Kelp Abundance Research Articles

Shifts in habitat use and demography of American lobsters in coastal Maine (USA) over the past quarter century

Some species are so linked to specific environments that their habitat association almost becomes a species-defining character and is used by managers and policymakers to direct their conservation. The American lobster Homarus americanus is among the most valuable fisheries species in North America and among the best studied benthic marine invertebrates in the world. Its populations and habitats have been studied and detailed in publications for over 35 yr. This lobster species was known to dwell in shelters, and their populations had historically been concentrated in shelter-providing boulder habitat. Our study revisited 20 long-term monitored sites at 10 m depth along more than 320 km of the Gulf of Maine. Surprisingly, we recorded fundamental changes in lobster abundance, habitat use, and distribution. Specifically, lobster population densities declined overall and occupancy in boulder habitats declined 60%, while densities on featureless ledge and sediment habitats increased 633 and 280%, respectively, from 2000 to 2019. Lobster rock shelter occupancy declined in recent years, but average body size increased, due in part to declines in smaller size classes. These demographic changes may result from both reduced recruitment and intraspecific competition resulting from the lower population densities. Habitat changes at our monitored sites included declines in kelp abundance, increases in diminutive algal turfs, and nearly 3°C warming of benthic water temperature in July (1995-2021), some of which may have contributed indirectly to those shifts. While these changes in shallow water habitat and demography have implications for the lobster fishery and stock assessments, it also illustrates previously undescribed behavioral plasticity.

Ocean warming undermines the recovery resilience of New England kelp forests following a fishery-induced trophic cascade.

Ecological theory predicts that kelp forests structured by trophic cascades should experience recovery and persistence of their foundation species when herbivores become rare. Yet, climate change may be altering the outcomes of top-down forcing in kelp forests, especially those located in regions that have rapidly warmed in recent decades, such as the Gulf of Maine. Here, using data collected annually from 30+ sites spanning >350 km of coastline, we explored the dynamics of Maine's kelp forests in the ~20 years after a fishery-induced elimination of sea urchin herbivores. Although forests (Saccharina latissima and Laminaria digitata) had broadly returned to Maine in the late 20th century, we found that forests in northeast Maine have since experienced slow but significant declines in kelp, and forest persistence in the northeast was juxtaposed by a rapid, widespread collapse in the southwest. Forests collapsed in the southwest apparently because ocean warming has-directly and indirectly-made this area inhospitable to kelp. Indeed, when modeling drivers of change using causal techniques from econometrics, we discovered that unusually high summer seawater temperatures the year prior, unusually high spring seawater temperatures, and high sea urchin densities each negatively impacted kelp abundance. Furthermore, the relative power and absolute impact of these drivers varied geographically. Our findings reveal that ocean warming is redefining the outcomes of top-down forcing in this system, whereby herbivore removal no longer predictably leads to a sustained dominance of foundational kelps but instead has led to a waning dominance (northeast) or the rise of a novel phase state defined by "turf" algae (southwest). Such findings indicate that limiting climate change and managing for low herbivore abundances will be essential for preventing further loss of the vast forests that still exist in northeast Maine. They also more broadly highlight that climate change is "rewriting the rules" of nature, and thus that ecological theory and practice must be revised to account for shifting species and processes.

Regime shifts in rocky intertidal communities associated with a marine heatwave and disease outbreak.

Long-term, large-scale experimental studies provide critical information about how global change influences communities. When environmental changes are severe, they can trigger abrupt transitions from one community type to another leading to a regime shift. From 2014 to 2016, rocky intertidal habitats in the northeast Pacific Ocean experienced extreme temperatures during a multi-year marine heatwave (MHW) and sharp population declines of the keystone predator Pisaster ochraceus due to sea star wasting disease (SSWD). Here we measured the community structure before, during and after the MHW onset and SSWD outbreak in a 15-year succession experiment conducted in a rocky intertidal meta-ecosystem spanning 13 sites on four capes in Oregon and northern California, United States. Kelp abundance declined during the MHW due to extreme temperatures, while gooseneck barnacle and mussel abundances increased due to reduced predation pressure after the loss of Pisaster from SSWD. Using several methods, we detected regime shifts from substrate- or algae-dominated to invertebrate-dominated alternative states at two capes. After water temperatures cooled and Pisaster population densities recovered, community structure differed from pre-disturbance conditions, suggesting low resilience. Consequently, thermal stress and predator loss can result in regime shifts that fundamentally alter community structure even after restoration of baseline conditions.

