Drivers of plankton community structure in intermittent and continuous coastal upwelling systems–from microbes and microscale in-situ imaging to large scale patterns

Eastern Boundary Systems support major fisheries whose early life stages depend on upwelling production. Upwelling can be highly variable at the regional scale, with substantial repercussions for new productivity and microbial loop activity. Studies that integrate the classic trophic web based on new production with the microbial loop are rare due to the range in body forms and sizes of the taxa. Underwater imaging can overcome this limitation, and with machine learning, enables fine resolution studies spanning large spatial scales. We used the In-situ Ichthyoplankton Imaging System (ISIIS) to investigate the drivers of plankton community structure in the northern California Current, sampled along the Newport Hydrographic (NH) and Trinidad Head (TR) lines, in OR and CA, respectively. The non-invasive imaging of particles and plankton over 1644km in the winters and summers of 2018 and 2019 yielded 1.194 billion classified plankton images. Combining nutrient analysis, flow cytometry, and 16S rRNA gene sequencing of the microbial community with mesoplankton underwater imaging enabled us to study taxa from 0.2µm to 15cm, including prokaryotes, copepods, ichthyoplankton, and gelatinous forms. To assess community structure, >2000 single-taxon distribution profiles were analyzed using high resolution spatial correlations. Co-occurrences on the NH line were consistently significantly higher off-shelf while those at TR were highest on-shelf. Random Forests models identified the concentrations of microbial loop associated taxa such as protists, Oithona copepods, and appendicularians as important drivers of co-occurrences at NH line, while at TR, cumulative upwelling and chlorophyll a were of the highest importance. Our results indicate that the microbial loop is driving plankton community structure in intermittent upwelling systems such as the NH line and supports temporal stability, and further, that taxa such as protists, Oithona copepods, and appendicularians connect a diverse and functionally redundant microbial community to stable plankton community structure. Where upwelling is more continuous such as at TR, primary production may dominate patterns of community structure, obscuring the underlying role of the microbial loop. Future changes in upwelling strength are likely to disproportionately affect plankton community structure in continuous upwelling regions, while high microbial loop activity enhances community structure resilience

The Influence of a Ubiquitous Filter Feeder on Coastal Microbial Communities.

Doliolids have a unique ability to impact the marine microbial community through bloom events and high filtration rates. Their predation on large eukaryotic microorganisms is established and evidence of predation on smaller prokaryotic microorganisms is beginning to emerge. We studied the retention of both eukaryotic and prokaryotic microbial taxa by wild-caught doliolids in the northern California Current system. We use qPCR to quantify the impact of doliolids on three important and globally abundant taxa: Synechococcus, SAR11 and diatoms. Doliolids were collected during bloom events identified at three different shelf locations with variable upwelling intensities. We discovered that in addition to eukaryotic phytoplankton, doliolids feed on a range of prokaryotic microbial functional groups. Prey included pelagic Archaea, Pelagibacter, and picocyanobacteria, expanding our understanding of doliolid feeding to the smallest and most numerous microbial community members of the ocean. We also found that doliolids retain SAR11, which is intriguing because some SAR11 lineages may evade predation by other benthic and pelagic tunicates through their surface properties. Given the ability of doliolids to clear large portions of seawater by filtration and their high abundance in this system, we suggest that doliolids are an important player in shaping microbial community structure, primary production, and carbon fate in an ecologically and economically important fisheries system

Temperature Influences Selective Mortality during the Early Life Stages of a Coral Reef Fish

For organisms with complex life cycles, processes occurring at the interface between life stages can disproportionately impact survival and population dynamics. Temperature is an important factor influencing growth in poikilotherms, and growth-related processes are frequently correlated with survival. We examined the influence of water temperature on growth-related early life history traits (ELHTs) and differential mortality during the transition from larval to early juvenile stage in sixteen monthly cohorts of bicolor damselfish Stegastes partitus, sampled on reefs of the upper Florida Keys, USA over 6 years. Otolith analysis of settlers and juveniles coupled with environmental data revealed that mean near-reef water temperature explained a significant proportion of variation in pelagic larval duration (PLD), early larval growth, size-at-settlement, and growth during early juvenile life. Among all cohorts, surviving juveniles were consistently larger at settlement, but grew more slowly during the first 6 d post-settlement. For the other ELHTs, selective mortality varied seasonally: during winter and spring months, survivors exhibited faster larval growth and shorter PLDs, whereas during warmer summer months, selection on PLD reversed and selection on larval growth became non-linear. Our results demonstrate that temperature not only shapes growth-related traits, but can also influence the direction and intensity of selective mortality

