with an adverse tag effect. By combining physiological experiments with select movement trajectories, I showed that pre-spawning migrations of alewife included an array of up and downstream directed movements with various interpretations. In my research, these downstream movements were unlikely to be related to tagging stress (Chapter 2), as the cortisol, glucose, and chlorides of tagged fish were not different from untagged fish (Appendix A, Physiology). Furthermore, I suggested metrics that should be recorded in telemetry studies to standardize how downstream fish movements are measured. In 2006, native fish were released at a downstream site (river km 6) and stocked fish were released upstream (rkm 25). I compared the behaviors of these same treatments across years. I showed that the behaviors of fish released in different years may differ based on temperature and discharge (Appendix B, Across Year Comparison). To determine the amount and location of potential spawning habitat, I undertook a habitat study that utilized a geographic information system (GIS) to maps the size and distribution of habitat types. I located multiple mainstem pools in the Ipswich River that may serve as suitable spawning grounds for alewife. Tagged fish were primarily located in these habitats (Appendix C, Habitat). To determine if juveniles were produced, I sampled various sites in the river for the presence of juveniles, using active and passive sampling techniques. Juveniles were not captured during these surveys (Appendix D, Juvenile Sampling). Before this research, little was known about the pre-spawning migrations of river herring. While river herring are assumed to be a generalist species, I found their behaviors to be complex. I have identified a number of gaps in the current knowledge of how these fish behave in the field. Restoration efforts must take into account the behavior of the fish, as well as the capacity of a system to accommodate those needs. Within the context of understanding fish behavior, protecting habitat, and providing regulatory restrictions on the fishery, stocking may contribute to broader management and restoration goals.

Over the last century anadromous alewife (Alosa psuedoharengus) and blueback herring (Alosa aestivalis), collectively called river herring, suffered drastic declines throughout their range from Newfoundland (Canada) to North Carolina (USA). A 2011 petition to include river herring in the Endangered Species Act (ESA) was rejected, partly due to inadequate information towards identifying coast-wide population status. Additionally, knowledge gaps were identified with basic ecology of the river herring life cycle in freshwater, including species reproductive strategies. In Chapter 2, I investigated how body size, spawning arrival time, and sex influence river herring reproductive success. I collaborated with the Massachusetts Division of Marine Fisheries to introduce adult river herring (421, 266, and 410 individuals in 2013, 2014, and 2015, respectively) into Pentucket Pond (Massachusetts, USA), which historically had river herring but is currently inaccessible to upstream migrants. Using fin clips from stocked adult fish and juveniles collected in the lake each summer, I genotyped individuals and constructed pedigrees with 15 microsatellites. River herring had small (mean =1.1) families and spawned multiple times with multiple mates from May to June. Females were more successful than males. Earlier arrival and larger body size were independent indicators of reproductive success. These results provide critical river herring life history information for the freshwater component of population models that will inform management of this at-risk species. Presently, most river herring populations are monitored using data from citizen counts of spawning adults entering freshwater. Involving citizen watershed groups in data collection and may provide ancillary benefits beyond collection of population-level count data. In Chapter 3, I used pre-and post-surveys to assess how involvement in one citizen monitoring program influenced participants’ environmental knowledge, attitudes, and behaviors. Participants began the program with high scores for broader scientific and pro-environmental attitudes and pro-environmental behaviors. After the program, participants reported increased connection to nature, citizen science involvement, river herring knowledge, and engagement in outdoor recreation. For participants, engagement with nature was the most important program benefit. These results provide an additional case study to the citizen science literature and demonstrate that citizen science programs can help participants connect with the environment.

Beginning in the early 1600s, dam construction in New England obstructed anadromous fish access to spawning grounds during migration. As a result, anadromous forage fish populations have declined, which has impacted freshwater, marine, and terrestrial ecosystems. To determine the impacts of dams on anadromous forage fish and freshwater ecosystems, I used historical and current data to estimate population changes in alewives (Alosa pseudoharengus) from 1600-1900. A significant reduction in spawning habitat occurred in New England as a result of 1,642 dams constructed between 1600 and 1900, resulting in 14.8% and 16.6% lake and stream habitat remaining by 1900, respectively. In eight New England watersheds, this translates to an estimated cumulative annual loss of 30 B juvenile alewives available as freshwater forage and 538 M year 1, 2 and 3 alewives available as marine forage. The cumulative annual lost number of adult return spawners was conservatively 17 M fish, or 3,642 metric tons. Lost marine-derived nutrients from adult return spawners were 11 T phosphorus, 64 T nitrogen, and 410 T carbon. A comparison of predator fish growth and condition in alewife and non-alewife lakes showed that white perch (Morone Americana) and yellow perch (Perca flavescens) have higher condition in early summer in lakes with alewives. Predator growth rates (length-at-age) were significantly higher in early life stages (ages 1 and 2) when alewives were present, but significantly lower in late life stages (ages 3 and older). Results indicate a greater maximum length obtained by mature fish when alewives are absent, and an earlier age and length at maturity when alewives are present. These results indicate significant ecosystem impacts of lost anadromous forage fish, with bottom-up trophic effects across multiple time scales and biological processes. An ecosystem-based management approach should be used by inland and marine aquatic managers, and ecosystem connectivity and trophic interactions should be considered when managing migratory fish and prioritizing restoration goals.