Impacts of marine heatwaves on top predator distributions are variable but predictable.

Marine heatwaves cause widespread environmental, biological, and socio-economic impacts, placing them at the forefront of 21st-century management challenges. However, heatwaves vary in intensity and evolution, and a paucity of information on how this variability impacts marine species limits our ability to proactively manage for these extreme events. Here, we model the effects of four recent heatwaves (2014, 2015, 2019, 2020) in the Northeastern Pacific on the distributions of 14 top predator species of ecological, cultural, and commercial importance. Predicted responses were highly variable across species and heatwaves, ranging from near total loss of habitat to a two-fold increase. Heatwaves rapidly altered political bio-geographies, with up to 10% of predicted habitat across all species shifting jurisdictions during individual heatwaves. The variability in predicted responses across species and heatwaves portends the need for novel management solutions that can rapidly respond to extreme climate events. As proof-of-concept, we developed an operational dynamic ocean management tool that predicts predator distributions and responses to extreme conditions in near real-time.

Suspected anthropogenic microparticle ingestion by Icelandic capelin.

Documented ingestion of microplastics and other anthropogenic microparticles (AMP) by marine fishes has increased over the last decade. However, baseline datasets on AMP ingestion in Arctic fish species are limited. This study documents AMP ingestion in Icelandic capelin (Mallotus villosus) and investigates how this relates to several biological variables. A total of 160 capelin gastrointestinal tracts were sampled and digested using 10 % potassium hydroxide and visual inspection protocols. A total frequency of occurrence (%FO) of 52.5 % and a mean AMP abundance of 1.33 ± 2.37 per individual was found (particles > 45 µm). The colour of AMP detected was significantly different between sampling locations. However, no differences in %FO or mean AMP abundance were detected between sampling locations or related to body size, gut fullness, or sex. This research contributes to the growing body of knowledge on AMP ingestion by Arctic marine species.

Whales in the carbon cycle: can recovery remove carbon dioxide?

The great whales (baleen and sperm whales), through their massive size and wide distribution, influence ecosystem and carbon dynamics. Whales directly store carbon in their biomass and contribute to carbon export through sinking carcasses. Whale excreta may stimulate phytoplankton growth and capture atmospheric CO2; such indirect pathways represent the greatest potential for whale-carbon sequestration but are poorly understood. We quantify the carbon values of whales while recognizing the numerous ecosystem, cultural, and moral motivations to protect them. We also propose a framework to quantify the economic value of whale carbon as populations change over time. Finally, we suggest research to address key unknowns (e.g., bioavailability of whale-derived nutrients to phytoplankton, species- and region-specific variability in whale carbon contributions).

Recovery of carbon benefits by overharvested baleen whale populations is threatened by climate change.

Despite the importance of marine megafauna on ecosystem functioning, their contribution to the oceanic carbon cycle is still poorly known. Here, we explored the role of baleen whales in the biological carbon pump across the southern hemisphere based on the historical and forecasted abundance of five baleen whale species. We modelled whale-mediated carbon sequestration through the sinking of their carcasses after natural death. We provide the first temporal dynamics of this carbon pump from 1890 to 2100, considering both the effects of exploitation and climate change on whale populations. We reveal that at their pre-exploitation abundance, the five species of southern whales could sequester 4.0 × 105 tonnes of carbon per year (tC yr-1). This estimate dropped to 0.6 × 105 tC yr-1 by 1972 following commercial whaling. However, with the projected restoration of whale populations under a RCP8.5 climate scenario, the sequestration would reach 1.7 × 105 tC yr-1 by 2100, while without climate change, recovered whale populations could sequester nearly twice as much (3.2 × 105 tC yr-1) by 2100. This highlights the persistence of whaling damages on whale populations and associated services as well as the predicted harmful impacts of climate change on whale ecosystem services.

Fast and Furious: Energetic Tradeoffs and Scaling of High-Speed Foraging in Rorqual Whales.

