Addressing the misidentification of sporangia as “embryonic sporophytes” in <i>Nereocystis luetkeana</i> (Mertens) Postels & Ruprecht

Kelp life history pathways alternate between macroscopic sporophytes that produce spores and microscopic gametophytes that produce gametes. Occasionally, an alternative pathway is seen. This study examined the circumstances by which the high latitude estuarine bull kelp, Nereocystis luetkeana, foregoes the “free-living microscopic stages by releasing embryonic sporophytes directly from sori. Sori were collected from adult N. luetkeana sporophytes from eight locations within Kachemak Bay, Alaska in 2018 and 2020 to examine spatial and temporal development of embryonic sporophytes on sori. Distinctions were made between sori collected from first-generation and overwintered adults to assess the influence of parental age on embryonic sporophyte release. Further distinctions were made between sori collected from attached and drifting individuals to assess the influence of the status of parental attachment to substrate on embryonic sporophyte release. Inspection of propagules released from sori after 48-h incubations indicated that embryonic sporophytes were occasionally released alongside viable spores. Though embryonic sporophytes were released from sori as early as spring, it was not evident that they were bound by seasonal or spatial limits. The percent of propagules that were embryonic sporophytes ranged from 0% to 100% but were not significantly different between first-generation and overwintered adults, nor were they different between attached and drifting individuals. Nevertheless, the characteristic of directly releasing embryonic sporophytes from adult sporophytes might have ecological advantages for N. luetkeana. Key words: Alaska; alternative; bull kelp; embryonic sporophytes; life history; Nereocystis luetkeana

Abstract. A series of accidents at the Fukushima Dai-ichi Nuclear Power Plant following the Great East Japan Earthquake and tsunami of 11 March 2011 resulted in the release of radioactive materials to the ocean by two major pathways: direct release from the accident site and atmospheric deposition. A 1 yr, regional-scale simulation of 137Cs activity in the ocean offshore of Fukushima was carried out, the sources of radioactivity being direct release, atmospheric deposition, and the inflow of 137Cs deposited into the ocean by atmospheric deposition outside the domain of the model. Direct releases of 137Cs were estimated for 1 yr after the accident by comparing simulated results and measured activities adjacent to the accident site. The contributions of each source were estimated by analysis of 131I/137Cs and 134Cs/137Cs activity ratios and comparisons between simulated results and measured activities of 137Cs. The estimated total amounts of directly released 131I, 137Cs, and 137Cs were 11.1 ± 2.2 PBq, 3.5 ± 0.7 PBq, and 3.6 ± 0.7 PBq, respectively. Simulated 137Cs activities attributable to direct release were in good agreement with measured 137Cs activities not only adjacent to the accident site, but also in a wide area in the model domain, therefore this implies that the estimated direct release rate was reasonable. Employment of improved nudging data by JCOPE2 improved both the offshore transport result and the reproducibility of 137Cs activities 30 km offshore. On the other hand, simulated 137Cs activities attributable to atmospheric deposition were low compared to measured activities. The rate of atmospheric deposition into the ocean was underestimated because of a lack of measurements of deposition into the ocean when atmospheric deposition rates were being estimated. Simulated 137Cs activities attributable to the inflow of 137Cs deposited into the ocean outside the domain of the model were in good agreement with measured activities in the open ocean within the model domain after June 2012. The consideration of inflow is important to simulate the 137Cs activity in this model region in the later period of the simulation. The contribution of inflow increased with time and was dominant (more than 99%) by the end of February 2012. The activity of directly released 137Cs, however, decreased exponentially with time and was detectable only in the coastal zone by the end of February 2012.

