Community dynamics of microbial eukaryotes in intertidal mudflats in the hypertidal Bay of Fundy.

Protists (microbial eukaryotes) are a critically important but understudied group of microorganisms. They are ubiquitous, represent most of the genetic and functional diversity among eukaryotes, and play essential roles in nutrient and energy cycling. Yet, protists remain a black box in marine sedimentary ecosystems like the intertidal mudflats in the Bay of Fundy. The harsh conditions of the intertidal zone and high energy nature of tides in the Bay of Fundy provide an ideal system for gaining insights into the major food web players, diversity patterns and potential structuring influences of protist communities. Our 18S rDNA metabarcoding study quantified seasonal variations and vertical stratification of protist communities in Bay of Fundy mudflat sediments. Three 'SAR' lineages were consistently dominant (in terms of abundance, richness, and prevalence), drove overall community dynamics and formed the core microbiome in sediments. They are Cercozoa (specifically thecate, benthic gliding forms), Bacillariophyta (mainly cosmopolitan, typically planktonic diatoms), and Dinophyceae (dominated by a toxigenic, bloom-forming species). Consumers were the dominant trophic functional group and were comprised mostly of eukaryvorous and bacterivorous Cercozoa, and omnivorous Ciliophora, while phototrophs were dominated by Bacillariophyta. The codominance of Apicomplexa (invertebrate parasites) and Syndiniales (protist parasites) in parasite assemblages, coupled with broader diversity patterns, highlighted the combined marine and terrestrial influences on microbial communities inhabiting intertidal sediments. Our findings, the most comprehensive in a hypertidal benthic system, suggest that synergistic interactions of both local and regional processes (notably benthic-pelagic coupling) may drive heterogenous microbial distribution in high-energy coastal systems.

Stochastic dispersal increases the rate of upstream spread: A case study with green crabs on the northwest Atlantic coast.

Dispersal heterogeneity is an important process that can compensate for downstream advection, enabling aquatic organisms to persist or spread upstream. Our main focus was the effect of year-to-year variation in larval dispersal on invasion spread rate. We used the green crab, Carcinus maenas, as a case study. This species was first introduced over 200 years ago to the east coast of North America, and once established has maintained a relatively consistent spread rate against the dominant current. We used a stage-structured, integro-difference equation model that couples a demographic matrix for population growth and dispersal kernels for spread of individuals within a season. The kernel describing larval dispersal, the main dispersive stage, was mechanistically modeled to include both drift and settlement rate components. It was parameterized using a 3-dimensional hydrodynamic model of the Gulf of St Lawrence, which enabled us to incorporate larval behavior, namely vertical swimming. Dispersal heterogeneity was modeled at two temporal scales: within the larval period (months) and over the adult lifespan (years). The kernel models variation within the larval period. To model the variation among years, we allowed the kernel parameters to vary by year. Results indicated that when dispersal parameters vary with time, knowledge of the time-averaged dispersal process is insufficient for determining the upstream spread rate of the population. Rather upstream spread is possible over a number of years when incorporating the yearly variation, even when there are only a few "good years" featured by some upstream dispersal among many "bad years" featured by only downstream dispersal. Accounting for annual variations in dispersal in population models is important to enhance understanding of spatial dynamics and population spread rates. Our developed model also provides a good platform to link the modeling of larval behavior and demography to large-scale hydrodynamic models.

Relative Importance of Biotic and Abiotic Forces on the Composition and Dynamics of a Soft-Sediment Intertidal Community.

Top-down, bottom-up, middle-out and abiotic factors are usually viewed as main forces structuring biological communities, although assessment of their relative importance, in a single study, is rarely done. We quantified, using multivariate methods, associations between abiotic and biotic (top-down, bottom-up and middle-out) variables and infaunal population/community variation on intertidal mudflats in the Bay of Fundy, Canada, over two years. Our analysis indicated that spatial structural factors like site and plot accounted for most of the community and population variation. Although we observed a significant relationship between the community/populations and the biotic and abiotic variables, most were of minor importance relative to the structural factors. We suggest that community and population structure were relatively uncoupled from the structuring influences of biotic and abiotic factors in this system because of high concentrations of resources that sustain high densities of infauna and limit exploitative competition. Furthermore, we hypothesize that the infaunal community primarily reflects stochastic spatial events, namely a "first come, first served" process.

Behavioral response of Corophium volutator to shorebird predation in the upper Bay of Fundy, Canada.