Although gigantic body size and obligate filter feeding mechanisms have evolved in multiple vertebrate lineages (mammals and fishes), intermittent ram (lunge) filter feeding is unique to a specific family of baleen whales: rorquals. Lunge feeding is a high cost, high benefit feeding mechanism that requires the integration of unsteady locomotion (i.e., accelerations and maneuvers); the impact of scale on the biomechanics and energetics of this foraging mode continues to be the subject of intense study. The goal of our investigation was to use a combination of multi-sensor tags paired with UAS footage to determine the impact of morphometrics such as body size on kinematic lunging parameters such as fluking timing, maximum lunging speed, and deceleration during the engulfment period for a range of species from minke to blue whales. Our results show that, in the case of krill-feeding lunges and regardless of size, animals exhibit a skewed gradient between powered and fully unpowered engulfment, with fluking generally ending at the point of both the maximum lunging speed and mouth opening. In all cases, the small amounts of propulsive thrust generated by the tail were unable to overcome the high drag forces experienced during engulfment. Assuming this thrust to be minimal, we predicted the minimum speed of lunging across scale. To minimize the energetic cost of lunge feeding, hydrodynamic theory predicts slower lunge feeding speeds regardless of body size, with a lower boundary set by the ability of the prey to avoid capture. We used empirical data to test this theory and instead found that maximum foraging speeds remain constant and high (∼4 m s-1) across body size, even as higher speeds result in lower foraging efficiency. Regardless, we found an increasing relationship between body size and this foraging efficiency, estimated as the ratio of energetic gain from prey to energetic cost. This trend held across timescales ranging from a single lunge to a single day and suggests that larger whales are capturing more prey-and more energy-at a lower cost.

Towards a North Pacific Ocean long-term monitoring program for plastic pollution: A review and recommendations for plastic ingestion bioindicators.

Marine debris is now a ubiquitous component of the Anthropocene global ocean. Plastic ingestion by marine wildlife was first reported in the 1960s and since that time, roughly one thousand marine species have been reported to consume this debris. This study focuses on plastic ingestion by marine invertebrates and vertebrates in the North Pacific Ocean. Specifically, we reviewed the scientific literature to assess the scope of the problem, identified key bioindicator species, and proposed guidelines for future monitoring of plastic debris in North Pacific marine ecosystems. Our meta-analysis confirmed that the North Pacific is among the most polluted ocean regions globally; roughly half of all fish and seabird specimens and more than three-quarters of sea turtles and bivalve specimens examined in this region had consumed plastic. While there are not enough standardized data to assess if these ingestion rates are changing, sampling standardization and reporting of methods are improving over time. Using a rubric-evaluation approach, we evaluated 352 species for their potential to serve as bioindicators of the prevalence of plastic pollution in the North Pacific. This analysis revealed a suite of 12 bioindicator species candidates which sample a variety of ecosystem components and cover a wide range of plastic size classes. Thus, we contend that these bioindicator candidates provide a key foundation for developing a comprehensive plastic monitoring program in the region. To enhance the utility of these bioindicators, we developed a framework for standardized data collection to minimize methodological variability across different studies and to facilitate the assessment of temporal trends over space and time. Tracking plastic ingestion by these bioindicators will help to assess the effectiveness of mitigation actions in the region, a critical step to evaluate progress towards sustainability and improved ocean health in the 21st century.

Enhancing the ecological realism of evolutionary mismatch theory.

Following rapid environmental change, why do some animals thrive, while others struggle? We present an expanded, cue-response framework for predicting variation in behavioral responses to novel situations. We show how signal detection theory can be used when individuals have three behavioral options (approach, avoid, or ignore). Based on this theory, we outline predictions about which animals are more likely to make mistakes around novel conditions (i.e., fall for a trap or fail to use an undervalued resource) and the intensity of that mismatch (i.e., severe versus moderate). Explicitly considering three options provides a more holistic perspective and allows us to distinguish between severe and moderate traps, which could guide management strategies in a changing world.

Baleen whale prey consumption based on high-resolution foraging measurements.

