Eelgrass-macroalgae interactions; context-dependency in upwelling-influenced estuaries

This dissertation investigates the context-dependency of species interactions between seagrass and macroalgae in upwelling-influenced estuaries. In all coastal systems, nutrient loading is multidirectional, resulting from mostly freshwater and marine inputs. The directionality of nutrient inputs may affect the rate of supply of organic matter to the system. In systems where freshwater nutrient loading dominates, and has increased through time, research shows that blooms of fast-growing macroalgae often result in loss of critical seagrass habitats. In upwelling-influenced systems, marine- based nutrient inputs dominate during the summer, also resulting in blooms of ulvoid macroalgae during these productive months. The dominance of marine nutrients in these estuaries, coupled with additional variation in the physicochemical characteristics of seagrass beds, present novel contexts to study the outcomes of species interactions between the seagrass (Zostera marina L. (eelgrass) and ulvoid macroalgae. I studied these interactions at two different spatial scales that both address the relative importance of marine versus terrestrial nutrient sources on interaction outcome. Regionally, I studied between-estuary, latitudinal patterns in species interactions relative to differences in marine and terrestrial drivers of nutrient loading. Within an estuary, I also compared interactions among zones along an estuarine gradient, where nutrient patterns were reflective of the relative contribution of marine- based nutrients. At both scales of inquiry I employed both observational and experimental approaches to quantify species interaction dynamics. At the regional scale, I used a 5- year observational dataset from four estuaries along the Oregon and Washington coasts to study the relationship between eelgrass and ulvoid macroalgae (Chapter 2). Across latitudes that span ~220 km, macroalgal production was highest in the southern estuaries, and associated with decreased eelgrass production compared to the northern estuaries. However, through time, no estuarine site, regardless of its macroalgal biomass, was associated with declining eelgrass biomass. Contrary to systems where macroalgal production is driven by terrestrial inputs, I found that blooms in upwelling- influenced systems were associated with both marine and terrestrial drivers of nutrient inputs and production. In Coos Bay (South Slough), at the within-estuary scale, I also found differences in macroalgal and eelgrass biomass among sites along an estuarine gradient. Here too, based on a 2-year seasonal dataset of producer dynamics, I found no temporal relationship between eelgrass and macroalgae producer dynamics (Chapter 3). I used a comparative-experimental framework to understand the impact of macroalgal manipulations (additions and removals) on interactions with eelgrass along this gradient. In intertidal seagrass beds in the marine and polyhaline zones of the estuary I found that interaction strength was neutral and sometimes positive. However, in the riverine zone, interaction strength was negative, caused by decreased eelgrass density following macroalgal manipulation. To further examine the mechanisms informing interaction outcomes in the marine zone, a large-scale macroalgal manipulation was conducted, coupled with a mesocosm experiment (Chapter 4). For the mesocosm experiment I manipulated macroalgae and nutrients as in the field, but found dissimilar results. In the mesocosms, where water movement was limited and no tidal action occurred, negative effects of macroalgal addition were found. These were associated with increased light attenuation and decreased sediment oxygen levels. Contrary to these results, I found no macroalgal, or covariate effects in the field experiment. I also manipulated water column nutrients in both experiments, and found limited effects of nutrient enrichment on eelgrass, but not macroalgae, in the mesocosm experiment. Throughout these studies I demonstrated that the mechanisms determining context-dependency in upwelling-influenced estuaries are informed by physical and biogeochemical conditions, coupled with high ambient marine-derived nutrient concentrations. These findings are important to coastal management because they suggest that the strength, direction and mechanisms of interactions are shaped by local abiotic conditions and long-term nutrient regimes, rather than high nutrient concentrations per se. Given the shifting nature of nutrient concentrations in coastal waters associated with both coastal development and climate change, knowledge of context dependency can also be used to assess and forecast future changes in species interactions

