Jarrod A. Santora

Increasing variance in North Pacific climate relates to unprecedented ecosystem variability off California

Changes in variance are infrequently examined in climate change ecology. We tested the hypothesis that recent high variability in demographic attributes of salmon and seabirds off California is related to increasing variability in remote, large-scale forcing in the North Pacific operating through changes in local food webs. Linear, indirect numerical responses between krill (primarily Thysanoessa spinifera) and juvenile rockfish abundance (catch per unit effort (CPUE)) explained >80% of the recent variability in the demography of these pelagic predators. We found no relationships between krill and regional upwelling, though a strong connection to the North Pacific Gyre Oscillation (NPGO) index was established. Variance in NPGO and related central Pacific warming index increased after 1985, whereas variance in the canonical ENSO and Pacific Decadal Oscillation did not change. Anthropogenic global warming or natural climate variability may explain recent intensification of the NPGO and its increasing ecological significance. Assessing non-stationarity in atmospheric-environmental interactions and placing greater emphasis on documenting changes in variance of bio-physical systems will enable insight into complex climate-marine ecosystem dynamics.

Spatio‐temporal dynamics of ocean conditions and forage taxa reveal regional structuring of seabird–prey relationships

Studies of predator–prey demographic responses and the physical drivers of such relationships are rare, yet essential for predicting future changes in the structure and dynamics of marine ecosystems. Here, we hypothesize that predator–prey relationships vary spatially in association with underlying physical ocean conditions, leading to observable changes in demographic rates, such as reproduction. To test this hypothesis, we quantified spatio-temporal variability in hydrographic conditions, krill, and forage fish to model predator (seabird) demographic responses over 18 years (1990–2007). We used principal component analysis and spatial correlation maps to assess coherence among ocean conditions, krill, and forage fish, and generalized additive models to quantify interannual variability in seabird breeding success relative to prey abundance. The first principal component of four hydrographic measurements yielded an index that partitioned “warm/weak upwelling” and “cool/strong upwelling” years. Partitioning of krill and forage fish time series among shelf and oceanic regions yielded spatially explicit indicators of prey availability. Krill abundance within the oceanic region was remarkably consistent between years, whereas krill over the shelf showed marked interannual fluctuations in relation to ocean conditions. Anchovy abundance varied on the shelf, and was greater in years of strong stratification, weak upwelling and warmer temperatures. Spatio-temporal variability of juvenile forage fish co-varied strongly with each other and with krill, but was weakly correlated with hydrographic conditions. Demographic responses between seabirds and prey availability revealed spatially variable associations indicative of the dynamic nature of “predator–habitat” relationships. Quantification of spatially explicit demographic responses, and their variability through time, demonstrate the possibility of delineating specific critical areas where the implementation of protective measures could maintain functions and productivity of central place foraging predators.

A spatially distinct history of the development of california groundfish fisheries.

During the past century, commercial fisheries have expanded from small vessels fishing in shallow, coastal habitats to a broad suite of vessels and gears that fish virtually every marine habitat on the globe. Understanding how fisheries have developed in space and time is critical for interpreting and managing the response of ecosystems to the effects of fishing, however time series of spatially explicit data are typically rare. Recently, the 1933–1968 portion of the commercial catch dataset from the California Department of Fish and Wildlife was recovered and digitized, completing the full historical series for both commercial and recreational datasets from 1933–2010. These unique datasets include landing estimates at a coarse 10 by 10 minute “grid-block” spatial resolution and extends the entire length of coastal California up to 180 kilometers from shore. In this study, we focus on the catch history of groundfish which were mapped for each grid-block using the year at 50% cumulative catch and total historical catch per habitat area. We then constructed generalized linear models to quantify the relationship between spatiotemporal trends in groundfish catches, distance from ports, depth, percentage of days with wind speed over 15 knots, SST and ocean productivity. Our results indicate that over the history of these fisheries, catches have taken place in increasingly deeper habitat, at a greater distance from ports, and in increasingly inclement weather conditions. Understanding spatial development of groundfish fisheries and catches in California are critical for improving population models and for evaluating whether implicit stock assessment model assumptions of relative homogeneity of fisheries removals over time and space are reasonable. This newly reconstructed catch dataset and analysis provides a comprehensive appreciation for the development of groundfish fisheries with respect to commonly assumed trends of global fisheries patterns that are typically constrained by a lack of long-term spatial datasets.

