David G. Foley

Tracking apex marine predator movements in a dynamic ocean

Pelagic marine predators face unprecedented challenges and uncertain futures. Overexploitation and climate variability impact the abundance and distribution of top predators in ocean ecosystems. Improved understanding of ecological patterns, evolutionary constraints and ecosystem function is critical for preventing extinctions, loss of biodiversity and disruption of ecosystem services. Recent advances in electronic tagging techniques have provided the capacity to observe the movements and long-distance migrations of animals in relation to ocean processes across a range of ecological scales. Tagging of Pacific Predators, a field programme of the Census of Marine Life, deployed 4,306 tags on 23 species in the North Pacific Ocean, resulting in a tracking data set of unprecedented scale and species diversity that covers 265,386 tracking days from 2000 to 2009. Here we report migration pathways, link ocean features to multispecies hotspots and illustrate niche partitioning within and among congener guilds. Our results indicate that the California Current large marine ecosystem and the North Pacific transition zone attract and retain a diverse assemblage of marine vertebrates. Within the California Current large marine ecosystem, several predator guilds seasonally undertake north–south migrations that may be driven by oceanic processes, species-specific thermal tolerances and shifts in prey distributions. We identify critical habitats across multinational boundaries and show that top predators exploit their environment in predictable ways, providing the foundation for spatial management of large marine ecosystems.

Migratory shearwaters integrate oceanic resources across the Pacific Ocean in an endless summer

Electronic tracking tags have revolutionized our understanding of broad-scale movements and habitat use of highly mobile marine animals, but a large gap in our knowledge still remains for a wide range of small species. Here, we report the extraordinary transequatorial postbreeding migrations of a small seabird, the sooty shearwater, obtained with miniature archival tags that log data for estimating position, dive depth, and ambient temperature. Tracks (262 ± 23 days) reveal that shearwaters fly across the entire Pacific Ocean in a figure-eight pattern while traveling 64,037 ± 9,779 km roundtrip, the longest animal migration ever recorded electronically. Each shearwater made a prolonged stopover in one of three discrete regions off Japan, Alaska, or California before returning to New Zealand through a relatively narrow corridor in the central Pacific Ocean. Transit rates as high as 910 ± 186 km·day−1 were recorded, and shearwaters accessed prey resources in both the Northern and Southern Hemisphere’s most productive waters from the surface to 68.2 m depth. Our results indicate that sooty shearwaters integrate oceanic resources throughout the Pacific Basin on a yearly scale. Sooty shearwater populations today are declining, and because they operate on a global scale, they may serve as an important indicator of climate change and ocean health.

Biological enhancement at cyclonic eddies tracked with GOES Thermal Imagery in Hawaiian waters

In November 1999, ship and satellite platforms were used to track, target and map the vertical and horizontal structure of two cyclonic eddies in the lee of Hawaii. Depth-integrated nitrate+nitrite levels within the photic zone of the eddy cores were 3- to 15- fold higher than those observed for control stations. The depth and magnitude of the deep chlorophyll maximum were significantly elevated in the more mature of the two eddies. HPLC analyses revealed that the enhanced chlorophyll was largely contributed by chromophyte microalgae. Modeled primary productivity rates were up to twofold higher at the stations within the eddies.

Zooplankton variability on the equator at 140°W during the JGOFS EqPac study

The vertical distributions of zooplankton biomass and community composition were measured on the equator at 140°W during March/April (19 days) and October 1992 (21 days). El Nino conditions prevailed during the March/April time series. The average integrated (200 m) zooplankton (>64 μm) biomass was 32 mM C m−2 in March/April and 41 mM C m−2 in October. The overall cruise means were not significantly different; however during October zooplankton were more aggregated in the upper euphotic zone, there were more copepods >200 μm, and there were more day/night differences in the vertical distribution of zooplankton biomass as a result of vertical migration behavior. Cyclopoid copepods were more abundant during the El Nino conditions of March compared to October. There were no apparent trends in integrated zooplankton biomass during the March/April time series. However, the species composition of the zooplankton community changed towards the end of the time series when meridional currents flowed to the south. Zooplankton biomass during the October cruise increased nearly five-fold with the passage of a tropical instability wave. A lag in the increase of zooplankton compared to nitrate and chlorophyll as well as changes in the zooplankton community composition over the time series reflect the response times of zooplankton to upwelling events.