Seagrass nursery function enhanced by habitat connectivity

Seascape connectivity is an emerging feature influencing marine ecosystem function. Connectivity, via the interplay of habitat configuration, its physical structure, and associated food webs, may alter the provision of nursery habitat, influencing the success of fish populations and harvests thereof. We quantified the effects of adjacent habitat type on seagrass nurseries for rockfish. We compared seagrass sites with contrasting structure and productivity in adjacent habitats: kelp forest and sand. To elucidate the spatially-dependent flow of material in the seagrass meadow, we estimated the proportion of basal contributions (producers) to the food web using an isotopic mixing model on YOY (young-of-the-year) rockfish consumers. To estimate population responses to spatial connectivity, we enumerated YOY rockfish recruitment by SCUBA, alongside invertebrate prey and fish predators at all sites. Using GLMMS, we modelled the relative effects of adjacent habitat type, seagrass trophic interactions (prey availability and predator presence), and structural complexity, on YOY rockfish recruitment. We found that kelp carbon was the highest basal contributor to YOY rockfish consumers across all seagrass sites, but auxiliary contributions varied among site. YOY rockfish at highly complex and productive seagrass sites (adjacent to kelp forests and interior meadow) consumed higher quality prey, corresponding to increased body condition. Adjacency to kelp forests, seagrass structural complexity, and prey biomass increased YOY rockfish densities. However, the positive effect of kelp adjacency was dampened by seagrass complexity; likely due to predator spill-over into complex seagrass habitat. Overall, recruitment and trophic dynamics (e.g. basal contributions, prey-predators interactions) in the seagrass meadow were influenced by seascape configuration, revealing that nursery function is spatially-mediated by adjacent habitats, and their structural complexity and productivity. With declining marine habitats worldwide, we illustrate the importance of considering connectivity among nearshore habitats for conservation planning, supporting an ecosystem approach that considers the full seascape matrix of intermixed habitats

Discerning population connectivity and natal origins of Pacific herring (Clupea pallasi): inferences on population structure from otolith chemistry

Pacific herring, Clupea pallasi, undertake annual migrations between feeding and spawning grounds that link life stages, habitats, populations, communities, and ecosystems. However, movement patterns of these highly mobile fish are poorly understood. Declines in Pacific herring abundance and slow population recoveries in the absence of fishing pressure have elevated concerns over the status of this ecologically, economically, and culturally important species. Pacific herring spawn on substrate in nearshore habitats where eggs and larvae develop for approximately two weeks before hatching. Early development within discrete spawning grounds could facilitate the incorporation of distinctive chemical signatures within otoliths that could be used as intrinsic markers to trace movements and mixing among groups or regions. Identifying the direction and strength of connectivity among groups can reveal source populations and promote the development of population- and ecosystem-based management strategies that reflect ecologically relevant spatial scales. We applied otolith microchemistry data to: 1) test the utility of elemental signatures to distinguish the natal origins of larval herring; 2) evaluate inter-annual variation in natal signatures within spawning sites; and 3) assess the similarity of edge and natal signatures of adult herring within and among spawning sites. In 2015 and 2016, we sampled actively spawning adult herring and their offspring in the northern Salish Sea and across British Columbia, Canada. Otoliths were extracted, aged, and their elemental composition analyzed using laser ablation inductively coupled plasma mass spectrometry. Cohort-specific analyses were applied to assess consistency among elemental signatures and broader, age-specific movement patterns. Our analyses show that otolith elemental signatures of Pacific herring can provide insight into complex population structure at scales of 10s – 1000 kms to inform and enhance spatially-explicit approaches to conservation and management

Pacific Herring Spawns Provide Temporal Subsidies to Nearshore Ecosystems

Pacific herring (Clupea pallasi), low trophic level fish at the base of many pelagic food webs, are integral to coastal social-ecological systems in North America. This migratory species provides a temporal subsidy to coastal systems by moving from offshore to nearshore waters for annual spawning events. Spawning migrations support subsistence and commercial fisheries, and provide an ecological pulse in prey availability. In coastal British Columbia, we quantified changes in rockfish (Sebastes maliger and caurinus) diet composition surrounding spawn events to understand the temporal importance of this subsidy. We found that the percentage of fish in rockfish diets switched from 30% fish tissue pre spawn to 34% herring roe post spawn, and that this shift peaked at 2-3 weeks post spawn events. This dietary change is most pronounced in females, where herring roe accounted for an average of 29% of stomach contents through the month following a spawn event. This may be due to differences in energy requirements for gravid females. Stable isotope concentrations of fish tissues (muscle, heart, liver, gonads) also showed evidence of herring nutrient assimilation. Determining these ecosystem-level associations is key to understanding both the social and ecological consequences of herring spawn declines

