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Surfzone mixing and transport on a sandy, steep (∼1/8 slope), reflective beach at Carmel River State Beach, California, are described for a range of wave and alongshore flow conditions. Depth-limited wave breaking occurred close to the shore due to the steepness of the beach, creating a narrow surf/swash zone (∼10 m wide). Fluorescent Rhodamine dye was released as a slug in the surfzone, and the temporal and spatial evolution was measured using in-situ dye sensors. Dye concentration measured as a function of time reveals sharp fronts that quickly decay resulting in narrow peaks near the dye release, which subsequently broaden and decrease in peak concentration with alongshore distance. The measurements indicate two stages of mixing and transport occur inside the surfzone on the steep beach. 1) In the near-field (50 m downstream from the dye release location), the mass transport was dominated by advection. The distance to the far-field is much shorter in the alongshore on a steep beach compared with a dissipative beach. Estimates of cross-shore and alongshore diffusion coefficients (κ x , κ y ) were found to be similar in magnitude within the surfzone. Outside the surfzone in the far-field, the results suggest that the mixing processes are independent of those inside the surfzone. The mixing and transport of material observed on this steep beach are found to be analogous to that previously measured on dissipative beaches, however the diffusion coefficients within and outside the surfzone were found to be smaller on this steep beach.

Many marine benthic invertebrates have a pelagic life stage, during which larvae need to accumulate enough reserves to complete metamorphosis to a settled benthic juvenile. Currently, very little is known about how ocean conditions affect quality of the larvae. We studied this for three settlement seasons (2017–2019) by collecting returning Dungeness crab (Metacarcinus magister) megalopae at the Oregon coast and analyzing them for fatty acid biomarkers. We found that the larvae are omnivorous and have versatile diets. The daily larval abundance was positively correlated with larval quality. Despite the relatively high day‐to‐day variation, we found pronounced seasonal and inter‐annual differences in the body condition (size and total fatty acid content) and biomarker composition of megalopae. Especially, the early season recruits of 2017 had lower content of lipids and polyunsaturated fatty acids known to be beneficial to crustaceans. This is likely related to lingering effects of the eastern Pacific marine heat wave (2014–2016) on pelagic communities. The larvae were rich in docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA) with lower levels in benthic juveniles, indicating an ontogenetic diet shift and likely lower availability of DHA and EPA in the benthic environment. The pulsed megalopa recruitment may provide substantial carbon and lipid inputs to the nearshore ecosystem as a form of pelagic subsidy. Our results reveal that ocean conditions may have an effect on the quality of returning larvae, which likely influence their recruitment performance and early juvenile success, and thus potentially also the population size and commercial catch 4 yr later. [ABSTRACT FROM AUTHOR]

The Intermittent Upwelling Hypothesis (IUH) posits that subsidies of larvae and phytoplankton to intertidal communities should vary unimodally along a gradient of upwelling from persistent upwelling to persistent downwelling with most subsidies occurring where upwelling is of intermediate strength and intermittent. Furthermore, the hypothesis states that larvae and phytoplankton are transported far offshore by strong, persistent upwelling and fail to subsidize nearshore communities, whereas weak upwelling or downwelling reduces nutrients for phytoplankton production limiting food for larvae and nearshore communities. We review studies conducted at sea and onshore and reanalyze published data to test the IUH and evaluate alternative hypotheses. To test the hypothesis, we examine five predictions that must hold if the IUH is true. (1) Larvae should inhabit the surface Ekman layer where they are transported offshore during upwelling. Larvae of many intertidal taxa occur deeper in the water column where currents flow shoreward during upwelling. (2) Larvae of nearshore species should occur farther offshore during upwelling than during relaxation or downwelling. Larvae of many nearshore species remain within several kilometers of shore during both conditions. (3) Larval settlement in intertidal communities should be lower during upwelling than relaxation or downwelling. Daily larval settlement has not observed to be higher during relaxation or downwelling events; settlement has most often been seen to vary with the fortnightly tidal cycle likely due to onshore larval transport by internal tides. (4) Larval settlement and recruitment in intertidal communities should be lower in areas of strong, persistent upwelling than where upwelling is weaker and less persistent. Recruitment of mussels and barnacles to artificial and natural substrates did not vary with the strength of upwelling, but did vary inversely with two measures of desiccation potential, and directly with indicators of surf zone hydrodynamics; larval recruitment was higher where surf zones were more dissipative with rip currents. (5) Phytoplankton subsidies to nearshore communities should be highest where upwelling is moderate and intermittent. Like larval subsidies, phytoplankton subsidies varied spatially with surf zone hydrodynamics rather than upwelling. This reconsideration of the evidence for the IUH finds the hypothesis unsupported.

