Submesoscale coupling of krill and whales revealed by aggregative Lagrangian coherent structures.

In the marine environment, dynamic physical processes shape biological productivity and predator-prey interactions across multiple scales. Identifying pathways of physical-biological coupling is fundamental to understand the functioning of marine ecosystems yet it is challenging because the interactions are difficult to measure. We examined submesoscale (less than 100 km) surface current features using remote sensing techniques alongside ship-based surveys of krill and baleen whale distributions in the California Current System. We found that aggregative surface current features, represented by Lagrangian coherent structures (LCS) integrated over temporal scales between 2 and 10 days, were associated with increased (a) krill density (up to 2.6 times more dense), (b) baleen whale presence (up to 8.3 times more likely) and (c) subsurface seawater density (at depths up to 10 m). The link between physical oceanography, krill density and krill-predator distributions suggests that LCS are important features that drive the flux of energy and nutrients across trophic levels. Our results may help inform dynamic management strategies aimed at reducing large whales ship strikes and help assess the potential impacts of environmental change on this critical ecosystem.

Pelagic calcifiers face increased mortality and habitat loss with warming and ocean acidification.

Global change is impacting the oceans in an unprecedented way, and multiple lines of evidence suggest that species distributions are changing in space and time. There is increasing evidence that multiple environmental stressors act together to constrain species habitat more than expected from warming alone. Here, we conducted a comprehensive study of how temperature and aragonite saturation state act together to limit Limacina helicina, globally distributed pteropods that are ecologically important pelagic calcifiers and an indicator species for ocean change. We co-validated three different approaches to evaluate the impact of ocean warming and acidification (OWA) on the survival and distribution of this species in the California Current Ecosystem. First, we used colocated physical, chemical, and biological data from three large-scale west coast cruises and regional time series; second, we conducted multifactorial experimental incubations to evaluate how OWA impacts pteropod survival; and third, we validated the relationships we found against global distributions of pteropods and carbonate chemistry. OWA experimental work revealed mortality increases under OWA, while regional habitat suitability indices and global distributions of L. helicina suggest that a multi-stressor framework is essential for understanding pteropod distributions. In California Current Ecosystem habitats, where pteropods are living close to their thermal maximum already, additional warming and acidification through unabated fossil fuel emissions (RCP 8.5) are expected to dramatically reduce habitat suitability.

Modeling predator and prey hotspots: Management implications of baleen whale co-occurrence with krill in Central California.

As global ocean-bound commerce increases, managing human activities has become important in reducing conflict with threatened wildlife. This study investigates environmental factors determining abundance and distribution of blue whales (Balaenoptera musculus), humpback whales (Megaptera novaeangliae) and their prey (Euphausia pacifica and Thysanoessa spinifera) in central California. We provide insights into environmental drivers of the ecology and distribution of these species, model whale distributions and determine coincident hotspots of whales and their prey that will help decrease human threats to whales and protect critical feeding habitat. We developed separate predictive models of whale abundances (using negative binomial regression on count data) and krill abundance (using a two-part hurdlemodel combining logistic and negative binomial regressions) over a 14 year period (2004-2017). Variables included in situ surface and midwater oceanographic measures (temperature, salinity, and fluorescence), basin-scale climate indices, and bathymetric- and distance-related data. Predictions were applied to 1 km2 cells spanning the study area for May, June, July, and September during each of the 14 years of surveys to identify persistent distribution patterns. Both whales and krill were found to consistently use the northeast region of Cordell Bank, the Farallon Escarpment, and the shelf-break waters. The main identified blue whale hotspots were also krill hotspots, while co-occurrence was more limited and varied seasonally for humpback whales and krill. These results are valuable in identifying patterns in important areas of ecological interaction to assist management of whales. Areas north of Cordell Bank are of particular management concern since they overlap with the end of the San Francisco Bay northern shipping lane. Our findings can help decrease threats to whales, particularly in important foraging areas, by supporting implementation of vessel management and informing potential conflicts with other human uses.

Modeling Nonresident Seabird Foraging Distributions to Inform Ocean Zoning in Central California.