The effect of sedimentation on spore settlement and recruitment of the endemic Arctic kelp, Laminaria solidungula (Phaeophyceae).

Environmental changes associated with rapid climate change in the Arctic, such as the increased rates of sedimentation from climatic or anthropogenic sources, can enhance the impact of abiotic stressors on coastal ecosystems. High sedimentation rates can be detrimental to nearshore kelp abundance and distribution, possibly due to increased mortality at the spore settlement stage. Spore settlement and viability of the Arctic kelp Laminaria solidungula were examined through a series of lab-based sedimentation experiments. Spores were exposed to increasing sediment loads in three experimental designs simulating different sedimentation scenarios: sediment deposition above settled spores, settlement of spores on sediment-covered substrate, and simultaneous suspension of spores and sediments during settlement. Spore settlement was recorded upon completion of each experiment, and gametophyte abundance was assessed following a growth period with sediments removed to examine short-term spore viability via a gametophyte-to-settled-spore ratio. In all three types of sediment exposure, the addition of sediments caused a 30%-40% reduction in spore settlement relative to a no-sediment control. Spore settlement decreased significantly between the low and high sediment treatments when spores were settled onto sediment-covered substrates. In all experiments, increasing amounts of sediment had no significant effect on spore viability, indicating that spores that had settled under different short-term sediment conditions were viable. Our results indicate that depending on spore-sediment interaction type, higher rates of sedimentation resulting from increased sediment loading could affect L. solidungula spore settlement success with potential impacts on the long-term persistence of a diverse and productive benthic habitat.

Local and regional variation in kelp loss and stability across coastal British Columbia

Kelp forests are among the most abundant coastal marine habitats but are vulnerable to climate change. The Northeast Pacific has experienced recent large-scale changes in kelp abundance and distribution, but little is known about changes north of the British Columbia (BC)-Washington border. Here, we assessed whether and how floating canopy kelp (Macrocystis pyrifera, Nereocystis luetkeana) distributions have changed in recent decades along the extensive coast of BC. We assembled and analysed available kelp distributional data, comparing snapshots of kelp linear extent from 1.5-3 decades ago (1994-2007) to recently collected data (2017-2021) across 11 different subregions spanning the province. We then leveraged timeseries, where available (n = 7 data sets), to contextualise patterns of change. In aggregate, the data suggest that kelp forests have declined considerably in some parts of the province, but with variable patterns of change across BC. In the warmest areas (southern BC), kelp persistence was negatively correlated with mean summer sea surface temperatures, which at times exceeded known thermal tolerances. In contrast, in northern subregions, top-down control by sea urchins and otters appeared to modulate kelp dynamics, with declines occurring in 2 subregions despite cool ocean temperatures. Timeseries data suggest that many declines occurred around the 2014-2016 marine heatwave, an event associated with sustained warming and altered trophic dynamics. Our results suggest that the extent of BC’s kelp forests has declined in some places in recent decades, but that regional and local-scale factors influence their responses to environmental change.

Loss, resilience and recovery of kelp forests in a region of rapid ocean warming.