Baleen whales influence their ecosystems through immense prey consumption and nutrient recycling1-3. It is difficult to accurately gauge the magnitude of their current or historic ecosystem role without measuring feeding rates and prey consumed. To date, prey consumption of the largest species has been estimated using metabolic models3-9 based on extrapolations that lack empirical validation. Here, we used tags deployed on seven baleen whale (Mysticeti) species (n = 321 tag deployments) in conjunction with acoustic measurements of prey density to calculate prey consumption at daily to annual scales from the Atlantic, Pacific, and Southern Oceans. Our results suggest that previous studies3-9 have underestimated baleen whale prey consumption by threefold or more in some ecosystems. In the Southern Ocean alone, we calculate that pre-whaling populations of mysticetes annually consumed 430 million tonnes of Antarctic krill (Euphausia superba), twice the current estimated total biomass of E. superba10, and more than twice the global catch of marine fisheries today11. Larger whale populations may have supported higher productivity in large marine regions through enhanced nutrient recycling: our findings suggest mysticetes recycled 1.2 × 104 tonnes iron yr-1 in the Southern Ocean before whaling compared to 1.2 × 103 tonnes iron yr-1 recycled by whales today. The recovery of baleen whales and their nutrient recycling services2,3,7 could augment productivity and restore ecosystem function lost during 20th century whaling12,13.

Scaling of oscillatory kinematics and Froude efficiency in baleen whales.

High efficiency lunate-tail swimming with high-aspect-ratio lifting surfaces has evolved in many vertebrate lineages, from fish to cetaceans. Baleen whales (Mysticeti) are the largest swimming animals that exhibit this locomotor strategy, and present an ideal study system to examine how morphology and the kinematics of swimming scale to the largest body sizes. We used data from whale-borne inertial sensors coupled with morphometric measurements from aerial drones to calculate the hydrodynamic performance of oscillatory swimming in six baleen whale species ranging in body length from 5 to 25 m (fin whale, Balaenoptera physalus; Bryde's whale, Balaenoptera edeni; sei whale, Balaenoptera borealis; Antarctic minke whale, Balaenoptera bonaerensis; humpback whale, Megaptera novaeangliae; and blue whale, Balaenoptera musculus). We found that mass-specific thrust increased with both swimming speed and body size. Froude efficiency, defined as the ratio of useful power output to the rate of energy input ( Sloop, 1978), generally increased with swimming speed but decreased on average with increasing body size. This finding is contrary to previous results in smaller animals, where Froude efficiency increased with body size. Although our empirically parameterized estimates for swimming baleen whale drag were higher than those of a simple gliding model, oscillatory locomotion at this scale exhibits generally high Froude efficiency as in other adept swimmers. Our results quantify the fine-scale kinematics and estimate the hydrodynamics of routine and energetically expensive swimming modes at the largest scale.

Microplastics and microfibers in surface waters of Monterey Bay National Marine Sanctuary, California.

Despite a recent report of high concentrations of microplastics and microfibers in the mesopelagic waters of Monterey Bay National Marine Sanctuary (MBNMS), little is known about these particles in surface waters. From 2017 to 2019, we sampled two nearshore and two offshore locations within MBNMS using a manta trawl and analyzed these samples for microplastics and microfibers. We found an average concentration of 1.32 ± 0.70 (SE) particles per m3. We found the highest concentration of particles closest to shore, and the lowest concentration above the remote Davidson Seamount. Fiber-like debris was more common in offshore, as compared to nearshore, sites. Overall, particles in our samples were primarily buoyant synthetic polymers, including polypropylene and polyethylene. Our results provide baseline data on the degree of microplastic and microfiber pollution in MBNMS surface waters and confirm that this pollution can be found in waters from the surface to at least 1000 m depth.

Plastic ingestion by marine fish is widespread and increasing.