Predator avoidance is an important component of predator-prey relationships and can affect prey availability for foraging animals. Each summer, the burrow-dwelling amphipod Corophium volutator is heavily preyed upon by Semipalmated Sandpipers (Calidris pusilla) on mudflats in the upper Bay of Fundy, Canada. We conducted three complementary studies to determine if adult C. volutator exhibit predator avoidance behavior in the presence of sandpipers. In a field experiment, we monitored vertical distribution of C. volutator adults in bird exclosures and adjacent control plots before sandpipers arrived and during their stopover. We also made polymer resin casts of C. volutator burrows in the field throughout the summer. Finally, we simulated shorebird pecking in a lab experiment and observed C. volutator behavior in their burrows. C. volutator adults were generally distributed deeper in the sediment later in the summer (after sandpipers arrived). In August, this response was detectably stronger in areas exposed to bird predation than in bird exclosures. During peak predator abundance, many C. volutator adults were beyond the reach of feeding sandpipers (>1.5 cm deep). However, burrow depth did not change significantly throughout the summer. Detailed behavioral observations indicated that C. volutator spent more time at the bottom of their burrow when exposed to a simulated predator compared to controls. This observed redistribution suggests that C. volutator adults move deeper into their burrows as an anti-predator response to the presence of sandpipers. This work has implications for predators that feed on burrow-dwelling invertebrates in soft-sediment ecosystems, as density may not accurately estimate prey availability.

Spatial variation in population structure and its relation to movement and the potential for dispersal in a model intertidal invertebrate.

Dispersal, the movement of an individual away from its natal or breeding ground, has been studied extensively in birds and mammals to understand the costs and benefits of movement behavior. Whether or not invertebrates disperse in response to such attributes as habitat quality or density of conspecifics remains uncertain, due in part to the difficulties in marking and recapturing invertebrates. In the upper Bay of Fundy, Canada, the intertidal amphipod Corophium volutator swims at night around the new or full moon. Furthermore, this species is regionally widespread across a large spatial scale with site-to-site variation in population structure. Such variation provides a backdrop against which biological determinants of dispersal can be investigated. We conducted a large-scale study at nine mudflats, and used swimmer density, sampled using stationary plankton nets, as a proxy for dispersing individuals. We also sampled mud residents using sediment cores over 3 sampling rounds (20-28 June, 10-17 July, 2-11 August 2010). Density of swimmers was most variable at the largest spatial scales, indicating important population-level variation. The smallest juveniles and large juveniles or small adults (particularly females) were consistently overrepresented as swimmers. Small juveniles swam at most times and locations, whereas swimming of young females decreased with increasing mud presence of young males, and swimming of large juveniles decreased with increasing mud presence of adults. Swimming in most stages increased with density of mud residents; however, proportionally less swimming occurred as total mud resident density increased. We suggest small juveniles move in search of C. volutator aggregations which possibly act as a proxy for better habitat. We also suggest large juveniles and small adults move if potential mates are limiting. Future studies can use sampling designs over large spatial scales with varying population structure to help understand the behavioral ecology of movement, and dispersal in invertebrate taxa.

Dispersal of marine benthic invertebrates through ice rafting.

Knowledge of dispersal vectors used by organisms is essential to the understanding of population and community dynamics. We report on ice rafting, a vector by which intertidal benthic invertebrates can be transported well outside their normal dispersal range during winter in temperate climates. We found multiple invertebrate taxa in sediment-laden ice blocks sampled in the intertidal zone. A large proportion of individuals were alive and active when freed from the ice. Using radio tracking, we found that ice blocks can travel over 20 km within a few days. Given the abundance of highly mobile ice blocks carrying viable invertebrates, we conclude that ice-rafting is likely an important dispersal vector, contributing to spatial community dynamics in intertidal systems. This mechanism helps explain observed genetic structure of populations, but it also raises concerns about potential negative impacts of climate change on connectivity between populations.

FORMATION AND PROPAGATION OF FEEDING FRONTS IN BENTHIC MARINE INVERTEBRATES: A MODELING APPROACH.

Feeding fronts are a striking pattern of spatial distribution observed in both marine and terrestrial ecological systems. These fronts not only determine the abundance and distribution of prey populations, but on a broader scale they may also affect the structure and dynamics of entire communities. Several mechanisms leading to the formation of feeding fronts have been proposed, and chemotaxis has been suggested as an important component. Here we develop two mathematical models that show front formation can occur with simple kinesis (and without chemotaxis) in two marine invertebrates with different feeding habits: a microphagous sea star (Oreaster reticulatus) that feeds on sediments and an herbivorous sea urchin (Strongylocentrotus droebachiensis) that grazes kelp beds. We utilize a large body of detailed empirical information on movement pattern, foraging behavior, and front dynamics for each species to develop, parameterize, and evaluate our models. We found that our model predictions for the rate of advance of a front and its relationship to the density of consumers at the front were in close agreement with independently collected, empirical observations in both systems. This work shows that simple local interactions between mobile consumers and a stationary resource can result in large-scale heterogeneous patterns of abundance of both species.