Changes in kelp abundances on regional scales have been highly variable over the past half-century owing to strong effects of local and regional drivers. Here, we assess patterns and dominant environmental variables causing spatial and interspecific variability in kelp persistence and resilience to change in Nova Scotia over the past 40 years. We conducted a survey of macrophyte abundance at 251 sites spanning the Atlantic coast of Nova Scotia from 2019 to 2022. We use this dataset to describe spatial variability in kelp species abundances, compare species occurrences to surveys conducted in 1982 and assess changes in kelp abundance over the past 22 years. We then relate spatial and temporal patterns in abundance and resilience to environmental metrics. Our results show losses of sea urchins and the cold-tolerant kelp species Alaria esculenta, Saccorhiza dermatodea and Agarum clathratum in Nova Scotia since 1982 in favour of the more warm-tolerant kelps Saccharina latissima and Laminaria digitata. Kelp abundances have increased slightly since 2000, and Saccharina latissima and L. digitata are widely abundant in the region today. The highest kelp cover occurs on wave-exposed shores and at sites where temperatures have remained below thresholds for growth (21 °C) and mortality (23 °C). Moreover, kelp has recovered from turf dominance following losses at some sites during a warm period from 2010 to 2012. Our results indicate that dramatic changes in kelp community composition and a loss of sea urchin herbivory as a dominant driver of change in the system have occurred in Nova Scotia over the past 40 years. However, a broad-scale shift to turf-dominance has not occurred, as predicted, and our results suggest that resilience and persistence are still a feature of kelp forests in the region despite rapid warming over the past several decades.

Management policies impact fishers’ income beyond tolerable limits: The case of the Chilean TURFs in Central Chile

Fish trade worldwide seeks sustainable fishery production which right-based-fisheries should provide. However, fisheries managed in TURF system (Territorial Use Rights for Fisheries), as occurs for loco (Concholepas concholepas) and kelp fisheries in central Chile, can turn out to be highly fluctuating. This can pose a challenge to fisher’s livelihood depending on the income variability this type of management produces. This work analyzes temporal variations of loco and kelp abundances and landings at increasing spatial scales, investigating which would be a TURF size that could dampen the influence of natural variations on fishers’ income enough for it to be tolerable. We analyzed loco and kelp abundance and landings data from hard bottom TURFs in Atacama (n = 22) and Coquimbo (n = 61) regions of Central Chile, which represent 81.4% of the total TURFs in these regions. Income variations less than 39% were considered tolerable, according to a study conducted by the National Consumer Service, where 61% of a person’s salary is necessary to cover essential needs in Chilean family. Loco and kelp landings variations exceeded that criterion at all analyzed spatial scales, suggesting that they cannot rely solely on those fisheries within a TURF. Even increasing the size of TURF including the coast of an entire region, does not dampen variability to a tolerable value. This explains the reason for former fisher migrations within these fisheries, something present regulations prevent, generating a complex situation which is necessary to revise in order not to maintain an incentive to side step regulations.

Marine invertebrates as proxies for early kelp use along the western coast of South America

Coastal societies have lived at the seaward edge of the Atacama Desert since at least 12,000 years ago. Kelp forest ecosystems provide evidence for important subsistence activity along the entire Chilean coast through fishing and gathering. Despite their importance, especially in hyperarid contexts with limited plant abundance, there is scarce evidence of kelp in archaeological contexts, hampering the study of kelp use in the past. In the present study, we use the presence of small marine invertebrates, inhabitants of stipes and holdfasts of macroalgae, as proxies that indicate past kelp presence. We analyze samples of three species of snails (Tegula atra,Tegula tridentata,andDiloma nigerrima) and one limpet (Scurria scurra) from nine archaeological sites dated between 7,000 and 500 cal years before present located around the area of Taltal (25°Lat S). Modern samples of these species were collected to reconstruct the size of fragmented archaeological shells and subsequently estimate the size of harvested kelps. Through this approach, we estimated the size and relative abundance of kelp used by coastal groups that inhabited the southern part of the Atacama Desert for around 6,500 years. Our results are a contribution to the scarce information on the presence and use of kelp in the prehistory of the Americas and contribute to comparative perspectives with other areas of the world where the use of kelp by humans in the past has already been explored.