Application of a data‐assimilative regional ocean modeling system for assessing California Current System ocean conditions, krill, and juvenile rockfish interannual variability

To be robust and informative, marine ecosystem models and assessments require parameterized biophysical relationships that rely on realistic water column characteristics at appropriate spatial and temporal scales. We examine how hydrographic properties off California from 1990 through 2010 during late winter and spring correspond to krill and juvenile rockfish (Sebastes spp.) abundances. We evaluated coherence among temperature, salinity, depth of 26.0 potential density isopycnal, and stratification strength at regionally and monthly time scales derived from shipboard and mooring observations, and a data-assimilative Regional Ocean Model System reanalysis. The reanalysis captures spatiotemporal physical variability of coastal ocean conditions in winter and spring months and elucidates mechanisms connecting the spatial and temporal upwelling and transport dynamics on observed krill and rockfish abundances in spring. This provides evidence for a mechanistic connection between the phenology of upwelling in the California Current System and seasonal development of the shelf ecosystem.

Persistence of trophic hotspots and relation to human impacts within an upwelling marine ecosystem

Human impacts (e.g., fishing, pollution, and shipping) on pelagic ecosystems are increasing, causing concerns about stresses on marine food webs. Maintaining predator-prey relationships through protection of pelagic hotspots is crucial for conservation and management of living marine resources. Biotic components of pelagic, plankton-based, ecosystems exhibit high variability in abundance in time and space (i.e., extreme patchiness), requiring investigation of persistence of abundance across trophic levels to resolve trophic hotspots. Using a 26-yr record of indicators for primary production, secondary (zooplankton and larval fish), and tertiary (seabirds) consumers, we show distributions of trophic hotspots in the southern California Current Ecosystem result from interactions between a strong upwelling center and a productive retention zone with enhanced nutrients, which concentrate prey and predators across multiple trophic levels. Trophic hotspots also overlap with human impacts, including fisheries extraction of coastal pelagic and groundfish species, as well as intense commercial shipping traffic. Spatial overlap of trophic hotspots with fisheries and shipping increases vulnerability of the ecosystem to localized depletion of forage fish, ship strikes on marine mammals, and pollution. This study represents a critical step toward resolving pelagic areas of high conservation interest for planktonic ecosystems and may serve as a model for other ocean regions where ecosystem-based management and marine spatial planning of pelagic ecosystems is warranted.

Timing of ice retreat alters seabird abundances and distributions in the southeast Bering Sea.

Timing of spring sea-ice retreat shapes the southeast Bering Sea food web. We compared summer seabird densities and average bathymetry depth distributions between years with early (typically warm) and late (typically cold) ice retreat. Averaged over all seabird species, densities in early-ice-retreat-years were 10.1% (95% CI: 1.1–47.9%) of that in late-ice-retreat-years. In early-ice-retreat-years, surface-foraging species had increased numbers over the middle shelf (50–150 m) and reduced numbers over the shelf slope (200–500 m). Pursuit-diving seabirds showed a less clear trend. Euphausiids and the copepod Calanus marshallae/glacialis were 2.4 and 18.1 times less abundant in early-ice-retreat-years, respectively, whereas age-0 walleye pollock Gadus chalcogrammus near-surface densities were 51× higher in early-ice-retreat-years. Our results suggest a mechanistic understanding of how present and future changes in sea-ice-retreat timing may affect top predators like seabirds in the southeastern Bering Sea.

Projecting marine mammal distribution in a changing climate

Climate-related shifts in marine mammal range and distribution have been observed in some populations; however, the nature and magnitude of future responses are uncertain in novel environments projected under climate change. This poses a challenge for agencies charged with management and conservation of these species. Specialized diets, restricted ranges, or reliance on specific substrates or sites (e.g., for pupping) make many marine mammal populations particularly vulnerable to climate change. High-latitude, predominantly ice-obligate, species have experienced some of the largest changes in habitat and distribution and these are expected to continue. Efforts to predict and project marine mammal distributions to date have emphasized data-driven statistical habitat models. These have proven successful for short time-scale (e.g., seasonal) management activities, but confidence that such relationships will hold for multi-decade projections and novel environments is limited. Recent advances in mechanistic modeling of marine mammals (i.e., models that rely on robust physiological and ecological principles expected to hold under climate change) may address this limitation. The success of such approaches rests on continued advances in marine mammal ecology, behavior, and physiology together with improved regional climate projections. The broad scope of this challenge suggests initial priorities be placed on vulnerable species or populations (those already experiencing declines or projected to undergo ecological shifts resulting from climate changes that are consistent across climate projections) and species or populations for which ample data already exist (with the hope that these may inform climate change sensitivities in less well observed species or populations elsewhere). The sustained monitoring networks, novel observations, and modeling advances required to more confidently project marine mammal distributions in a changing climate will ultimately benefit management decisions across time-scales, further promoting the resilience of marine mammal populations.