Climate change is affecting marine ecosystems in many ways, including raising temperatures and leading to ocean acidification. From 2014 to 2016, an extensive marine heat wave extended along the west coast of North America and had devastating effects on numerous species, including bull kelp (Nereocystis luetkeana). Bull kelp is an important foundation species in coastal ecosystems and can be affected by marine heat waves and ocean acidification; however, the impacts have not been investigated on sensitive early life stages. To determine the effects of changing temperatures and carbonate levels on Northern California's bull kelp populations, we collected sporophylls from mature bull kelp individuals in Point Arena, CA. At the Bodega Marine Laboratory, we released spores from field-collected bull kelp, and cultured microscopic gametophytes in a common garden experiment with a fully factorial design crossing modern conditions (11.63 ± 0.54°C and pH 7.93 ± 0.26) with observed extreme climate conditions (15.56 ± 0.83°C and 7.64 ± 0.32 pH). Our results indicated that both increased temperature and decreased pH influenced growth and egg production of bull kelp microscopic stages. Increased temperature resulted in decreased gametophyte survival and offspring production. In contrast, decreased pH had less of an effect but resulted in increased gametophyte survival and offspring production. Additionally, increased temperature significantly impacted reproductive timing by causing female gametophytes to produce offspring earlier than under ambient temperature conditions. Our findings can inform better predictions of the impacts of climate change on coastal ecosystems and provide key insights into environmental dynamics regulating the bull kelp lifecycle.

A fundamental goal in ecology is to understand the drivers of stability in natural ecosystems in the face of disturbances. However, this is challenging when biotic and abiotic stressors operate simultaneously across multiple spatial scales. Such is the case for bull kelp forests (Nereocystis luetkeana) in northern California, where losses of predators combined with marine heatwaves have led to shifts from kelp forest to sea urchin barren states. However, despite the >90% loss of bull kelp forests since 2014, some patches remain. Here, we investigate the bull kelp community assemblage in these remnant patches as well as the drivers of bull kelp forest resistance. We used a combination of in situ field surveys (years 2020–2022), remote sensing data (years 2016–2022), and a laboratory grazing experiment with urchins (Strongylocentrotus purpuratus). We found that, in addition to the two dominant states (kelp forest vs. urchin barren), there is a third community state dominated by understory canopy‐forming macroalgae that stays subsurface. Moreover, bull kelp abundance and cover were positively associated with freshwater flow and proximity to freshwater sources, and bull kelp persistence was positively associated with sand cover, all of which seem to diminish sea urchin abundance and the negative effects of sea urchin herbivory on bull kelp. This was also shown in the laboratory experiment where sea urchin herbivory rates on bull kelp decreased with decreasing salinity. Overall, these results suggest that freshwater influence in shallow coastal environments could prevent loss of bull kelp and show that land–sea connections should be considered for species‐specific management and conservation actions.

The Northern kelp crab (Pugettia producta) and the graceful kelp crab (Pugettia gracilis) are common primary consumers in bull kelp beds near the San Juan Islands (Salish Sea, NE Pacific). In this system, urchins (often considered the most voracious herbivores exerting top-down control on kelp beds) tend to remain sedentary because of the high availability of detrital macroalgae, but the extent to which kelp crabs consume kelp (and other food options) is largely unknown. I conducted four types of laboratory feeding experiments to evaluate kelp crab feeding patterns: (1) feeding electivity between bull kelp (Nereocystis luetkeana) and seven species of co-occurring local macroalgae; (2) feeding electivity on aged vs. fresh bull kelp; (3) feeding preference between N. luetkeana and small snails (Lacuna sp.); and (4) scaling of feeding rate with body size in P. producta and P. gracilis. In choice experiments, P. producta consumed greater mass of N. luetkeana than of other macroalgal species offered and elected to eat fresh bull kelp over aged. However, P. producta also consumed snails (Lacuna sp.), indicating more generalized feeding than previously suspected. Feeding rates for P. producta exceeded the expected 3∕4 scaling rule of metabolic rates, indicating that larger P. producta may have a disproportionately large impact on bull kelp. A subtidal field experiment, designed to assess the influence of consumers on juvenile bull kelp net tissue gain, found that only fully enclosed (protected) bull kelp increased in wet mass and blade length. Herbivory by kelp crabs, among other consumers, is likely to play a previously unrecognized role in mediating the growth and survival of this annual kelp species within the Salish Sea.