Plastic pollution has pervaded almost every facet of the biosphere, yet we lack an understanding of consumption risk by marine species at the global scale. To address this, we compile data from research documenting plastic debris ingestion by marine fish, totaling 171,774 individuals of 555 species. Overall, 386 marine fish species have ingested plastic debris including 210 species of commercial importance. However, 148 species studied had no records of plastic consumption, suggesting that while this evolutionary trap is widespread, it is not yet universal. Across all studies that accounted for microplastics, the incidence rate of plastic ingested by fish was 26%. Over the last decade this incidence has doubled, increasing by 2.4 ± 0.4% per year. This is driven both by increasing detection of smaller sized particles as a result of improved methodologies, as well as an increase in fish consuming plastic. Further, we investigated the role of geographic, ecological, and behavioral factors in the ingestion of plastic across species. These analyses revealed that the abundance of plastic in surface waters was positively correlated to plastic ingestion. Demersal species are more likely to ingest plastic in shallow waters; in contrast, pelagic species were most likely to consume plastic below the mixed layer. Mobile predatory species had the highest likelihood to ingest plastic; similarly, we found a positive relationship between trophic level and plastic ingestion. We also find evidence that surface ingestion-deep sea egestion of microplastics by mesopelagic myctophids is likely a key mechanism for the export of microplastics from the surface ocean to the seafloor, a sink for marine debris. These results elucidate the role of ecology and biogeography underlying plastic ingestion by marine fish and point toward species and regions in urgent need of study.

Odors from marine plastic debris elicit foraging behavior in sea turtles.

Pfaller et al. report that sea turtles respond to odors from biofouled plastic debris with the same behavior that is elicited by food odors, providing a possible unifying explanation for why sea turtles interact with marine plastic.

Scaling of swimming performance in baleen whales.

The scale dependence of locomotor factors has long been studied in comparative biomechanics, but remains poorly understood for animals at the upper extremes of body size. Rorqual baleen whales include the largest animals, but we lack basic kinematic data about their movements and behavior below the ocean surface. Here, we combined morphometrics from aerial drone photogrammetry, whale-borne inertial sensing tag data and hydrodynamic modeling to study the locomotion of five rorqual species. We quantified changes in tail oscillatory frequency and cruising speed for individual whales spanning a threefold variation in body length, corresponding to an order of magnitude variation in estimated body mass. Our results showed that oscillatory frequency decreases with body length (∝length-0.53) while cruising speed remains roughly invariant (∝length0.08) at 2 m s-1 We compared these measured results for oscillatory frequency against simplified models of an oscillating cantilever beam (∝length-1) and an optimized oscillating Strouhal vortex generator (∝length-1). The difference between our length-scaling exponent and the simplified models suggests that animals are often swimming non-optimally in order to feed or perform other routine behaviors. Cruising speed aligned more closely with an estimate of the optimal speed required to minimize the energetic cost of swimming (∝length0.07). Our results are among the first to elucidate the relationships between both oscillatory frequency and cruising speed and body size for free-swimming animals at the largest scale.

Memory and resource tracking drive blue whale migrations.

In terrestrial systems, the green wave hypothesis posits that migrating animals can enhance foraging opportunities by tracking phenological variation in high-quality forage across space (i.e., "resource waves"). To track resource waves, animals may rely on proximate cues and/or memory of long-term average phenologies. Although there is growing evidence of resource tracking in terrestrial migrants, such drivers remain unevaluated in migratory marine megafauna. Here we present a test of the green wave hypothesis in a marine system. We compare 10 years of blue whale movement data with the timing of the spring phytoplankton bloom resulting in increased prey availability in the California Current Ecosystem, allowing us to investigate resource tracking both contemporaneously (response to proximate cues) and based on climatological conditions (memory) during migrations. Blue whales closely tracked the long-term average phenology of the spring bloom, but did not track contemporaneous green-up. In addition, blue whale foraging locations were characterized by low long-term habitat variability and high long-term productivity compared with contemporaneous measurements. Results indicate that memory of long-term average conditions may have a previously underappreciated role in driving migratory movements of long-lived species in marine systems, and suggest that these animals may struggle to respond to rapid deviations from historical mean environmental conditions. Results further highlight that an ecological theory of migration is conserved across marine and terrestrial systems. Understanding the drivers of animal migration is critical for assessing how environmental changes will affect highly mobile fauna at a global scale.