Individual and population‐level variation in susceptibility to temperature in early life history stages of giant kelp

AbstractBecause foundation species create structure in a community, understanding their ecological and evolutionary responses to global change is critical for predicting the ecological and economic management of species and communities that rely on them. Giant kelp (Macrocystis pyrifera) is a globally distributed foundation species with seasonal fluctuations in abundance in response to local nutrient levels, storm intensity, and ocean temperatures. Here we examine genetic variation in individual and population‐level responses of early life history stages (zoospore settlement, survival, and gametogenesis) to increased temperatures to determine the potential for natural selection on temperature‐tolerant individuals that would allow adaptation to a changing climate. We collected fertile M. pyrifera sporophyll blades from three sites along the California coast (Los Angeles, Santa Barbara, Monterey Bay) and induced zoospore release in the lab. Spores settled on microscope slides at three treatment temperatures (16, 20, and 22°C), matured for 21 days, and were imaged weekly to determine settlement, survival, and maturation success. On average, individuals from all sites showed lower rates of settlement and maturation in response to increasing temperature. However, the magnitude of the responses to temperature varied among populations. Survival tended to increase with temperature in Los Angeles and Santa Barbara populations but decreased with increasing temperature for the Monterey Bay population. We observed little genetic variation in temperature responses among individuals within sites, suggesting little scope for evolution within populations to increase the resilience of M. pyrifera populations to warming ocean temperatures and predicted declines in kelp abundance. Yet sufficient dispersal among populations could allow for adaptation of early life history traits among populations via evolutionary rescue of declining populations.

Nearshore subtidal community response during and after sediment disturbance associated with dam removal

Dam removal is used increasingly to restore aquatic ecosystems and remove unnecessary or high-risk infrastructure. As the number of removals increases, there is a growing understanding about the hydrologic, geomorphic, and ecological responses to these removals. Most dam removal studies, however, focus on river and watershed responses to dam removal. The removal of two dams on the Elwha River provided a unique opportunity to characterize the response of nearshore (coastal) ecosystems. We conducted SCUBA surveys between 2011 and 2022 to quantify trajectories of change in a nearshore ecosystem during and after dam removal. We focused on the degree to which the abundances of kelp, benthic invertebrates, and fish changed in response to patterns of sediment fluxes during and after dam removal. Our findings point to two pathways of response depending on the disturbance mechanism and species type. Sites with persistent sediment deposition were characterized by wholesale community changes that did not recover to a before dam removal condition. Instead, the sites were colonized by new species that were largely absent prior to dam removal. Sites that experienced high turbidity but lacked persistent seafloor deposition were primarily characterized by a reduction in the abundance of kelp and other algae during dam removal and a rapid recovery after sediment flux to the nearshore declined. Dam removal influences on invertebrates and fish at these sites were more variable, benefiting some species and disadvantaging others. In addition to dam removal, sea star wasting syndrome and a marine heatwave exerted distinct controls on subtidal communities during the same period. The loss of the predatory sea star Pycnopodia helianthoides was associated with gains in some of its prey species, and kelp community changes reflected regional trends in ocean temperature and kelp abundance. The results presented here have important implications for understanding the response of marine ecosystems to future dam removals and similar sediment perturbation events.

High-latitude kelps and future oceans: A review of multiple stressor impacts in a changing world.

Kelp forests worldwide are threatened by both climate change and localized anthropogenic impacts. Species with cold-temperate, subpolar, or polar distributions are projected to experience range contractions over the coming decades, which may be exacerbated by climatic events such as marine heatwaves and increased freshwater and sediment input from rapidly contracting glaciers. The northeast Pacific has an extensive history of harvesting and cultivating kelps for subsistence, commercial, and other uses, and, therefore, declines in kelp abundance and distributional shifts will have significant impacts on this region. Gaps in our understanding of how cold-temperate kelp species respond to climate stressors have limited our ability to forecast the status of kelp forests in future oceans, which hampers conservation and management efforts. Here, we conducted a structured literature review to provide a synthesis of the impacts of multiple climate-related stressors on kelp forests in the northeast Pacific, assess existing knowledge gaps, and suggest potential research priorities. We chose to focus on temperature, salinity, sediment load, and light as the stressors most likely to vary and impact kelps as climate change progresses. Our results revealed biases in the existing literature toward studies investigating the impacts of temperature, or temperature in combination with light. Other stressors, particularly salinity and sediment load, have received much less focus despite rapidly changing conditions in high-latitude regions. Furthermore, multiple stressor studies appear to focus on kelp sporophytes, and it is necessary that we improve our understanding of how kelp microstages will be affected by stressor combinations. Finally, studies that investigate the potential of experimental transplantation or selective cultivation of genotypes resilient to environmental changes are lacking and would be useful for the conservation of wild populations and the seaweed aquaculture industry.