Surf zones, regions of breaking waves, are at the interface between the shore and coastal ocean. Surf zone hydrodynamics may affect delivery of phytoplankton subsidies to the intertidal zone. Over a month of daily sampling at an intermediate surf zone with bathymetric rip currents and a reflective surf zone, we measured surf zone hydrodynamics and compared concentrations of coastal phytoplankton taxa in the surf zones to concentrations offshore. At the intermediate surf zone, ~80% of the variability in the concentration of coastal phytoplankton taxa within the surf zone was explained by their variation offshore; however, concentrations were much higher and lower than those offshore in samples from a bathymetric rip current and over the adjacent shoal, respectively. Hydrodynamics at this intermediate surf zone did not hinder the delivery of coastal phytoplankton to the surf zone, but the bathymetric rip current system appeared to redistribute phytoplankton concentrating them within eddies. At the reflective shore, we sampled surf zones at a beach and two adjacent rocky intertidal sites. Concentrations of typical coastal phytoplankton taxa were usually an order of magnitude or more lower than those offshore, even when offshore samples were collected just 20 m beyond the breakers. The phytoplankton assemblages inside and outside the surf zone often appeared to be disconnected. Surf zone hydrodynamics at the steep, reflective shore coupled with low phytoplankton concentrations in near-surface water appeared to limit delivery of phytoplankton subsidies to the surf zone. Surf zone hydrodynamics may be a key factor in the alongshore variation in phytoplankton subsidies to coastal communities.

We investigated whether cross-shore distributions of coastal phytoplankton to the surf zone are controlled by hydrodynamics and their biological characteristics. Data from a rip-channeled beach indicate that concentrations of phytoplankton are higher in the surf zone than offshore. To examine how phytoplankton is transported toward the shore, we used a coupled biophysical model, comprised of a 3D physical model of coastal dynamics and an individual-based model (IBM) for tracking phytoplankton on the rip-channeled beach. Waves and wind in the biophysical model were parameterized by the conditions during the sampling period. Previous studies indicated that growth rates of phytoplankton can be enhanced by high turbulence, which might contribute to high phytoplankton concentration in the surf zone. Some numerical and laboratory works showed that turbulence can also increase the downward velocity of phytoplankton, which could be carried by onshore bottom currents and remain in the surf zone. Furthermore, we adapted the IBM with the theoretical model of diurnal vertical migration (DVM) for phytoplankton. The theoretical DVM works as follows: in the morning, phytoplankton cells adhere to air bubbles and stay at the surface and close to the shore in the daytime because onshore wind and surface current direction is usually onshore; in the late afternoon, the cells switch their attachment from air bubbles to sand grains and sink to the bottom where the water flow is normally onshore at night. Finally, depth-varying growth of phytoplankton was also incorporated into the DVM module. Simulations using neutral passive particles do not give the expected results of observed patterns. All tested mechanisms, i.e., wind- and wave-driven currents, rip-current circulation, turbulence-driven growth and sinking, DVM, and depth-varying growth, enhanced onshore phytoplankton migration and cell concentrations in the surf zone, indicating that both biological traits and physical factors can be essential to phytoplankton cross-shore transport and spatial variability. Our model is open to be modified and re-parameterized, followed by further analysis and validation, so that it can be more adequate for ecological assessment of coastal areas.