Inter-Cohort Competition Drives Density Dependence and Selective Mortality in a Marine Fish

For organisms with complex life cycles, the transition between life stages and between habitats can act as a significant demographic and selective bottleneck. In particular, competition with older and larger conspecifics and heterospecifics may influence the number and characteristics of individuals successfully making the transition. We investigated whether the availability of enemy-free space mediated the interaction between adult goldspot gobies (Gnatholepis thompsoni), a. common tropical reef fish, and juvenile conspecifics that had recently settled from the plankton. We added rocks, which provide refuge from predators, to one-half of each of five entire coral reefs in the Bahamas and measured the survival and growth of recent settlers in relation to adult goby densities. We also evaluated whether mortality was selective with respect to three larval traits (age at settlement, size at settlement, and presettlement growth rate) and measured the influence of refuge availability and adult goby density on selection intensity. Selective mortality was measured by comparing larval traits of newly settled gobies (≤ 5 postsettlement) with those of survivors (2-3 week postsettlement juveniles). We detected a negative relationship between juvenile survival and adult goby density in both low- and high-refuge habitats, though experimental refuge addition reduced the intensity of this density dependence. Juvenile growth also declined with increasing adult goby density, but this effect was similar in both low- and high-refuge habitats. Refuge availability had no consistent effect on selective mortality, but adult goby density was significantly related to the intensity of size-selective mortality: bigger juveniles were favored where adults were abundant, and smaller juveniles were favored where adults were rare. Given the typically large difference in sizes of juveniles and adults, similar stage-structured interactions may be common but underappreciated in many marine species

Larval dispersal in a changing ocean with an emphasis on upwelling regions

Dispersal of benthic species in the sea is mediated primarily through small, vulnerable larvae that must survive minutes to months as members of the plankton community while being transported by strong, dynamic currents. As climate change alters ocean conditions, the dispersal of these larvae will be affected, with pervasive ecological and evolutionary consequences. We review the impacts of oceanic changes on larval transport, physiology, and behavior. We then discuss the implications for population connectivity and recruitment and evaluate life history strategies that will affect susceptibility to the effects of climate change on their dispersal patterns, with implications for understanding selective regimes in a future ocean. We find that physical oceanographic changes will impact dispersal by transporting larvae in different directions or inhibiting their movements while changing environmental factors, such as temperature, pH, salinity, oxygen, ultraviolet radiation, and turbidity, will affect the survival of larvae and alter their behavior. Reduced dispersal distance may make local adaptation more likely in well-connected populations with high genetic variation while reduced dispersal success will lower recruitment with implications for fishery stocks. Increased dispersal may spur adaptation by increasing genetic diversity among previously disconnected populations as well as increasing the likelihood of range expansions. We hypothesize that species with planktotrophic (feeding), calcifying, or weakly swimming larvae with specialized adult habitats will be most affected by climate change. We also propose that the adaptive value of retentive larval behaviors may decrease where transport trajectories follow changing climate envelopes and increase where transport trajectories drive larvae toward increasingly unsuitable conditions. Our holistic framework, combined with knowledge of regional ocean conditions and larval traits, can be used to produce powerful predictions of expected impacts on larval dispersal as well as the consequences for connectivity, range expansion, or recruitment. Based on our findings, we recommend that future studies take a holistic view of dispersal incorporating biological and oceanographic impacts of climate change rather than solely focusing on oceanography or physiology. Genetic and paleontological techniques can be used to examine evolutionary impacts of altered dispersal in a future ocean, while museum collections and expedition records can inform modern-day range shifts