Temporal dynamics of the deep-sea pink urchin Strongylocentrotus fragilis on the Northeast Pacific continental margin

The Northeast Pacific continental margin is characterized by strong seasonal upwelling, which drives high primary productivity, and supports high diversity and biomass of benthic megafauna. The recent occurrence of a marine heat wave (“The Blob”, sensu Kintisch, 2015) in 2013–2016 resulted in changes to phytoplankton community composition and loss of coastal kelp abundance and diversity, reducing gross primary productivity in the region. However, cumulative effects of marine heat waves and ongoing basin-scale deoxygenation in deep-sea ecosystems remain poorly understood. Here, we use a 7-year time series of physicochemical and video imagery data from Ocean Networks Canada's NEPTUNE observatory to investigate temporal dynamics of the deep-sea pink urchin Strongylocentrotus fragilis in relation to multi-year environmental variability. Using generalized additive models, we show that local S. fragilis density at Barkley Upper Slope (420 m) fluctuated over time and was partially explained by changes in dissolved oxygen concentration and suspended particulate matter in the benthic boundary layer (ADCP backscatter), with high urchin density corresponding to high oxygen and low backscatter. Seafloor dissolved oxygen ranged from 0.80 to 1.89 mL/L and varied seasonally, exhibiting a clear negative correlation with sea surface primary productivity (MODIS satellite Chl-a data), corresponding with the onset of yearly upwelling conditions. However, during the anomalously warm years affected by ‘The Blob’, dampened upwelling maintained higher dissolved oxygen conditions near the seafloor. S. fragilis density declined during ‘Blob’ conditions, likely in response to reduced kelp and phytodetritus subsidies from coastal waters. We propose a foraging-respiration trade-off hypothesis, whereby S. fragilis forages in deeper water during weak upwelling and migrates to shallower habitats during low oxygen conditions. S. fragilis is an important bioturbator and detritivore; changes in the density and distribution of this species may directly affect sediment turnover rates and nutrient cycling on the continental margin, with consequences for surface and coastal productivity.https://github.com/rjcommand/.

Using unoccupied aerial vehicles to map and monitor changes in emergent kelp canopy after an ecological regime shift

AbstractKelp forests are complex underwater habitats that form the foundation of many nearshore marine environments and provide valuable services for coastal communities. Despite their ecological and economic importance, increasingly severe stressors have resulted in declines in kelp abundance in many regions over the past few decades, including the North Coast of California, USA. Given the significant and sustained loss of kelp in this region, management intervention is likely a necessary tool to reset the ecosystem and geospatial data on kelp dynamics are needed to strategically implement restoration projects. Because canopy‐forming kelp forests are distinguishable in aerial imagery, remote sensing is an important tool for documenting changes in canopy area and abundance to meet these data needs. We used small unoccupied aerial vehicles (UAVs) to survey emergent kelp canopy in priority sites along the North Coast in 2019 and 2020 to fill a key data gap for kelp restoration practitioners working at local scales. With over 4,300 hectares surveyed between 2019 and 2020, these surveys represent the two largest marine resource‐focused UAV surveys conducted in California to our knowledge. We present remote sensing methods using UAVs and a repeatable workflow for conducting consistent surveys, creating orthomosaics, georeferencing data, classifying emergent kelp and creating kelp canopy maps that can be used to assess trends in kelp canopy dynamics over space and time. We illustrate the impacts of spatial resolution on emergent kelp canopy classification between different sensors to help practitioners decide which data stream to select when asking restoration and management questions at varying spatial scales. Our results suggest that high spatial resolution data of emergent kelp canopy from UAVs have the potential to advance strategic kelp restoration and adaptive management.

Kelp carbon sink potential decreases with warming due to accelerating decomposition.

