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.

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.

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.

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.

Demographic connectivity is a fundamental process influencing the dynamics and persistence of spatially structured populations. Consequently, quantifying connectivity is essential for properly designing networks of protected areas so that they achieve their core ecological objective of maintaining population persistence. Recently, many empirical studies in marine systems have provided essential, and historically challenging to obtain, data on patterns of larval dispersal and export from marine protected areas (MPAs). Here, we review the empirical studies that have directly quantified the origins and destinations of individual larvae and assess those studies' relevance to the theory of population persistence and MPA design objectives. We found that empirical studies often do not measure or present quantities that are relevant to assessing population persistence, even though most studies were motivated or contextualized by MPA applications. Persistence of spatial populations, like nonspatial populations, depends on replacement, whether individuals reproduce enough in their lifetime to replace themselves. In spatial populations, one needs to account for the effect of larval dispersal on future recruitment back to the local population through local retention and other connectivity pathways. The most commonly reported descriptor of larval dispersal was the fraction of recruitment from local origin (self-recruitment). Self-recruitment does not inform persistence-based MPA design because it is a fraction of those arriving, not a fraction of those leaving (local retention), so contains no information on replacement. Some studies presented connectivity matrices, which can inform assessments of persistence with additional knowledge of survival and fecundity after recruitment. Some studies collected data in addition to larval dispersal that could inform assessments of population persistence but which were not presented in that way. We describe how three pieces of empirical information are needed to fully describe population persistence in a network of MPAs: (1) lifetime fecundity, (2) the proportion of larvae that are locally retained (or the full connectivity matrix), and (3) survival rate after recruitment. We conclude by linking theory and data to provide detailed guidance to empiricists and practitioners on field sampling design and data presentation that better informs the MPA objective of population persistence.