Nonmetric multidimensional scaling (NMDS) is a powerful statistical tool which enables complex multivariate data sets to be visualized in a reduced number of dimensions. Users typically evaluate the fit of an NMDS ordination via ordination “stress” (i.e., data distortion) against a commonly accepted set of heuristic guidelines. However, these guidelines do not account for the mathematical relationship which links ordination stress to sample size. Consequently, researchers working with large data sets may unnecessarily present ordinations in an intractable number of dimensions, subdivide their data, or forego the use of NMDS entirely and lose the benefits of this highly flexible and useful technique. In order to overcome the limitations of these practices, we advocate for an alternative approach to the evaluation of NMDS ordination fit via the usage of permutation‐based ecological null models. We present the rationale for this approach from a theoretical basis, supported by a brief literature review, and an example usage of the methodology. Our literature review shows that NMDS analyses often far exceed the number of observations under which the original stress guidelines were formulated—with a significant increasing trend in recent decades. Adoption of a permutation‐based approach will consequently provide a more flexible and quantitative evaluation of NMDS fit and allow for the continued application of NMDS in an era of increasingly large datasets.

Nonindigenous aquatic species are becoming increasingly common in coastal and inland waters, largely due to the global transport of zooplankton via commercial shipping and recreational boating. The cost of mitigation and lost income due to invasive zooplankton is estimated in the billions of dollars annually, yet we know little about the temporal dynamics of these invaders. Analysis of an 8.5-year (June 2005-December 2013) zooplankton time series from the Columbia River revealed contrasting patterns of invasion dynamics between species, cyclical periods of community invasion, and key environmental variables that constrain populations of invasive zooplankton. We identified four seasonal zooplankton communities (autumn/ invaded, winter/barren, spring/rotifer, and transitional) that are strongly correlated with changes in chlorophyll content and water temperature, with peak abundances of invasive zooplankters occurring during periods of maximum water temperature. Additionally, we observed contrasting patterns of phenology between persistent and ephemeral invasive populations, with successful invaders exhibiting delayed annual peaks in population abundance. Two invasive zooplankters-the copepod Pseudodiaptomus forbesi and larval Asian clam Corbicula fluminea-dominate the zooplankton community in late summer and early autumn. Likewise, our results support conclusions from a growing body of literature that delayed phenology may be a key functional trait for successful invaders.

We present the first comprehensive analysis of the Pacific Northwest estuaries (PNWE) zooplankton time series, which encompasses 38 estuaries distributed across more than 1000 km of the North American Pacific Coast. With observations spanning more than 20 yr, we here examine biogeographic trends among zooplankton communities, patterns of biological invasion across the region, and environmental correlates with dominant native and invasive taxa. Our results show that some estuaries across the region are invaded by multiple zooplankton species and that the geographic extent of invasion is far greater than previously reported for at least five species of copepods: Pseudodiaptomus inopinus, Pseudodiaptomus forbesi, Oithona davisae, Limnoithona sinensis, Sinocalanus doerrii, and the cladoceran Bosmina coregoni. Some of these species appear to be rapidly spreading across the region, while others have occupied a relatively static geographic range for decades. The copepod, P. inopinus, is by far the most abundant and geographically widespread of these invaders, comprising more than 90% of all zooplankton abundance at some sites. We propose that the geographic distribution of these invaders is strongly constrained by geomorphic characteristics that define the salinity and mixing regimes in these estuaries, reflecting the strong role that physical forces play in structuring estuarine zooplankton communities.

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Although parasite-mediated selection is a major driver of host evolution, its influence on genetic variation for parasite resistance is not yet well understood. We monitored resistance in a large population of the planktonic crustacean Daphnia magna over 8 years, as it underwent yearly epidemics of the bacterial pathogen Pasteuria ramosa . We observed cyclic dynamics of resistance: Resistance increased throughout the epidemics, but susceptibility was restored each spring when hosts hatched from sexual resting stages. Host resting stages collected across the year showed that largely resistant host populations can produce susceptible sexual offspring. A genetic model of resistance developed for this host-parasite system, based on multiple loci and strong epistasis, is in partial agreement with our findings. Our results reveal that, despite strong selection for resistance in a natural host population, genetic slippage after sexual reproduction can be a strong factor for the maintenance of genetic diversity of host resistance.

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.

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.

Large river estuaries experience multiple anthropogenic stressors. Understanding plankton community dynamics in these estuaries provides insights into the patterns of natural variability and effects of human activity. We undertook a 2-year study in the Columbia River Estuary to assess the potential impacts of abiotic and biotic factors on planktonic community structure over multiple time scales. We measured microplankton and zooplankton abundance, biomass and composition monthly, concurrent with measurements of chlorophyll a, nutrient concentrations, temperature and salinity, from a dock in the lower estuary. We then statistically assessed the associations among the abundances of planktonic groups and environmental and biological factors. During the late spring high flow period of both years, the lower estuary was dominated by freshwater and low salinity-adapted planktonic taxa, and zooplankton grazers were more strongly associated with the autotroph-dominated microplankton assemblage than abiotic factors. During the early winter period of higher salinity and lower flow, nutrient (P) availability exerted a strong influence on microplankton taxa, while only temperature and upwelling strength were associated with the zooplankton assemblage. Our results indicate that the relative influence of biotic (grazers) and abiotic (salinity, flow, nutrients and upwelling) factors varies seasonally and inter-annually, and among different size classes in the estuarine food web.