Cycling of organic carbon in the ocean has the potential to mitigate or exacerbate global climate change, but major questions remain about the environmental controls on organic carbon flux in the coastal zone. Here, we used a field experiment distributed across 28° of latitude, and the entire range of 2 dominant kelp species in the northern hemisphere, to measure decomposition rates of kelp detritus on the seafloor in relation to local environmental factors. Detritus decomposition in both species were strongly related to ocean temperature and initial carbon content, with higher rates of biomass loss at lower latitudes with warmer temperatures. Our experiment showed slow overall decomposition and turnover of kelp detritus and modeling of coastal residence times at our study sites revealed that a significant portion of this production can remain intact long enough to reach deep marine sinks. The results suggest that decomposition of these kelp species could accelerate with ocean warming and that low-latitude kelp forests could experience the greatest increase in remineralization with a 9% to 42% reduced potential for transport to long-term ocean sinks under short-term (RCP4.5) and long-term (RCP8.5) warming scenarios. However, slow decomposition at high latitudes, where kelp abundance is predicted to expand, indicates potential for increasing kelp-carbon sinks in cooler (northern) regions. Our findings reveal an important latitudinal gradient in coastal ecosystem function that provides an improved capacity to predict the implications of ocean warming on carbon cycling. Broad-scale patterns in organic carbon decomposition revealed here can be used to identify hotspots of carbon sequestration potential and resolve relationships between carbon cycling processes and ocean climate at a global scale.

Choosing the optimal method of stock assessment for Saccharina japonica in the northwestern Tatar Strait

Experiments on comparing the methods for assessment of Saccharina japonica commercial stock were conducted in 2021. In total, 11 sites in the northwestern Tatar Strait southward from the Sovetskaya Gavan Bay, along the 44 km stretch of coastline between Cape Krasny Partizan (48о58′ N 140о23′ E) and Cape Korovin (48о37′ N 140о11′ E), were surveyed aboard RV Ubezhdenny and small boats. This is a traditional area of Saccharina japonica commercial harvesting. Four methods were compared: 1) visual assessment of projective cover from the sea surface by two observers independently; 2) assessment of projective cover by two observers independently on the images obtained from a remotely operated underwater vehicle (ROV); 3) estimation of projective cover by a diver; and 4) assessment on the data of quantitative diving samples. The former three methods are comparable and differ by 1.2–1.7 times. The estimates made by the latter method of diving sampling exceed them by 2.4–4.1 times because of artifacts specific to this method. To eliminate inaccuracies, diving samples should be collected under control from the sea surface by ROV. Evaluation of the projective cover by diver is inadvisable. Visual assessments of the projective cover from the sea surface are simple and cheap but inconvenient for documentation. The stock assessment with micro-ROV is recognized as the optimal method provided an adequate accounting of the kelp abundance and complete documenting of the survey materials.

Winter oceanographic conditions predict summer bull kelp canopy cover in northern California.

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.

Southeast Alaskan kelp forests: inferences of process from large-scale patterns of variation in space and time.

Humans were considered external drivers in much foundational ecological research. A recognition that humans are embedded in the complex interaction networks we study can provide new insight into our ecological paradigms. Here, we use time-series data spanning three decades to explore the effects of human harvesting on otter–urchin–kelp trophic cascades in southeast Alaska. These effects were inferred from variation in sea urchin and kelp abundance following the post fur trade repatriation of otters and a subsequent localized reduction of otters by human harvest in one location. In an example of a classic trophic cascade, otter repatriation was followed by a 99% reduction in urchin biomass density and a greater than 99% increase in kelp density region wide. Recent spatially concentrated harvesting of otters was associated with a localized 70% decline in otter abundance in one location, with urchins increasing and kelps declining in accordance with the spatial pattern of otter occupancy within that region. While the otter–urchin–kelp trophic cascade has been associated with alternative community states at the regional scale, this research highlights how small-scale variability in otter occupancy, ostensibly due to spatial variability in harvesting or the risk landscape for otters, can result in within-region patchiness in these community states.

Toward a conceptual framework for managing and conserving marine habitats: A case study of kelp forests in the Salish Sea.

