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1. Kelp-associated variability in seawater chemistry during a marine heatwave event connects to transgenerational effects in the purple urchin Strongylocentrotus purpuratus

   https://doi.org/10.3354/meps14523  

   Kozal, LC; Nelson, JC; Hofmann, GE (April 2024, Marine Ecology Progress Series)

   Giant kelpMacrocystis pyriferaprovides the foundation for immense biodiversity on the coast of California, USA. Kelp forests can change seawater retention time, altering water chemistry, including pH and dissolved oxygen (DO), as well as the magnitude and predictability of variability in the same properties. Environmental heterogeneity across space and time could drive organismal performance and processes such as transgenerational plasticity (TGP), where parental experience modifies the offspring phenotype, potentially conferring tolerance to future environmental stress. We monitored environmental variability by deploying temperature, pH, and DO sensors inside and outside a temperate kelp forest in the Santa Barbara Channel (SBC) throughout the gametogenesis period of a key herbivore, the purple urchinStrongylocentrotus purpuratus. Over the 6 mo period, pH and temperature were slightly elevated inside the kelp forest, accompanied by more predictable, low-frequency variability relative to outside. AdultS. purpuratuswere conditioned inside and outside the kelp spanning gametogenesis. The urchins were spawned and their larvae were raised under high (1053 µatm) and lowpCO₂(435 µatm) at 15°C in the laboratory to assess their physiological response to the maternal and developmental environments. Larvae raised under highpCO₂were more susceptible to acute thermal stress; however, within each larval treatment, progeny from outside-conditioned mothers had a 0.4°C higher lethal temperature (LT₅₀). Our results indicate that heterogeneity in abiotic factors associated with kelp can have transgenerational effects in the field, and interactions between factors, including temperature and pH, will impact purple urchins as local variability associated with marine heatwaves and upwelling evolves with climate change. 

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2. Drivers of kelp forest refugia during successive disturbance events

   https://doi.org/10.1111/1365-2745.70072  

   Cavanaugh, Katherine C; Weiss, Robert E; Bell, Tom W; Rogers‐Bennett, Laura; Cavanaugh, Kyle C (August 2025, Journal of Ecology)

   Abstract Increased ocean temperatures have led to large‐scale declines in many ecologically important species, including kelp forests. Spatial heterogeneity across seascapes could protect kelp individuals and small populations from thermal stress and nutrient limitation. Habitat features within upwelling regions may facilitate the transport of deep, cold water into shallow systems, but little is known about the spatiotemporal occurrence or stability of these climate refugia. Kelp in climate refugia may, however, also experience other stressors, such as overgrazing by kelp herbivores, reducing their effectiveness.Here, we use high‐resolution kelp canopy maps generated from CubeSat constellation data to characterize kelp persistence in northern California following a dramatic decline in kelp abundance due to increased temperature and nutrient limitation during a severe marine heatwave and continued intense grazing pressure by purple sea urchins.Kelp persistence was associated with local areas of relatively cool water temperature and seascape features such as shallow depths and low‐complexity bathymetry, which may have provided refuge from overgrazing. However, a very small percentage of kelp forests in the region exhibited high persistence, with many forests present in only one or two of the 9 years studied. Most kelp patches were not spatially stable over time. Initially, kelp presence aligned with climate refugia, but as overgrazing emerged as the dominant driver of kelp distributions post‐2019, kelp shifted to areas that offered protection from grazing pressure.Synthesis. Cooler areas with localized upwelling acted as climate refugia during the increased ocean temperatures from the 2014–2016 marine heatwave, supporting nutrient‐rich environments and mitigating heat stress for kelp forests. However, these temperature refugia often did not spatially overlap with areas providing protection from grazing pressure, leaving kelp forests vulnerable to future warming even within temperature refugia if grazing pressure remains high. 

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3. Southward decrease in the protection of persistent giant kelp forests in the northeast Pacific

   https://doi.org/10.1038/s43247-021-00177-9  

   Arafeh-Dalmau, Nur; Cavanaugh, Kyle C.; Possingham, Hugh P.; Munguia-Vega, Adrian; Montaño-Moctezuma, Gabriela; Bell, Tom W.; Cavanaugh, Kate; Micheli, Fiorenza (December 2021, Communications Earth & Environment)

   null (Ed.)

   Abstract Kelp forests are globally important and highly productive ecosystems, yet their persistence and protection in the face of climate change and human activity are poorly known. Here, we present a 35-year time series of high-resolution satellite imagery that maps the distribution and persistence of giant kelp ( Macrocystis pyrifera ) forests along ten degrees of latitude in the Northeast Pacific Ocean. We find that although 7.7% of giant kelp is protected by marine reserves, when accounting for persistence only 4% of kelp is present and protected. Protection of giant kelp decreases southerly from 20.9% in Central California, USA, to less than 1% in Baja California, Mexico, which likely exacerbates kelp vulnerability to marine heatwaves in Baja California. We suggest that a two-fold increase in the area of kelp protected by marine reserves is needed to fully protect persistent kelp forests and that conservation of climate-refugia in Baja California should be a priority. 