Persistence of hotspots and variability of seabird species richness and abundance in the southern California Current

Aggregations of seabirds at sea may provide information on centers of enhanced trophic interactions and concentrating mechanisms, however, to date most studies lack quantification of persistence, a key hotspot characteristic. Persistence statistics may reduce uncertainty about seabird habitat use, improve understanding of the spatio-temporal scales of pelagic food web dynamics, and inform conservation planning. Using 26 years (1987–2012, 47 surveys) of shipboard surveys from a 300K km2 study area within the southern California Current Ecosystem, we conduct a spatial assessment of the inter-annual and seasonal dynamics of the persistence of seabird hotspots and identify recurring sites of elevated seabird species richness and abundance. Previous studies document declines in abundance, but were based on broad spatial standardizations to assess where declines may have occurred. Here, we refine the hypothesis that seabird populations have declined off southern California by focusing on persistently used habitats in nearshore or offshore domains. We demonstrate that spatio-temporal variability of seabird distribution and abundance is characterized by anomalous events embedded within trends. In addition to identifying the locations of persistence of seabird aggregations, we found significant declines in species richness and the density of sooty shearwater (Puffinus griseus) and Leachs storm petrel (Oceanodroma leucorhoa); in contrast, black-footed albatross (Phoebastria nigripes) abundance appear to be increasing. This assessment provides a spatially-explicit framework for future evaluations of biophysical drivers of seabird hotspots and their associations and impacts on forage fish and zooplankton populations.

Ecosystem Oceanography of Seabird Hotspots: Environmental Determinants and Relationship with Antarctic Krill Within an Important Fishing Ground

The discipline of ecosystem oceanography provides a framework for assessing the role of mesoscale physical processes on the formation and occurrence of biological hotspots. We used shipboard surveys over nine years to investigate environmental determinants of seabird hotspots near the Antarctic Peninsula, a region experiencing rapid climate change and an expanding krill fishery. We hypothesize that seabird hotspots are structured by mesoscale ocean conditions that reflect differences in prey distribution within oceanic and coastal waters. We used generalized additive models to quantify functional relationships of seabird hotspots with krill biomass, and a suite of remotely sensed environmental variables, such as eddy kinetic energy. The spatial organization, changes in intensity, and distribution shifts of seabird hotspots indicate different environmental drivers within coastal and oceanic domains and reflect the seasonal variability of the ecosystem. Our results indicate at least eight mesoscale hotspot zones that represent ecologically important areas where significant krill and predator biomass may be concentrated. Our ecosystem assessment of seabird hotspots identified critical foraging habitat and provided reference points to benefit research on estimating their trophic impacts on Antarctic ecosystems and potential effects from the krill fishery. Our approach is generally applicable to other pelagic ecosystems that are structured by hydrographic fronts and eddies, and containing schooling forage species shared by multiple wide-ranging predators. Furthermore, identification of biological hotspots is useful for the designation of marine protected areas most critical to potentially endangered wildlife and fisheries resources.

Modeling spatiotemporal dynamics of krill aggregations: size, intensity, persistence, and coherence with seabirds

Understanding aggregation dynamics of forage species is important for evaluating biophysical scaling in marine ecosystems and heterogeneity of trophic interactions. In particular, zooplankton aggregations are fundamental units of many pelagic systems, but are difficult to observe continuously through space and time. Using an established modeling framework that encompasses a coupled regional oceanographic and individual-based modeling system, we test the hypothesis that persistence (duration) of krill aggregations is dependent on their size, intensity, and location of formation within the coastal upwelling region of the California Current. In support of this hypothesis, we found that aggregation size is positively related to intensity, whereas persistence has a parabolic response to aggregation size and intensity, indicating the likelihood that large and highly persistent aggregations are rare. Persistence of krill aggregations also depends on formation location within coastal upwelling areas. We found that krill aggregations were more likely to form near a major seabird colony and that some coastal upwelling areas act as sources of aggregations for other areas. Observations of seabird aggregations were used to evaluate the potential structural realism of predicted krill aggregations. Seabird aggregations displayed marked coherence with predicted krill aggregations in space, providing important criteria on the scaling and availability of krill aggregations to breeding and migratory species. Predicting scales of krill aggregation dynamics will benefit ecosystem assessments, and numerical modeling of predator foraging and marine spatial management aimed at ensuring protection of ecologically important areas. This article is protected by copyright. All rights reserved.