Longwaves and primary productivity variations in the equatorial pacific at 0°, 140dgW

Abstract High temporal resolution measurements of physical and bio-optical variables were made in the upper ocean using a mooring located at 0°, 140°W from 9 February 1992 to 15 March 1993 as part of the equatorial Pacific Ocean (EgPac) study. Chlorophyll and primary productivity time-series records were generated using the mooring data. Primary productivity varied by about 50% around the mean on time scales of weeks and by over a factor of four within our observational period. The mooring observations encompassed both El Niho and cool conditions. Kelvin waves were evident during the El Nifio phase, and tropical instability waves (TIWs) were dominant during the cool phase. The two extreme conditions also were observed concurrently with complementary ship-based measurements. In addition, bio-optical drifters provided simultaneous spatial data concerning net phytoplankton growth rates during passage of a TIW. The collective data sets have been used to examine the causes of the observed variability in phytoplankton biomass and productivity. Our joint results and analyses appear to support the hypothesis that the vertical transport of iron into the upper layer and primary production rates are modulated by variability of the depth of the Equatorial Undercurrent and by equatorial longwaves. In particular, our results are consonant with the suggestion of Barber et al. (1996) that passage of a TIW may be considered to be a natural analog of a small iron enrichment experiment. Predicting primary productivity and, thus, carbon flux in the equatorial Pacific requires continuous, long-term observations of a few physical, biological, and optical properties that can be used to parameterize the biological variability.

Influence of a cyclonic eddy on microheterotroph biomass and carbon export in the lee of Hawaii

heterotroph biomass and 234 Th-derived carbon export rates within the eddy were 2 to 3 times higher than those observed for adjacent waters. If this eddy is representative of other cyclonic eddies that are frequently formed in the lee of Hawaii, then eddy activity may significantly enhance the areal efficiency of the biological pump and facilitate the transfer of organic carbon to organisms inhabiting the mesopelagic and abyssal-benthic zones of this subtropical ecosystem. INDEX TERMS: 4520 Oceanography: Physical: Eddies and mesoscale processes; 4806 Oceanography: Biological and Chemical: Carbon cycling; 4855 Oceanography: Biological and Chemical:Plankton;4866Oceanography:BiologicalandChemical: Sorptive scavenging; 4870 Oceanography: Biological and Chemical: Stable isotopes. Citation: Bidigare, R. R., C. BenitezNelson, C. L. Leonard, P. D. Quay, M. L. Parsons, D. G. Foley, and M. P. Seki, Influence of a cyclonic eddy on microheterotroph biomass and carbon export in the lee of Hawaii,Geophys.Res.Lett., 30(6), 1318, doi:10.1029/2002GL016393, 2003.

Quantifying climatological ranges and anomalies for Pacific coral reef ecosystems.

Coral reef ecosystems are exposed to a range of environmental forcings that vary on daily to decadal time scales and across spatial scales spanning from reefs to archipelagos. Environmental variability is a major determinant of reef ecosystem structure and function, including coral reef extent and growth rates, and the abundance, diversity, and morphology of reef organisms. Proper characterization of environmental forcings on coral reef ecosystems is critical if we are to understand the dynamics and implications of abiotic–biotic interactions on reef ecosystems. This study combines high-resolution bathymetric information with remotely sensed sea surface temperature, chlorophyll-a and irradiance data, and modeled wave data to quantify environmental forcings on coral reefs. We present a methodological approach to develop spatially constrained, island- and atoll-scale metrics that quantify climatological range limits and anomalous environmental forcings across U.S. Pacific coral reef ecosystems. Our results indicate considerable spatial heterogeneity in climatological ranges and anomalies across 41 islands and atolls, with emergent spatial patterns specific to each environmental forcing. For example, wave energy was greatest at northern latitudes and generally decreased with latitude. In contrast, chlorophyll-a was greatest at reef ecosystems proximate to the equator and northern-most locations, showing little synchrony with latitude. In addition, we find that the reef ecosystems with the highest chlorophyll-a concentrations; Jarvis, Howland, Baker, Palmyra and Kingman are each uninhabited and are characterized by high hard coral cover and large numbers of predatory fishes. Finally, we find that scaling environmental data to the spatial footprint of individual islands and atolls is more likely to capture local environmental forcings, as chlorophyll-a concentrations decreased at relatively short distances (>7 km) from 85% of our study locations. These metrics will help identify reef ecosystems most exposed to environmental stress as well as systems that may be more resistant or resilient to future climate change.