Plankton are transported onshore, providing subsidies of food and new recruits to surf-zone and intertidal communities. The transport of plankton to the surf zone is influenced by wind, wave, and tidal forcing, and whether they enter the surf zone depends on alongshore variation in surf-zone hydrodynamics caused by the interaction of breaking waves with coastal morphology. Areas with gently sloping shores and wide surf zones typically have orders-of-magnitude-higher concentrations of plankton in the surf zone and dense larval settlement in intertidal communities because of the presence of bathymetric rip currents, which are absent in areas with steep shores and narrow surf zones. These striking differences in subsidies have profound consequences; areas with greater subsidies support more productive surf-zone communities and possibly more productive rocky intertidal communities. Recognition of the importance of spatial subsidies for rocky community dynamics has recently advanced ecological theory, and incorporating surf-zone hydrodynamics would be an especially fruitful line of investigation.

Surf zone hydrodynamics vary along shorelines potentially affecting the delivery of larvae and zooplankton subsidies to intertidal communities, and, hence, the intensity of postsettlement interactions, growth and reproduction of filter-feeding foundation species and planktivorous fishes. We investigated the ability of zooplankton assemblages to enter the wide surf zone of the rip-channeled, more dissipative beach at Sand City, California, and the narrow surf zone of the steep reflective beach at nearby Carmel River State Beach. Every day for a month, we surveyed zooplankton inside and outside the surf zone and concomitant larval settlement of the dominant invertebrate onshore at each site in this upwelling regime. At the more dissipative surf zone, all zooplankters were far more concentrated inside than outside the surf zone. Many taxa increased in the surf zone and the predominant invertebrate on beaches, Emerita analoga, settled abundantly when prevailing northwesterly winds relaxed and waves were small. At the reflective surf zone, concentrations of zooplankters of most taxa were far greater outside than inside the surf zone, and many taxa increased in the surf zone when waves were small. Twice as many taxa were positively correlated inside and outside the surf zone at the dissipative than the reflective surf zone, indicating that zooplankters were more freely exchanged although behavior also played a role. Thus, spatial and temporal variation in surf zone hydrodynamics may regulate subsidies of zooplankton food and larval recruits to nearshore communities with potential cascading effects on community dynamics and structure.

Larvae in the coastal ocean are transported toward shore by a variety of mechanisms. Crossing the surf zone is the last step in a shoreward migration and surf zones may act as semipermeable barriers altering delivery of larvae to the shore. We related variation in the structure of intertidal barnacle populations to surf zone width (surf zone hydrodynamics proxy), wave height, alongshore wind stress (upwelling proxy), solar radiation, and latitude at 40 rocky intertidal sites from San Diego, California to the Olympic Peninsula, Washington. We measured daily settlement and weekly recruitment of barnacles at selected sites and related these measures to surf zone width. Chthamalus density varied inversely with that of Balanus, and the density of Balanus and new recruits was negatively related to solar radiation. Across the region, long-term mean wave height and an indicator of upwelling intensity and frequency did not explain variation in Balanus or new recruit densities. Balanus and new recruit densities, daily settlement, and weekly recruitment were up to three orders of magnitude higher at sites with wide (>50 m), more dissipative surf zones with bathymetric rip currents than at sites with narrow (

Larvae of intertidal species develop at sea and must return to adult habitats to replenish populations. Similarly, nutrients, detritus and plankton provide important subsidies spurring growth and reproduction of macroalgae and filter-feeding invertebrates that form the foundation of intertidal communities. Together, these factors determine the density and intensity of interactions among community members. We hypothesized that spatial variation in surfzone hydrodynamics affects the delivery of plankton subsidies. We compared entire zooplankton communities inside and outside the surf zone daily while monitoring physical conditions for one month each at two shores with different surfzone characteristics. Opposite cross-shore distributions of larvae and other zooplankters occurred at the two sites: zooplankton was much more abundant inside the mildly sloping dissipative surf zone (DSZ) with rip currents and was more abundant outside the steep reflective surf zone (RSZ). Biophysical numerical simulations demonstrated that zooplankters were concentrated in rip channels of the DSZ and were mostly unable to enter the RSZ, indicating the hydrodynamic processes behind the observed spatial variation of zooplankters in the surf zone. Differences in the concentration of larvae and other zooplankters between the inner shelf and surf zone may be an underappreciated, key determinant of spatial variation in inshore communities.

