Early detection monitoring for larval dreissenid mussels: how much plankton sampling is enough?

The development of quagga and zebra mussel (dreissenids) monitoring programs in the Pacific Northwest provides a unique opportunity to evaluate a regional invasive species detection effort early in its development. Recent studies suggest that the ecological and economic costs of a dreissenid infestation in the Pacific Northwest of the USA would be significant. Consequently, efforts are underway to monitor for the presence of dreissenids. However, assessments of whether these efforts provide for early detection are lacking. We use information collected from 2012 to 2014 to characterize the development of larval dreissenid monitoring programs in the states of Idaho, Montana, Oregon, and Washington in the context of introduction and establishment risk. We also estimate the effort needed for high-probability detection of rare planktonic taxa in four Columbia and Snake River reservoirs and assess whether the current level of effort provides for early detection. We found that the effort expended to monitor for dreissenid mussels increased substantially from 2012 to 2014, that efforts were distributed across risk categories ranging from high to very low, and that substantial gaps in our knowledge of both introduction and establishment risk exist. The estimated volume of filtered water required to fully census planktonic taxa or to provide high-probability detection of rare taxa was high for the four reservoirs examined. We conclude that the current level of effort expended does not provide for high-probability detection of larval dreissenids or other planktonic taxa when they are rare in these reservoirs. We discuss options to improve early detection capabilities.

Environmental influence on cyanobacteria abundance and microcystin toxin production in a shallow temperate lake.

The increasing frequency of harmful cyanobacterial blooms in freshwater systems is a commonly recognized problem due to detrimental effects on water quality. Vancouver Lake, a shallow, tidally influenced lake in the flood plain of the Columbia River within the city of Vancouver, WA, USA, has experienced numerous summertime cyanobacterial blooms, dominated by Aphanizomenon sp. and Anabaena sp. Cyanobacteria abundance and toxin (microcystin) levels have been monitored in this popular urban lake for several years; however, no previous studies have identified which cyanobacteria species produce toxins, nor analyzed how changes in environmental variables contribute to the fluctuations in toxic cyanobacteria populations. We used a suite of molecular techniques to analyze water samples from Vancouver Lake over two summer bloom cycles (2009 and 2010). Both intracellular and extracellular microcystin concentrations were measured using an ELISA kit. Intracellular microcystin concentrations exceeded WHO guidelines for recreational waters several times throughout the sampling period. PCR results demonstrated that Microcystis sp. was the sole microcystin-producing cyanobacteria species present in Vancouver Lake, although Microcystis sp. was rarely detected in microscopical counts. qPCR results indicated that the majority of the Microcystis sp. population contained the toxin-producing gene (mcyE), although Microcystis sp. abundance rarely exceeded 1 percent of overall cyanobacteria abundance. Non-metric multidimensional scaling (NMDS) revealed that PO4-P was the main environmental variable influencing the abundance of toxic and non-toxic cyanobacteria, as well as intracellular microcystin concentrations. Our study underscores the importance of using molecular genetic techniques, in addition to traditional microscopy, to assess the importance of less conspicuous species in the dynamics of harmful algal blooms.

The influence of water quality variables on cyanobacterial blooms and phytoplankton community composition in a shallow temperate lake.

Cyanobacterial blooms and their detrimental effects on water quality have become a worldwide problem. Vancouver Lake, a tidally influenced shallow temperate freshwater lake in Washington state, U.S.A., exhibits annual summer cyanobacterial blooms that are of concern to local resource managers. Our objectives were to describe changes in phytoplankton community composition in Vancouver Lake over seasonal, annual, and interannual time scales, and to identify strong water quality predictors of phytoplankton community structure, with an emphasis on cyanobacterial blooms, from 2007 through 2010. Cluster analysis, indicator species analysis, and non-metric multidimensional scaling were used to identify significantly different phytoplankton community groupings and to determine which environmental factors influenced community changes. From 2007 through 2009, depletion of NO3-N followed by elevated PO4-P concentration was associated with increased biomass and duration of each cyanobacterial bloom. Time-lag analysis suggested that NO3-N availability contributed to interannual changes within the summer phytoplankton community. Specifically, in summer 2010, a distinct cyanobacteria community was not present, potentially due to increased NO3-N and decreased PO4-P and NH4-N availability. Our study provides a comprehensive assessment of species-level responses to water quality variables in a shallow non-stratifying temperate lake, contributes to a better understanding of phytoplankton dynamics, and may aid in predicting and managing cyanobacterial blooms.

Zooplankton invasions in the early 21st century: a global survey of recent studies and recommendations for future research.

We present a comprehensive survey of the scientific literature pertaining to non-indigenous and invasive zooplankton published across the first decades of the twenty-first century (i.e., 2000-2018). We provide a concise summary of the manner in which the scientific community has allocated its efforts to this issue in recent decades, and to illuminate trends that emerge from the literature. Our search yielded 620 publications encompassing 139 invasive zooplankton species, with invasive zooplankton reported from every region of the planet-including the Arctic and Antarctic. Most taxa were reported in a small number of publications, with the majority being mentioned in only a single paper. In contrast, approximately half of the surveyed publications concerned just four species: Bythotrephes longimanus, Mnemioposis leidyi, Cercopagis pengoi, and Daphnia lumholtzi. Our survey reveals strong geographic patterns among the literature, with most publications arising from economically developed western nations. We found that the majority of publications pertained to holoplanktonic organisms from freshwater habitats, especially from the North American Great Lakes. Based on these results, we present several recommendations for future research topics that may hold considerable opportunity for growth in our understanding of the invasion process.