Quantifying marine debris associated with coastal golf courses.

Identifying terrestrial sources of debris is essential to suppress the flow of plastic to the ocean. Here, we report a novel source of debris to the marine environment. From May 2016 to June 2018, we collected golf balls from coastal environments associated with five courses in Carmel, California. Our 75 collections recovered 39,602 balls from intertidal and nearshore environments adjacent to, or downriver from, the golf courses. Combining our collections with concurrent efforts of the Monterey Bay National Marine Sanctuary and the Pebble Beach Corporation, we report the retrieval of 50,681 balls, totaling approximately 2.5 tons of debris. We also examined decomposition patterns in the collected balls, which illustrate that degradation and loss of microplastic from golf balls to the marine environment may be of concern. Our findings will help to develop and direct mitigation procedures for this region and others with coastal golf courses.

Odours from marine plastic debris induce food search behaviours in a forage fish.

Plastic pollution is an anthropogenic stressor in marine ecosystems globally. Many species of marine fish (more than 50) ingest plastic debris. Ingested plastic has a variety of lethal and sublethal impacts and can be a route for bioaccumulation of toxic compounds throughout the food web. Despite its pervasiveness and severity, our mechanistic understanding of this maladaptive foraging behaviour is incomplete. Recent evidence suggests that the chemical signature of plastic debris may explain why certain species are predisposed to mistaking plastic for food. Anchovy (Engraulis sp.) are abundant forage fish in coastal upwelling systems and a critical prey resource for top predators. Anchovy ingest plastic in natural conditions, though the mechanism they use to misidentify plastic as prey is unknown. Here, we presented wild-caught schools of northern anchovy (Engraulis mordax) with odour solutions made of plastic debris and clean plastic to compare school-wide aggregation and rheotactic responses relative to food and food odour presentations. Anchovy schools responded to plastic debris odour with increased aggregation and reduced rheotaxis. These results were similar to the effects food and food odour presentations had on schools. Conversely, these behavioural responses were absent in clean plastic and control treatments. To our knowledge, this is the first experimental evidence that adult anchovy use odours to forage. We conclude that the chemical signature plastic debris acquires in the photic zone can induce foraging behaviours in anchovy schools. These findings provide further support for a chemosensory mechanism underlying plastic consumption by marine wildlife. Given the trophic position of forage fish, these findings have considerable implications for aquatic food webs and possibly human health.

Marine plastic debris emits a keystone infochemical for olfactory foraging seabirds.

Plastic debris is ingested by hundreds of species of organisms, from zooplankton to baleen whales, but how such a diversity of consumers can mistake plastic for their natural prey is largely unknown. The sensory mechanisms underlying plastic detection and consumption have rarely been examined within the context of sensory signals driving marine food web dynamics. We demonstrate experimentally that marine-seasoned microplastics produce a dimethyl sulfide (DMS) signature that is also a keystone odorant for natural trophic interactions. We further demonstrate a positive relationship between DMS responsiveness and plastic ingestion frequency using procellariiform seabirds as a model taxonomic group. Together, these results suggest that plastic debris emits the scent of a marine infochemical, creating an olfactory trap for susceptible marine wildlife.

Evidence that dimethyl sulfide facilitates a tritrophic mutualism between marine primary producers and top predators.

Tritrophic mutualistic interactions have been best studied in plant-insect systems. During these interactions, plants release volatiles in response to herbivore damage, which, in turn, facilitates predation on primary consumers or benefits the primary producer by providing nutrients. Here we explore a similar interaction in the Southern Ocean food web, where soluble iron limits primary productivity. Dimethyl sulfide has been studied in the context of global climate regulation and is an established foraging cue for marine top predators. We present evidence that procellariiform seabird species that use dimethyl sulfide as a foraging cue selectively forage on phytoplankton grazers. Their contribution of beneficial iron recycled to marine phytoplankton via excretion suggests a chemically mediated link between marine top predators and oceanic primary production.