Nitrogen and carbon stable isotope data sets are commonly used to assess complex population to ecosystem responses to natural or anthropogenic changes at regional to global spatial scales, and monthly to decadal timescales. Measured in the tissues of consumers, nitrogen isotopes (δ15 N) are primarily used to estimate trophic position while carbon isotopes (δ13 C) describe habitat associations and feeding pathways. Models of both δ15 N and δ13 C values and their associated variance can be used to estimate likely dietary contributions and niche width and provide inferences about consumer movement and migration. Stable isotope data have added utility when used in combination with other empirical data sets (e.g., stomach content, movement tracking, bioregionalization, contaminant, or fisheries data) and are increasingly relied upon in food web and ecosystem models. While numerous regional studies publish tables of mean δ15 N and δ13 C values, limited individual records have been made available for wider use. Such a deficiency has impeded full utility of the data, which otherwise would facilitate identification of macroscale patterns. The data provided here consist of 4,498 records of individuals of three tuna species, Thunnus alalunga, T. obesus, and T. albacares sampled from all major ocean basins from 2000 to 2015. For each individual tuna, we provide a record of the following: species name, sampling date, sampling location, tuna length, muscle bulk and baseline corrected δ15 N values, and muscle bulk and, where available, lipid corrected δ13 C values. We provide these individual records to support comparative studies and more robust modeling projects seeking to improve understanding of complex marine ecosystem dynamics and their responses to a changing environment. There are no copyright restrictions for research and/or teaching purposes. Users are requested to acknowledge their use of the data in publications, research proposals, websites, and other outlets following the citation instructions in Class III, Section B.

The eastern Pacific Ocean (EPO) covers ~ 55 million km2 of complex, dynamic habitats that can affect the distribution of ecologically and commercially important skipjack tuna, Katsuwonus pelamis, and its forage. The relationships between oceanography, ontogeny, and forage were quantitatively assessed to better understand skipjack tuna trophic ecology, and for parameterizing ecosystem models. Predator–prey dynamics of 320 skipjack tuna sampled from 1992 to 1994 and 2003 to 2005 were characterized—based on mean prey biomass (% $$\overline{W}$$ )—with respect to oceanographic biomes (Longhurst provinces), Oceanic Niño Index (ONI), year and skipjack tuna size, using a classification tree approach. Longhurst province was the most influential variable in explaining diet. Spatially, prey taxa richness was significantly lower, and taxonomic composition different, in predominant upwelling provinces compared to the primarily oligotrophic waters of the tropical offshore provinces. Anchovies comprised over 50% $$\overline{W}$$ of diet composition in the ‘upwelling’ provinces whereas various epipelagic (51% $$\overline{W}$$ ) and mesopelagic fishes (28% $$\overline{W}$$ ) were important in the ‘central American coastal and tropical offshore’ provinces. An ontogenetic breakpoint in diet was identified by the classification tree at 547 mm fork length, with small skipjack tuna primarily consuming anchovies, Humboldt squid and krill and large skipjack tuna consuming epipelagic fishes and red crab. Identifying skipjack tuna foraging arenas by Longhurst province can help facilitate the development of a spatially explicit ecosystem model, while defining ‘life stanzas’ as at the 547 mm breakpoint in the EPO will provide more realistic representations of diet composition, consumption rates, and losses to predation and fishing in such models.

Ecological risk assessment (ERA), including Productivity-Susceptibility Analysis (PSA), is becoming increasingly used to assess the relative vulnerability of data-limited non-target species to the impacts by fishing. PSA was developed for the eastern Pacific Ocean (EPO) tuna purse-seine fishery to assess the vulnerability of incidentally-caught species for three set types, “dolphin sets”, “unassociated sets” and “floating-object sets”, during 2005–2013. Because of operational differences between these set types, susceptibility values were combined for each species across the three set types to produce an overall fleet-wide susceptibility estimate. Vulnerability was highest for elasmobranchs, namely the giant manta ray, bigeye and pelagic thresher sharks, smooth and scalloped hammerhead sharks, and silky shark. Billfishes, dolphins, other rays, ocean sunfish, and yellowfin and bigeye tunas were classified as moderately vulnerable while the remaining species, all teleosts, had the lowest vulnerability scores. This purse-seine fleet-wide PSA identified potentially vulnerable species that can be compared with PSAs for other fisheries operating in the EPO, once detailed catch information becomes available for those fisheries. Such information can assist managers with prioritising fishery- and species-specific research programs and/or mitigation measures.