Modeling the trophic impacts of invasive zooplankton in a highly invaded river.

The lower Columbia River (Washington and Oregon, USA) has been heavily invaded by a large number of planktonic organisms including the invasive copepod Pseudodiaptomus forbesi and the planktonic juveniles of the invasive clam, Corbicula fluminea. In order to assess the ecological impacts of these highly abundant invaders, we developed a multivariate auto-regressive (MAR) model of food web dynamics based upon a 12-year time-series of plankton community and environmental data from the Columbia River. Our model results indicate that plankton communities in the lower Columbia River are strongly impacted by the copepod P. forbesi at multiple trophic levels. We observed different ecological effects across different life stages of P. forbesi, with nauplii negatively impacting ciliates and autotrophs, and copepodite stages negatively impacting Daphnia and cyclopoid copepods. Although juvenile C. fluminea were highly abundant in the summer and autumn of each year, our best fit MAR model did not show significant C. fluminea impacts. Our results illustrate the strong ecological impact that some zooplankton invaders may cause within rivers and estuarine systems, and highlight the need for further research on the feeding ecology of the planktonic life-stage of C. fluminea. Overall, our study demonstrates the manner in which long-term, high resolution data sets can be used to better understand the ecological impacts of invasive species among complex and highly dynamic communities.

Linking functional response and bioenergetics to estimate juvenile salmon growth in a reservoir food web.

Juvenile salmon (Oncorhynchus spp.) use of reservoir food webs is understudied. We examined the feeding behavior of subyearling Chinook salmon (O. tshawytscha) and its relation to growth by estimating the functional response of juvenile salmon to changes in the density of Daphnia, an important component of reservoir food webs. We then estimated salmon growth across a broad range of water temperatures and daily rations of two primary prey, Daphnia and juvenile American shad (Alosa sapidissima) using a bioenergetics model. Laboratory feeding experiments yielded a Type-II functional response curve: C = 29.858 P *(4.271 + P)-1 indicating that salmon consumption (C) of Daphnia was not affected until Daphnia densities (P) were < 30 · L-1. Past field studies documented Daphnia densities in lower Columbia River reservoirs of < 3 · L-1 in July but as high as 40 · L-1 in August. Bioenergetics modeling indicated that subyearlings could not achieve positive growth above 22°C regardless of prey type or consumption rate. When feeding on Daphnia, subyearlings could not achieve positive growth above 20°C (water temperatures they commonly encounter in the lower Columbia River during summer). At 16-18°C, subyearlings had to consume about 27,000 Daphnia · day-1 to achieve positive growth. However, when feeding on juvenile American shad, subyearlings had to consume 20 shad · day-1 at 16-18°C, or at least 25 shad · day-1 at 20°C to achieve positive growth. Using empirical consumption rates and water temperatures from summer 2013, subyearlings exhibited negative growth during July (-0.23 to -0.29 g · d-1) and August (-0.05 to -0.07 g · d-1). By switching prey from Daphnia to juvenile shad which have a higher energy density, subyearlings can partially compensate for the effects of higher water temperatures they experience in the lower Columbia River during summer. However, achieving positive growth as piscivores requires subyearlings to feed at higher consumption rates than they exhibited empirically. While our results indicate compromised growth in reservoir habitats, the long-term repercussions to salmon populations in the Columbia River Basin are unknown.

Predation on the Invasive Copepod, Pseudodiaptomus forbesi, and Native Zooplankton in the Lower Columbia River: An Experimental Approach to Quantify Differences in Prey-Specific Feeding Rates.

Invasive planktonic crustaceans have become a prominent feature of aquatic communities worldwide, yet their effects on food webs are not well known. The Asian calanoid copepod, Pseudodiaptomus forbesi, introduced to the Columbia River Estuary approximately 15 years ago, now dominates the late-summer zooplankton community, but its use by native aquatic predators is unknown. We investigated whether three species of planktivorous fishes (chinook salmon, three-spined stickleback, and northern pikeminnow) and one species of mysid exhibited higher feeding rates on native copepods and cladocerans relative to P. forbesi by conducting `single-prey' feeding experiments and, additionally, examined selectivity for prey types with `two-prey' feeding experiments. In single-prey experiments individual predator species showed no difference in feeding rates on native cyclopoid copepods (Cyclopidae spp.) relative to invasive P. forbesi, though wild-collected predators exhibited higher feeding rates on cyclopoids when considered in aggregate. In two-prey experiments, chinook salmon and northern pikeminnow both strongly selected native cladocerans (Daphnia retrocurva) over P. forbesi, and moreover, northern pikeminnow selected native Cyclopidae spp. over P. forbesi. On the other hand, in two-prey experiments, chinook salmon, three-spined stickleback and mysids were non- selective with respect to feeding on native cyclopoid copepods versus P. forbesi. Our results indicate that all four native predators in the Columbia River Estuary can consume the invasive copepod, P. forbesi, but that some predators select for native zooplankton over P. forbesi, most likely due to one (or both) of two possible underlying casual mechanisms: 1) differential taxon-specific prey motility and escape responses (calanoids > cyclopoids > daphnids) or 2) the invasive status of the zooplankton prey resulting in naivety, and thus lower feeding rates, of native predators feeding on invasive prey.