Hydrodynamic regime determines the magnitude of surface sediment \u27blue carbon\u27 stocks in British Columbia eelgrass meadows

Carbon storage in marine vegetated habitats (blue carbon) is increasingly being considered in carbon financing and coastal ecosystem management. Amongst other ecosystem functions, seagrass meadows are reported to be highly efficient at sequestering and storing significant amounts of carbon. However, seagrass blue carbon data remain sparse and regionally biased to tropical regions. In the Pacific Northwest, we lack information on the magnitude and variability of carbon stocks, as well as local drivers of variability. We collected sediment cores from six eelgrass meadows on the central coast of British Columbia, Canada, to quantify sedimentary organic carbon (Corg) stocks and accumulation rates. Carbon stocks exhibited 10-fold variability (335.35 – 3664. 48 g Corg m-2) in the top 20cm of sediment; these values align with other temperate eelgrass meadows, but are lower than reported global seagrass values. On average, Corg stocks within seagrass beds and adjacent unvegetated habitat overlapped, although stocks in the meadow interior (1392.05 g Corg m-2) were greater than those along meadow edges (1129.82 g Corg m-2) and in adjacent unvegetated sediments (977.10 g Corg m-2). Corg accumulation rates ranged from 12.57 to 50.45 g Corg m-2 year-1, lower than the global average reported for seagrasses (138 g Corg m-2 year-1), but again similar to other eelgrass meadows. Further, Corg in sediments beneath eelgrass meadows appears to be largely from non-eelgrass sources (terrestrial, benthic microalgae and macroalgae). Generalized linear mixed effects models suggest that hydrodynamic regime is the strongest driver of carbon stocks in the top 5cm of sediment, more important than the structural complexity of seagrass beds, or the proportion of fine sediments. Lower water velocities may allow greater deposition of particles within meadows, together with reduced erosion and resuspension. These results support physical characteristics over seagrass features as primary determinants of blue carbon storage in nearshore soft sediment habitats

Developing an Introductory UAV/Drone Mapping Training Program for Seagrass Monitoring and Research

Unoccupied Aerial Vehicles (UAVs), or drone technologies, with their high spatial resolution, temporal flexibility, and ability to repeat photogrammetry, afford a significant advancement in other remote sensing approaches for coastal mapping, habitat monitoring, and environmental management. However, geographical drone mapping and in situ fieldwork often come with a steep learning curve requiring a background in drone operations, Geographic Information Systems (GIS), remote sensing and related analytical techniques. Such a learning curve can be an obstacle for field implementation for researchers, community organizations and citizen scientists wishing to include introductory drone operations into their work. In this study, we develop a comprehensive drone training program for research partners and community members to use cost-effective, consumer-quality drones to engage in introductory drone mapping of coastal seagrass monitoring sites along the west coast of North America. As a first step toward a longer-term Public Participation GIS process in the study area, the training program includes lessons for beginner drone users related to flying drones, autonomous route planning and mapping, field safety, GIS analysis, image correction and processing, and Federal Aviation Administration (FAA) certification and regulations. Training our research partners and students, who are in most cases novice users, is the first step in a larger process to increase participation in a broader project for seagrass monitoring in our case study. While our training program originated in the United States, we discuss our experiences for research partners and communities around the globe to become more confident in introductory drone operations for basic science. In particular, our work targets novice users without a strong background in geographic research or remote sensing. Such training provides technical guidance on the implementation of a drone mapping program for coastal research, and synthesizes our approaches to provide broad guidance for using drones in support of a developing Public Participation GIS process

North to south: ecosystem features determine seagrass community response to sea otter foraging

We compared sea otter recovery in California (CA) and British Columbia (BC) to determine how key ecosystem properties shape top-down effects in seagrass communities. Potential ecosystem drivers of sea otter foraging in CA and BC seagrass beds that we examined include the role of coastline complexity and environmental stress on sea otter effects. In BC, we found greater species richness across seagrass trophic assemblages. Furthermore, Cancer spp. crabs, an important link in the seagrass trophic cascade observed in CA, was less common. Additionally, the more recent reintroduction of sea otters, more complex coastline, and reduced environmental stress in BC seagrass habitats supported the hypothesis that sea otter foraging pressure is currently reduced in more northern latitudes. In order to manage the ecosystem features that lead to regional differences in top predator effects in seagrass communities, we review our findings, their spatial and temporal constraints, and present a social-ecological framework for future re- search