The Making of a Productivity Hotspot in the Coastal Ocean

Background Highly productive hotspots in the ocean often occur where complex physical forcing mechanisms lead to aggregation of primary and secondary producers. Understanding how hotspots persist, however, requires combining knowledge of the spatio-temporal linkages between geomorphology, physical forcing, and biological responses with the physiological requirements and movement of top predators. Methodology/Principal Findings Here we integrate remotely sensed oceanography, ship surveys, and satellite telemetry to show how local geomorphology interacts with physical forcing to create a region with locally enhanced upwelling and an adjacent upwelling shadow that promotes retentive circulation, enhanced year-round primary production, and prey aggregation. These conditions provide an area within the upwelling shadow where physiologically optimal water temperatures can be found adjacent to a region of enhanced prey availability, resulting in a foraging hotspot for loggerhead sea turtles (Caretta caretta) off the Baja California peninsula, Mexico. Significance/Conclusions We have identified the set of conditions that lead to a persistent top predator hotspot, which increases our understanding of how highly migratory species exploit productive regions of the ocean. These results will aid in the development of spatially and environmentally explicit management strategies for marine species of conservation concern.

Combined climate‐ and prey‐mediated range expansion of Humboldt squid (Dosidicus gigas), a large marine predator in the California Current System

Climate-driven range shifts are ongoing in pelagic marine environments, and ecosystems must respond to combined effects of altered species distributions and environmental drivers. Hypoxic oxygen minimum zones (OMZs) in midwater environments are shoaling globally; this can affect distributions of species both geographically and vertically along with predator-prey dynamics. Humboldt (jumbo) squid (Dosidicus gigas) are highly migratory predators adapted to hypoxic conditions that may be deleterious to their competitors and predators. Consequently, OMZ shoaling may preferentially facilitate foraging opportunities for Humboldt squid. With two separate modeling approaches using unique, long-term data based on in situ observations of predator, prey, and environmental variables, our analyses suggest that Humboldt squid are indirectly affected by OMZ shoaling through effects on a primary food source, myctophid fishes. Our results suggest that this indirect linkage between hypoxia and foraging is an important driver of the ongoing range expansion of Humboldt squid in the northeastern Pacific Ocean.

Identification of distinct movement patterns in Pacific leatherback turtle populations influenced by ocean conditions

Interactions with fisheries are believed to be a major cause of mortality for adult leatherback turtles (Dermochelys coriacea), which is of particular concern in the Pacific Ocean, where they have been rapidly declining. In order to identify where these interactions are occurring and how they may be reduced, it is essential first to understand the movements and behavior of leatherback turtles. There are two regional nesting populations in the East Pacific (EP) and West Pacific (WP), comprising multiple nesting sites. We synthesized tracking data from the two populations and compared their movement patterns. A switching state-space model was applied to 135 Argos satellite tracks to account for observation error, and to distinguish between migratory and area-restricted search behaviors. The tracking data, from the largest leatherback data set ever assembled, indicated that there was a high degree of spatial segregation between EP and WP leatherbacks. Area-restricted search behavior mainly occurred in the southeast Pacific for the EP leatherbacks, whereas the WP leatherbacks had several different search areas in the California Current, central North Pacific, South China Sea, off eastern Indonesia, and off southeastern Australia. We also extracted remotely sensed oceanographic data and applied a generalized linear mixed model to determine if leatherbacks exhibited different behavior in relation to environmental variables. For the WP population, the probability of area-restricted search behavior was positively correlated with chlorophyll-a concentration. This response was less strong in the EP population, but these turtles had a higher probability of search behavior where there was greater Ekman upwelling, which may increase the transport of nutrients and consequently prey availability. These divergent responses to oceanographic conditions have implications for leatherback vulnerability to fisheries interactions and to the effects of climate change. The occurrence of leatherback turtles within both coastal and pelagic areas means they have a high risk of exposure to many different fisheries, which may be very distant from their nesting sites. The EP leatherbacks have more limited foraging grounds than the WP leatherbacks, which could make them more susceptible to any temperature or prey changes that occur in response to climate change.