Extending from central California to Alaska, bull kelp (Nereocystis luetkeana) forms seasonal kelp forests that are iconic coastal ecosystems in much of the eastern Pacific. Historical and ongoing field surveys and aerial imagery are used to provide biological data on kelp canopy cover and health, but satellite remote sensing provides the opportunity to generate consistent, long-term datasets over a large spatial scale. Robust satellite-based timeseries measurements of giant kelp (Macrocystis pyrifera) are available for much of the California coastline (e.g., Santa Barbara to Santa Cruz), but there have been no equivalent publications for bull kelp. Recent loss of bull kelp in northern California emphasized the need for more expansive long-term monitoring of canopy trends. We tested various kelp classification approaches using Multiple Endmember Spectral Mixture Analysis (MESMA) applied to Landsat imagery, which allowed sufficient temporal and spatial data collection during bull kelp's narrow seasonal maximum, and compare with the California Department of Fish and Wildlife (CDFW) aerial survey canopy area product. We addressed five main topics that have relevance to enabling Landsat in long-term monitoring of bull kelp canopy coverage in northern California: (1) the effect of MESMA configurations applied to Landsat imagery, including software dependencies and endmember configurations, (2) comparison of Landsat to traditional surveys, (3) differences across Landsat sensors, (4) tidal influence on canopy area, and (5) trends in the decadal timeseries. We found that there was no statistical difference (p = 0.53) between MESMA platforms (IDL-based and a Python-scripted method; RMSE 1.5 km2 or 17.6% when normalized by the range in CDFW), and that a 7-endmember MESMA model provided the lowest RMSE (1.4 km2 or 16.9%). Furthermore, while tidal phases can submerge or emerge kelp canopy and thus potentially affect kelp detection, we found a weak and statistically insignificant correlation between tides and performance of our remote estimate of kelp canopy area. Canopy estimations from Landsat-8 images yielded a higher NRMSE than Landsat-4 and -5 images, but the lack of matchup limits comparison. Imagery from early fall yielded the largest coverage estimates. Overall, our results show that Landsat enables broad remote measurement of bull kelp canopy coverage to supplement existing survey methods and increase continuity of timeseries for monitoring long-term trends.

A series of accidents at the Fukushima Dai-ichi Nuclear Power Plant (1F NPP) following the Great East Japan Earthquake and tsunami of 11 March 2011 resulted in the release of radioactive materials to the ocean. We used the Regional Ocean Model System (ROMS) to simulate the 137Cs activity in the oceanic area off Fukushima, with the sources of radioactivity being direct release, atmospheric deposition, river discharge, and inflow across the domain boundary. The direct release rate of 137Cs after the accident until the end of 2016 was estimated by comparing simulated results with measured 137Cs activities adjacent to the 1F NPP. River discharge rates of 137Cs were estimated by multiplying simulated river flow rates by the dissolved 137Cs activities, which were estimated by an empirical function. Inflow of 137Cs across the domain boundary was set according to the results of a North Pacific Ocean model. Because the spatiotemporal variability of 137Cs activity was large, the simulated results were compared with the annual averaged observed 137Cs activity distribution. Normalized annual averaged 137Cs activity distributions in the regional ocean were similar for each year from 2013 to 2016. This result suggests that the annual averaged distribution is predictable. Simulated 137Cs activity attributable to direct release was in good agreement with measurement data from the coastal zone adjacent to the 1F NPP. Comparison of the simulated results with measured activity in the offshore area indicated that the simulation slightly underestimated the activity attributable to inflow across the domain boundary. This result suggests that recirculation of subducted 137Cs to the surface layer was underestimated by the North Pacific model. During the study period, the effect of river discharge on oceanic 137Cs activity was small compared to the effect of directly released 137Cs.