Seabird aggregations at sea have been shown to be associated with concentrations of prey. Previous research identified Central California as a highly used foraging area for seabirds, with locally breeding seabirds foraging close to their colonies on Southeast Farallon Island. Herein, we focus on nonresident (i.e. non-locally breeding) seabird species off of Central California. We hypothesized that high-use foraging areas for nonresident seabirds would be influenced by oceanographic and bathymetric factors and that spatial and temporal distributions would be similar within planktivorous and generalist foraging guilds but would differ between them. With data collected by the Applied California Current Ecosystem Studies (ACCESS) partnership during cruises between April and October from 2004-2013, we developed generalized linear models to identify high-use foraging areas for each of six nonresident seabird species. The four generalist species are Phoebastria nigripes (black-footed albatross), Ardenna griseus (sooty shearwater), Ardenna creatopus (pink-footed shearwater), and Fulmarus glacialis (northern fulmar). The two planktivorous species are Phalaropus lobatus (red-necked phalarope) and Phalaropus fulicarius (red phalarope). Sea surface temperature was significant for generalist species and sea surface salinity was important for planktivorous species. The distance to the 200-m isobath was significant in five of six models, Pacific Decadal Oscillation with a 3-month lag in four models, and sea surface fluorescence, the distance to Cordell Bank, and depth in three models. We did not find statistically significant differences between distributions of individual seabird species within a foraging guild or between guilds, with the exception of the sooty shearwater. Model results for a multi-use seabird foraging area highlighted the continental shelf break, particularly within the vicinity of Cordell Bank, as the highest use areas as did Marxan prioritization. Our research methods can be implemented elsewhere to identify critical habitat that needs protection as human development pressures continue to expand to the ocean.

Spatial Distribution and Temporal Patterns of Cassin's Auklet Foraging and Their Euphausiid Prey in a Variable Ocean Environment.

Krill (Euphausiids) play a vital ecosystem role in many of the world's most productive marine regions, providing an important trophic linkage. We introduce a robust modeling approach to link Cassin's auklet (Ptychoramphus aleuticus) abundance and distribution to large-scale and local oceanic and atmospheric conditions and relate these patterns to similarly modeled distributions of an important prey resource, krill. We carried out at-sea strip transect bird surveys and hydroacoustic assessments of euphausiids (2004-2013). Data informed separate, spatially-explicit predictive models of Cassin's auklet abundance (zero-inflated negative binomial regression) and krill biomass (two-part model) based on these surveys. We established the type of prey responsible for acoustic backscatter by conducting net tows of the upper 50 m during surveys. We determined the types of prey fed to Cassin's auklet chicks by collecting diet samples from provisioning adults. Using time-depth-recorders, we found Cassin's auklets utilized consistent areas in the upper water column, less than 30 m, where krill could be found (99.5% of dives were less than 30 m). Birds primarily preyed upon two species of euphausiids, Euphausia pacifica and Thysanoessa spinifera, which were available in the upper water column. Cassin's auklet abundance was best predicted by both large scale and localized oceanic processes (upwelling) while krill biomass was best predicted by local factors (temperature, salinity, and fluorescence) and both large scale and localized oceanic processes (upwelling). Models predicted varying krill and bird distribution by month and year. Our work informs the use of Cassin's auklet as a valuable indicator or krill abundance and distribution and strengthens our understanding of the link between Cassin's auklet and its primary prey. We expect future increases in frequency and magnitude of anomalous ocean conditions will result in decreased availability of krill leading to declines in the Farallon Islands population of Cassin's auklets.

Using seabird habitat modeling to inform marine spatial planning in central California's National Marine Sanctuaries.

Understanding seabird habitat preferences is critical to future wildlife conservation and threat mitigation in California. The objective of this study was to investigate drivers of seabird habitat selection within the Gulf of the Farallones and Cordell Bank National Marine Sanctuaries to identify areas for targeted conservation planning. We used seabird abundance data collected by the Applied California Current Ecosystem Studies Program (ACCESS) from 2004-2011. We used zero-inflated negative binomial regression to model species abundance and distribution as a function of near surface ocean water properties, distances to geographic features and oceanographic climate indices to identify patterns in foraging habitat selection. We evaluated seasonal, inter-annual and species-specific variability of at-sea distributions for the five most abundant seabirds nesting on the Farallon Islands: western gull (Larus occidentalis), common murre (Uria aalge), Cassin's auklet (Ptychorampus aleuticus), rhinoceros auklet (Cerorhinca monocerata) and Brandt's cormorant (Phalacrocorax penicillatus). The waters in the vicinity of Cordell Bank and the continental shelf east of the Farallon Islands emerged as persistent and highly selected foraging areas across all species. Further, we conducted a spatial prioritization exercise to optimize seabird conservation areas with and without considering impacts of current human activities. We explored three conservation scenarios where 10, 30 and 50 percent of highly selected, species-specific foraging areas would be conserved. We compared and contrasted results in relation to existing marine protected areas (MPAs) and the future alternative energy footprint identified by the California Ocean Uses Atlas. Our results show that the majority of highly selected seabird habitat lies outside of state MPAs where threats from shipping, oil spills, and offshore energy development remain. This analysis accentuates the need for innovative marine spatial planning efforts and provides a foundation on which to build more comprehensive zoning and management in California's National Marine Sanctuaries.