Deeper habitats and cooler temperatures moderate a climate-driven seagrass disease

Eelgrass creates critical coastal habitats worldwide and fulfills essential ecosystem functions as a foundation seagrass. Climate warming and disease threaten eelgrass, causing mass mortalities and cascading ecological impacts. Subtidal meadows are deeper than intertidal and may also provide refuge from the temperature-sensitive seagrass wasting disease. From cross-boundary surveys of 5761 eelgrass leaves from Alaska to Washington and assisted with a machine-language algorithm, we measured outbreak conditions. Across summers 2017 and 2018, disease prevalence was 16% lower for subtidal than intertidal leaves; in both tidal zones, disease risk was lower for plants in cooler conditions. Even in subtidal meadows, which are more environmentally stable and sheltered from temperature and other stressors common for intertidal eelgrass, we observed high disease levels, with half of the sites exceeding 50% prevalence. Models predicted reduced disease prevalence and severity under cooler conditions, confirming a strong interaction between disease and temperature. At both tidal zones, prevalence was lower in more dense eelgrass meadows, suggesting disease is suppressed in healthy, higher density meadows. These results underscore the value of subtidal eelgrass and meadows in cooler locations as refugia, indicate that cooling can suppress disease, and have implications for eelgrass conservation and management under future climate change scenarios

A review of the opportunities and challenges for using remote sensing for management of surface-canopy forming kelps

© The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Cavanaugh, K. C., Bell, T., Costa, M., Eddy, N. E., Gendall, L., Gleason, M. G., Hessing-Lewis, M., Martone, R., McPherson, M., Pontier, O., Reshitnyk, L., Beas-Luna, R., Carr, M., Caselle, J. E., Cavanaugh, K. C., Miller, R. F., Hamilton, S., Heady, W. N., Hirsh, H. K., Hohman R., Lee L. C., Lorda J., Ray J., Reed D. C., Saccomanno V. R., Schroeder, S. B. A review of the opportunities and challenges for using remote sensing for management of surface-canopy forming kelps. Frontiers in Marine Science, 8, (2021): 753531, https://doi.org/10.3389/fmars.2021.753531.Surface-canopy forming kelps provide the foundation for ecosystems that are ecologically, culturally, and economically important. However, these kelp forests are naturally dynamic systems that are also threatened by a range of global and local pressures. As a result, there is a need for tools that enable managers to reliably track changes in their distribution, abundance, and health in a timely manner. Remote sensing data availability has increased dramatically in recent years and this data represents a valuable tool for monitoring surface-canopy forming kelps. However, the choice of remote sensing data and analytic approach must be properly matched to management objectives and tailored to the physical and biological characteristics of the region of interest. This review identifies remote sensing datasets and analyses best suited to address different management needs and environmental settings using case studies from the west coast of North America. We highlight the importance of integrating different datasets and approaches to facilitate comparisons across regions and promote coordination of management strategies.Funding was provided by the Nature Conservancy (Grant No. 02042019-5719), the U.S. National Science Foundation (Grant No. OCE 1831937), and the U.S. Department of Energy ARPA-E (Grant No. DE-AR0000922)

Low-Altitude UAV Imaging Accurately Quantifies Eelgrass Wasting Disease From Alaska to California

Declines in eelgrass, an important and widespread coastal habitat, are associated with wasting disease in recent outbreaks on the Pacific coast of North America. This study presents a novel method for mapping and predicting wasting disease using Unoccupied Aerial Vehicle (UAV) with low-altitude autonomous imaging of visible bands. We conducted UAV mapping and sampling in intertidal eelgrass beds across multiple sites in Alaska, British Columbia, and California. We designed and implemented a UAV low-altitude mapping protocol to detect disease prevalence and validated against in situ results. Our analysis revealed that green leaf area index derived from UAV imagery was a strong and significant (inverse) predictor of spatial distribution and severity of wasting disease measured on the ground, especially for regions with extensive disease infection. This study highlights a novel, efficient, and portable method to investigate seagrass disease at landscape scales across geographic regions and conditions