Evaluation of radioactive cesium impact from atmospheric deposition and direct release fluxes into the North Pacific from the Fukushima Daiichi nuclear power plant

Distribution of oceanic 137Cs from the Fukushima Dai-ichi Nuclear Power Plant simulated numerically by a regional ocean model

Emissions of C9 – C16 hydrocarbons from kelp species on Vancouver Island: Alaria marginata (winged kelp) and Nereocystis luetkeana (bull kelp) as an atmospheric source of limonene

Canopy-forming kelp, which include giant kelp (Macrocystis integrifolia) and bull kelp (Nereocystis luetkeana) have been identified as ecologically significant species on the coast of British Columbia (BC). Giant kelp and bull kelp provide crucial habitat for multiple fish and invertebrate species, and they play a key role in nearshore nutrient and flow regimes. The spatial distribution of both kelp species is a crucial and currently missing input for marine protected area network planning and long-term ecological research. An ongoing collaboration between the Hakai Institute, based on the Central Coast of BC, and the Pacific Region of Fisheries and Oceans Canada, seeks to examine the application of satellite imagery for mapping kelp extent on the BC coast. A primary objective of this research is to determine whether object-based image analysis (OBIA) could be used to differentiate and delineate the extent of both giant and bull kelp using high resolution satellite imagery. A subset of pansharpened WorldView-2 imagery (0.5 m resolution) was selected for a region (McMullin Island group) on the Central Coast of BC where both species are known to be present. Knowledge of the region is extended via available field data and local ecological knowledge. While bull kelp and giant kelp have very similar spectral signatures (both are brown algae), they have very different morphologies. These morphological differences indicate that texture analysis would be best for species differentiation. Using recursive feature elimination, image feature variables (both spectral and textural) showed high differentiation between species. These variables were used as inputs for OBIA using eCognition software to test four scales of image segmentation and three different image classifiers. The results of our study demonstrate high classification accuracy for mapping bull and giant kelp. We obtained user's and producer's accuracies greater than 90% for both kelp species using a random forest classifier. We also observed an effect of the scale of image segmentation on the results of the classifications. Overall, these methods and results demonstrate a first and novel instance of the application of OBIA for mapping multiple co-occurring species of canopy-forming kelp. Future research will include examining the effects of tide height and geographical location as well as developing methods for modelling kelp biomass.

Giant kelps (which may reach lengths of 45 m) are a prominent exception to the general rule that wave-swept organisms are small. The ability of these kelps to maintain their large size in the presence of ocean waves has been attributed to their extreme flexibility and the concomitant tendency to 'go with the flow', a tendency that reduces the hydrodynamic forces imposed on the plant. However, the flexibility of giant kelps carries with it the potential for the organism to apply an inertial load to its own structure if the blade mass reaches the end of its tether. Here, we examine the complex trade-off between flexibility and inertial loading using a simple computational model of the bull kelp Nereocystis luetkeana. In field and laboratory tests, the model accurately predicts the forces and motions imposed on flexible structures in wave-induced flows. Subsequent predictions from the model suggest that mature N. luetkeana can indeed benefit from moving with the flow, but that the forces imposed on juveniles are actually increased by the plant's flexibility. Furthermore, the benefit accrued from going with the flow is sensitive to the shape of the plant. If the bull kelp were to grow while maintaining a juvenile shape, the stress placed on its stipe would be drastically increased by dynamic loading, and these inappropriately shaped plants would be subjected to a high risk of breakage. For certain combinations of wave height, wave period and stipe length, the increased stress in hypothetical 'small'-shaped plants may be associated with chaotic motion of the blade mass.