Most harmful algal blooms (HAB) originate away from the shore and, for them to endanger human health, they must be first transported to shore after which they must enter the surf zone where they can be feed upon by filter feeders. The last step in this sequence, entrance into the surf zone, depends on surf zone hydrodynamics. During two 30-day periods, we sampled Pseudo-nitzschia and particulate domoic acid (pDA) in and offshore of a more dissipative surf zone at Sand City, California (2010) and sampled Pseudo-nitzschia in and out of reflective surf zones at a beach and rocky shores at Carmel River State Beach, California (2011). At Sand City, we measured domoic acid in sand crabs, Emerita analoga. In the more dissipative surf zone, concentrations of Pseudo-nitzschia and pDA were an order of magnitude higher in samples from a rip current than in samples collected just seaward of the surf zone and were 1000 times more abundant than in samples from the shoals separating rip currents. Domoic acid was present in all the Emerita samples and varied directly with the concentration of pDA and Pseudo-nitzschia in the rip current. In the more reflective surf zones, Pseudo-nitzschia concentrations were 1-2 orders of magnitude lower than in samples from 125 and 20m from shore. Surf zone hydrodynamics affects the ingress of Pseudo-nitzschia into surf zones and the exposure of intertidal organisms to HABs on the inner shelf.

Competent larvae of intertidal invertebrates have to migrate toward shore for settlement; however, their migration through the surf zone is not understood. We investigated larval transport mechanisms at a rip-channeled beach. Because tracking larvae in the surf zone is infeasible, we used a three-dimensional biophysical model to simulate the processes. The coupled model consists of a physical module for currents and waves, and a biological module for adding larval traits and behaviors as well as Stokes drift to Lagrangian particles. Model calculations were performed with and without onshore wind forcing. Without wind, wave-driven onshore streaming occurs in the bottom boundary layer outside the surf zone. With onshore wind, onshore currents occur near the surface. In the surf zone, offshore-directed rip currents and compensating onshore-directed currents over shoals are formed in both no-wind and wind cases. In the biological module, neutral, negative, and positive buoyant particles were released offshore. Additionally, particles either sank in the presence of turbulence or not. Two scenarios achieved successful onshore migration: Negatively buoyant larvae without wind forcing sink in the turbulent bottom boundary layer and are carried onshore by streaming; positively buoyant larvae drift toward shore in wind-driven surface currents to the surf zone, then sink in the turbulent surf zone and remain near the bottom while transported shoreward. In both cases, the larval concentration is highest in the rip channel, consistent with field data. This successful result is only obtained if turbulence-dependent sinking behavior and Stokes drift are included in the transport of larvae. © 2014, by the Association for the Sciences of Limnology and Oceanography, Inc.

Larvae of intertidal species develop in the coastal ocean, and the last body of water they must cross while migrating back to shore is the surf zone. We hypothesized that the surf zone is a semipermeable barrier to this shoreward migration and that differences in water exchange across the surf zone result in temporal and spatial variation in larval delivery to the shore. We tested the hypotheses that larval delivery 1) should increase with larger waves and 2) should be higher on more dissipative beaches than on more reflective beaches. We found a significant positive correlation between the daily averaged ratio of wave height to wave period (H/. T) and daily cyprid settlement at Dike Rock, California and Bastendorff Beach, Oregon, USA. We tested the second hypothesis by comparing populations of barnacles, limpets, and benthic algae on rocks on four more dissipative and six more reflective sandy beaches in northern California and southern Oregon. Newly recruited barnacles and limpets were significantly more abundant at more dissipative than reflective beaches, and the higher abundance was most likely due to differences in settlement rather than post-settlement mortality. The density and percent cover of barnacles and the density of limpets were significantly higher at more dissipative beaches. In contrast, the density and percent cover of algae were significantly higher at more reflective beaches. The results are consistent with the hypothesis that the surf zone is a semipermeable barrier to the shoreward migration of larvae and that differences in water exchange across the surf zone as function of the beach hydrodynamics result in temporal and spatial variation in larval delivery to the shore. © 2010 Elsevier B.V.