Considerable uncertainty remains over how increasing atmospheric CO2 and anthropogenic climate changes are affecting open-ocean marine ecosystems from phytoplankton to top predators. Biological time series data are thus urgently needed for the world's oceans. Here, we use the carbon stable isotope composition of tuna to provide a first insight into the existence of global trends in complex ecosystem dynamics and changes in the oceanic carbon cycle. From 2000 to 2015, considerable declines in δ13 C values of 0.8‰-2.5‰ were observed across three tuna species sampled globally, with more substantial changes in the Pacific Ocean compared to the Atlantic and Indian Oceans. Tuna recorded not only the Suess effect, that is, fossil fuel-derived and isotopically light carbon being incorporated into marine ecosystems, but also recorded profound changes at the base of marine food webs. We suggest a global shift in phytoplankton community structure, for example, a reduction in 13 C-rich phytoplankton such as diatoms, and/or a change in phytoplankton physiology during this period, although this does not rule out other concomitant changes at higher levels in the food webs. Our study establishes tuna δ13 C values as a candidate essential ocean variable to assess complex ecosystem responses to climate change at regional to global scales and over decadal timescales. Finally, this time series will be invaluable in calibrating and validating global earth system models to project changes in marine biota.

Aim We examined potential environmental drivers of broad-scale spatial patterns in the trophic structure of marine ecosystems as represented by nitrogen stable isotopes in globally distributed marine predators. Additionally, we assessed the effects of spatial scale on the predictive capabilities of environmental variables. LocationTime periodGlobal oceans. 2000 to 2015. Major taxa studiedMethodsTunas: Thunnus albacares, Thunnus obesus, Thunnus alalunga. We undertook a global compilation and meta-analysis of the bulk nitrogen stable isotope ratios (N-15 values) of three tuna species (n=4,281). After adjusting for regional variations in baseline N-15 values using a global ocean biogeochemistry model, generalized additive mixed models were employed to infer global-scale oceanographic controls of trophic structure, using cosmopolitan tuna species as a model. ResultsMain conclusionsFor the three tuna species, variation in trophic position estimated using bulk N-15 values was largely explained by geographical location and the corresponding oxygen minimum layer depth. Tuna trophic positions declined in areas with reduced oxygen at depth. Food-chain length, as captured by maximum trophic position, was longer in areas of the western Pacific Ocean and shorter in the northern Atlantic and eastern Pacific Oceans. Trophic adaptability of the tuna predators, as indicated by intraspecific variability, was highest in the western and central Pacific Ocean and lowest in the northern Atlantic Ocean. Our analysis demonstrated that while tunas share similar functional trophic roles, deeper-foraging tuna species had higher trophic positions globally. The predictive capacity of environmental variables decreased at finer (regional) spatial scales. Our work suggests that habitat compression resulting from the predicted global expansion of oxygen minimum zones with ocean warming will impact the trophic structure of marine food webs and the corresponding foraging habits of marine predators. Spatial scale analyses highlighted the importance of representing differences in regional ecological dynamics in global-scale trophic and ecosystem models.

Amino acid compound-specific isotope analysis (AA-CSIA) of tissue samples from four distinct trophic groups across the food web in the pelagic eastern tropical Pacific Ocean;

Predator-prey interactions for three commercially valuable tuna species: yellowfin (Thunnus albacares), bigeye (T. obesus), and albacore (T. alalunga), collected over a 40-year period from the Pacific, Indian, and Atlantic Oceans, were used to quantitatively assess broad, macro-scale trophic patterns in pelagic ecosystems. Analysis of over 14,000 tuna stomachs, using a modified classification tree approach, revealed for the first time the global expanse of pelagic predatory fish diet and global patterns of micronekton diversity. Ommastrephid squids were consistently one of the top prey groups by weight across all tuna species and in most ocean bodies. Interspecific differences in prey were apparent, with epipelagic scombrid and mesopelagic paralepidid fishes globally important for yellowfin and bigeye tunas, respectively, while vertically-migrating euphausiid crustaceans were important for albacore tuna in the Atlantic and Pacific Oceans. Diet diversity showed global and regional patterns among tuna species. In the central and western Pacific Ocean, characterized by low productivity, a high diversity of micronekton prey was detected while low prey diversity was evident in highly productive coastal waters where upwelling occurs. Spatial patterns of diet diversity were most variable in yellowfin and bigeye tunas while a latitudinal diversity gradient was observed with lower diversity in temperate regions for albacore tuna. Sea-surface temperature was a reasonable predictor of the diets of yellowfin and bigeye tunas, whereas chlorophyll-a was the best environmental predictor of albacore diet. These results suggest that the ongoing expansion of warmer, less productive waters in the world's oceans may alter foraging opportunities for tunas due to regional changes in prey abundances and compositions.