Kelp forests are in decline across much of their range due to place‐specific combinations of local and global stressors. Declines in kelp abundance can lead to cascading losses of biodiversity and productivity with far‐reaching ecological and socioeconomic consequences. The Salish Sea is a hotspot of kelp diversity where many species of kelp provide critical habitat and food for commercially, ecologically, and culturally important fish and invertebrate species. However, like other regions, kelp forests in much of the Salish Sea are in rapid decline. Data gaps and limited long‐term monitoring have hampered attempts to identify and manage for specific drivers of decline, despite the documented urgency to protect these important habitats. To address these knowledge gaps, we gathered a focus group of experts on kelp in the Salish Sea to identify perceived direct and indirect stressors facing kelp forests. We then conducted a comprehensive literature review of peer‐reviewed studies from the Salish Sea and temperate coastal ecosystems worldwide to assess the level of support for the pathways identified by the experts, and we identified knowledge gaps to prioritize future research. Our results revealed major research gaps within the Salish Sea and highlighted the potential to use expert knowledge for making informed decisions in the region. We found high support for the pathways in the global literature, with variable consensus on the relationship between stressors and responses across studies, confirming the influence of local ecological, oceanographic, and anthropogenic contexts and threshold effects on stressor–response relationships. Finally, we prioritized areas for future research in the Salish Sea. This study demonstrates the value expert opinion has to inform management decisions. These methods are readily adaptable to other ecosystem management contexts, and the results of this case study can be immediately applied to kelp management.

Long-term changes in reef fish assemblages after 40 years of no-take marine reserve protection

Marine reserves can increase the abundance and size of harvested fish species, but the indirect effects of protection on wider fish assemblages are less well understood. Better understanding how marine reserves indirectly effect reef fish is essential to interpreting changes in non-targeted taxa and informing management expectations as to the long-term effects of marine protection. We investigated how reef fish assemblages and habitats have changed within New Zealand's oldest marine reserve after 40 years of no-take protection (1978 to 2018), and how they differed inside and outside the reserve in 2018. Reef fish assemblages differed between 1978 and 2018, and between reserve and fished sites, and these effects were greatest at depths where habitats had changed. We report an overall increase in fish biomass in the reserve, but a long-term decline in total fish density. The increase in biomass under protection was largely driven by an increase in six target species. Eleven species exhibited long-term declines in the marine reserve; three likely a result of indirect effects related to habitat change and/or increased predation, while other declines likely reflect large-scale changes independent of reserve protection. Predicted indirect effects associated with large-scale habitat change were not apparent, except for an increase in one non-target herbivore species alongside increased kelp abundance. While further work is needed to confirm the mechanisms driving individual indirect effects, the results demonstrate that the long-term indirect effects of protection on reef fish are less common, more complex and more difficult to predict than direct effects.

Diet of a threatened endemic fox reveals variation in sandy beach resource use on California Channel Islands.

The coastal zone provides foraging opportunities for insular populations of terrestrial mammals, allowing for expanded habitat use, increased dietary breadth, and locally higher population densities. We examined the use of sandy beach resources by the threatened island fox (Urocyon littoralis) on the California Channel Islands using scat analysis, surveys of potential prey, beach habitat attributes, and stable isotope analysis. Consumption of beach invertebrates, primarily intertidal talitrid amphipods (Megalorchestia spp.) by island fox varied with abundance of these prey across sites. Distance-based linear modeling revealed that abundance of giant kelp (Macrocystis pyrifera) wrack, rather than beach physical attributes, explained the largest amount of variation in talitrid amphipod abundance and biomass across beaches. δ13C and δ15N values of fox whisker (vibrissae) segments suggested individualism in diet, with generally low δ13C and δ15N values of some foxes consistent with specializing on primarily terrestrial foods, contrasting with the higher isotope values of other individuals that suggested a sustained use of sandy beach resources, the importance of which varied over time. Abundant allochthonous marine resources on beaches, including inputs of giant kelp, may expand habitat use and diet breadth of the island fox, increasing population resilience during declines in terrestrial resources associated with climate variability and long-term climate change.