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4. Limited biogeochemical modification of surface waters by kelp forest canopies: Influence of kelp metabolism and site‐specific hydrodynamics

   https://doi.org/10.1002/lno.11999  

   Traiger, Sarah B.; Cohn, Brian; Panos, Demetra; Daly, Margaret; Hirsh, Heidi K.; Martone, Maria; Gutierrez, Isabella; Mucciarone, David A.; Takeshita, Yuichiro; Monismith, Stephen G.; et al (December 2021, Limnology and Oceanography)

   Climate change is causing decreases in pH and dissolved oxygen (DO) in coastal ecosystems. Canopy-forming giant kelp can locally increase DO and pH through photosynthesis, with the most pronounced effect expected in surface waters where the bulk of kelp biomass resides. However, limited observations are available from waters in canopies and measurements at depth show limited potential of giant kelp to ameliorate chemical conditions. We quantified spatiotemporal variability of surface biogeochemistry and assessed the role of biological and physical drivers in pH and DO modification at two locations differing in hydrodynamics inside and outside of two kelp forests in Monterey Bay, California in summer 2019. pH, DO, dissolved inorganic carbon (DIC), and temperature were measured at and near the surface, in conjunction with physical parameters (currents and pressure), nutrients, and metrics of phytoplankton and kelp biological processes. DO and pH were highest, with lower DIC, at the surface inside kelp forests. However, differences inside vs. outside of kelp forests were small (DO 6–8%, pH 0.05 higher in kelp). The kelp forest with lower significant wave height and slower currents had greater modification of surface biogeochemistry as indicated by larger diel variation and slightly higher mean DO and pH, despite lower kelp growth rates. Differences between kelp forests and offshore areas were not driven by nutrients or phytoplankton. Although kelp had clear effects on biogeochemistry, which were modulated by hydrodynamics, the small magnitude and spatial extent of the effect limits the potential of kelp forests to mitigate acidification and hypoxia. 

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5. Persistence of southern California giant kelp beds and alongshore variation in nutrient exposure driven by seasonal upwelling and internal waves

   https://doi.org/10.3389/fmars.2023.1007789  

   Leichter, James J.; Ladah, Lydia B.; Parnell, P. Ed; Stokes, M. Dale; Costa, Matthew T.; Fumo, James; Dayton, Paul K. (March 2023, Frontiers in Marine Science)

   Kelp beds provide significant ecosystem services and socioeconomic benefits globally, and prominently in coastal zones of the California Current. Their distributions and abundance, however, vary greatly over space and time. Here, we describe long-term patterns of Giant Kelp (Macrocystis pyrifera) sea surface canopy area off the coast of San Diego County from 1983 through 2019 along with recent patterns of water column nitrate (NO₃^-) exposure inferred fromin situtemperature data in 2014 and 2015 at sites spanning 30 km of the coastline near San Diego California, USA. Site-specific patterns of kelp persistence and resilience were associated with ocean and climate dynamics, with total sea surface kelp canopy area varying approximately 33-fold over the almost 4 decades (min 0.34 km²in 1984; max 11.25 km²in 2008, median 4.79 km²). Site-normalized canopy areas showed that recent kelp persistence since 2014 was greater at Point Loma and La Jolla, the largest kelp beds off California, than at the much smaller kelp bed off Cardiff. NO₃^-exposure was estimated from an 11-month time series ofin situwater column temperature collected in 2014 and 2015 at 4 kelp beds, using a relationship between temperature and NO₃^-concentration previously established for the region. The vertical position of the 14.5°C isotherm, an indicator of the main thermocline and nutricline, varied across the entire water column at semidiurnal to seasonal frequencies. We use a novel means of quantifying estimated water column NO₃^-exposure integrated through time (mol-days m^-2) adapted from degree days approaches commonly used to characterize thermal exposures. Water column integrated NO₃^-exposure binned by quarters of the time series showed strong seasonal differences with highest exposure in Mar - May 2015, lowest exposure in Sep - Dec 2014, with consistently highest exposure off Point Loma. The water column integrated NO₃^-signal was filtered to provide estimates of the contribution to total nitrate exposure from high frequency variability (ƒ >= 1 cycle 30 hr^-1) associated predominantly with internal waves, and low frequency variability driven predominantly by seasonal upwelling. While seasonal upwelling accounted for > 90% of NO₃^-exposure across the full year, during warm periods when seasonal upwelling was reduced or absent and NO₃^-exposure was low overall, the proportion due to internal waves increased markedly to 84 to 100% of the site-specific total exposure. The high frequency variability associated with internal waves may supply critical nutrient availability during anomalously warm periods. Overall, these analyses support a hypothesis that differences in NO₃^-exposure among sites due to seasonal upwelling and higher frequency internal wave forcing contribute to spatial patterns in Giant Kelp persistence in southern California. The study period includes anomalously warm surface conditions and the marine heatwave associated with the “Pacific Warm Blob” superimposed on the seasonal thermal signal and corresponding to the onset of a multi-year decline in kelp canopy area and marked differences in kelp persistence among sites. Our analysis suggests that, particularly during periods of warm surface conditions, variation in NO₃^-exposure associated with processes occurring at higher frequencies, including internal waves can be a significant source of NO₃^-exposure to kelp beds in this region. The patterns described here also offer a view of the potential roles of seasonal and higher frequency nutrient dynamics for Giant Kelp persistence in southern California under continuing ocean surface warming and increasing frequency and intensity of marine heatwaves. 

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