Bull kelp, Nereocystis luetkeana, is a northeastern Pacific kelp with broad distribution from Alaska to central California. Its population declines have caused severe concerns in northern California, the Salish Sea in Washington, and recently in some populations in Oregon. Despite bull kelp's accumulated ecological and physiological studies, an assembled and annotated genomic reference was still unavailable. Here, we report the complete and annotated genome of Nereocystis luetkeana, produced by the California Conservation Genomics Project (CCGP), which aims to reveal genomic diversity patterns across California by sequencing the complete genomes of approximately 150 carefully selected species. The genome was assembled into 1562 scaffolds with 449.82Mb, 80x of coverage and 22 952 gene models. BUSCO assembly showed a completeness score of 72% for the stramenopiles gene set. The mitochondria and chloroplast genome sequences have 37 Kb and 131Mb, respectively. The orthology analysis between 10 Phaeophycean genomes showed 1065 expanded and 286 unique orthogroups for this species. Pairwise comparisons showed 542 orthogroups present only in N. luetkeana and M. pyrifera, another large-body kelp. The enrichment analysis of these orthogroups showed important functions related to central metabolism and signaling due to ATPases enrichment in these two species. This genome assembly will provide an essential resource for the ecology, evolution, conservation, and breeding of bull kelp.

This study examines the effects on bull kelp (Nereocystis luetkeana) and ribbon kelp (Alaria marginata) of combinations of three climate-related stressors relevant to high-latitude kelp forests: temperature, salinity, and sediment load. Fertile specimens of both species were collected from Juneau, Alaska. Spores produced were cultivated over 40 days in four ecologically relevant stressor treatments: control (all stressor levels normal; CTRL), increased glacial melt (normal temperature, low salinity, high sediment load; GLAC), increased runoff (high temperature, low salinity, normal sediment load; MELT) and climate change (high temperature, low salinity, high sediment load; CLIM). Gametophyte density in both species was reduced in treatments involving high sediment load. Gametophyte density in bull kelp was also reduced in the increased runoff treatment, while ribbon kelp appeared resilient. Gametophytes of A. marginata grew equally in the increased glacial melt treatment as in the control and exhibited some growth in the increased runoff treatment. Conversely, gametophytes of N. luetkeana exhibited low growth in all treatments other than the control. A large number of gametophytes of both species were unidentifiable as either male or female in high-temperature treatments. This likely had impacts on reproduction, as neither species was able to produce eggs or sporophytes in these treatments. The results presented here show that both N. luetkeana (a subtidal canopy-former) and A. marginata (an intertidal subcanopy species) are sensitive to combinations of thermal, hyposaline, and sediment stress. This may have an impact on the development of gametophytes and successful reproduction in these species and may therefore have implications for the ongoing persistence of wild kelp populations in future ocean conditions.

Bull kelp, Nereocystis luetkeana, is an iconic kelp forest species of the Northeast Pacific that provides a wide range of ecosystem services to coastal marine species and society. In northern California, U.S.A., Nereocystis abundance declined sharply in 2014 and has yet to recover. While abiotic and biotic stressors were present prior to 2014, the population collapse highlights the need for a better understanding of how environmental conditions impact Nereocystis. In this study, we used a newly-developed, satellite-based dataset of bull kelp abundance, proxied by canopy cover over 20 years, to test the hypothesis that winter oceanographic conditions determine summer Nereocystis canopy cover. For the years before the collapse (1991 through 2013), wintertime ocean conditions, synthesized in a Multivariate Ocean Climate Indicator (MOCI), were indeed a good predictor of summer Nereocystis canopy cover (R2 = 0.40 to 0.87). We attribute this relationship to the effects of upwelling and/or temperature on nutrient availability. South of Point Arena, California, winter ocean conditions had slightly lower explanatory power than north of Point Arena, also reflective of spring upwelling-driven nutrient entrainment. Results suggest that the Nereocystis gametophytes and/or early sporophytes are sensitive to winter oceanographic conditions. Furthermore, environmental conditions in winter 2014 could have been used to predict the Nereocystis collapse in summer 2014, and for kelp north of Point Arena, a further decline in 2015. Importantly, environmental models do not predict changes in kelp after 2015, suggesting biotic factors suppressed kelp recovery, most likely extreme sea urchin herbivory. Conditions during winter, a season that is often overlooked in studies of biophysical interactions, are useful for predicting summer Nereocystis kelp forest canopy cover, and will be useful in supporting kelp restoration actions in California and perhaps elsewhere in the world.