The effective evaluation of trophic interactions in pelagic food webs is essential for understanding food web ecology, conservation biology, and management. We tested the applicability of compound‐specific isotope analysis of amino acids (CSIA‐AA) for (1) characterizing trophic positions (TPs) of nine species from four trophic groups (tunas, squids, myctophids, and euphausiids) within a pelagic food web in the eastern tropical Pacific (ETP) Ocean, (2) evaluating trophic discrimination factors (TDFs) of each trophic group, and (3) detecting spatial changes in TPs and food chain length across a region with heterogeneous productivity. Although δ15N values of bulk tissues generally increased from south‐to‐north, CSIA‐AA revealed that trophic positions were uniform throughout our study area. These results suggest that variability in δ15N values were largely driven by nitrogen cycling dynamics in the ETP, which highlights the importance of these processes for the interpretation of δ15N values in food web studies. Absolute TP estimates were unrealistic for higher‐level species, and TDFs (tunas: 4.0‰, squids: 4.6‰, myctophids: 5.0‰, and euphausiids: 7.0‰) were lower than a widely used ecosystem TDF. We used remotely sensed oceanographic data to evaluate the physical oceanography and biological productivity throughout our study area and found significant relationships between δ15N values, nitrate concentrations, and SST across our study area. We did not find a gradient in phytoplankton cell size co‐occurring with an expected productivity gradient across our sampling region, which substantiated our isotope results indicating non‐significant spatial changes in TP and food chain length across the ETP.

We examined the stomach contents of 3 vertically migrating myctophid fish species from the eastern tropical Pacific (ETP) Ocean and used a classification tree to examine the influence of spatial, biological, and oceanographic predictor variables on diet. Myctophum nitidulum (n = 299), Symbolophorus reversus (n = 199), and Gonichthys tenuiculus, (n = 82) were collected with dip nets from surface waters, and prey taxa were quantified from bongo net tows from August through November 2006. A classification tree produced splits with longitude and sea surface salinity (SSS), thereby separating 3 geographically and oceanographically distinct regions of the ETP (offshore, nearshore, and intermediate), where diet was similar among the 3 species. Myctophids consumed, primarily, ostracods offshore (76.4% mean percentage by number [MNi]), euphausiids nearshore (45.0%), and copepods (66.6%) in the intermediate region. The offshore region was characterized by a greater abundance of ostracods in the zooplankton community (17.5% by number) and within a deep mixed- layer depth (MLD) (mean 52.6 m, max 93.0 m). SSS was low in the nearshore region (< 32.9 psu) and the MLD was shallow. The intermediate region represented a transition zone between the oceanographic condition of the offshore and nearshore regions. Our results indicate that these 3 myctophid species share a similar regional diet that is strongly influenced by longitude, ostracod availability, SSS, and MLD.

Tunas are highly specialized predators that have evolved numerous adaptations for a lifestyle that requires large amounts of energy consumption. Here we review our understanding of the bioenergetics and feeding dynamics of tunas on a global scale, with an emphasis on yellowfin, bigeye, skipjack, albacore, and Atlantic bluefin tunas. Food consumption balances bioenergetics expenditures for respiration, growth (including gonad production), specific dynamic action, egestion, and excretion. Tunas feed across the micronekton and some large zooplankton. Some tunas appear to time their life history to take advantage of ephemeral aggregations of crustacean, fish, and molluscan prey. Ontogenetic and spatial diet differences are substantial, and significant interdecadal changes in prey composition have been observed. Diet shifts from larger to smaller prey taxa highlight ecosystem-wide changes in prey availability and diversity and provide implications for changing bioenergetics requirements into the future. Where tunas overlap, we show evidence of niche separation between them; resources are divided largely by differences in diet percentages and size ranges of prey taxa. The lack of long-term data limits the ability to predict impacts of climate change on tuna feeding behaviour. We note the need for systematic collection of feeding data as part of routine monitoring of these species, and we highlight the advantages of using biochemical techniques for broad-scale analyses of trophic relations. We support the continued development of ecosystem models, which all too often lack the regional-specific trophic data needed to adequately investigate climate and fishing impacts.

Diet studies are an essential component of ecosystem-based approaches to fisheries management. In the eastern Pacific Ocean (EPO), the silky shark (Carcharhinus falciformis) is the most common species of shark in the bycatch of the tuna purse-seine fishery. A rare, comprehensive dataset of stomach contents of 786 silky sharks sampled in mostly tropical regions of the EPO (25°N–15°S; 79°W–162°W) during 1992–1994 and 2003–2005 was analyzed via classification tree and quantile regression methodologies to gain insight into its ecosystem role. Results suggest that the silky shark is an opportunistic predator that forages on a variety of prey. Broad-scale spatial and shark size covariates explained the feeding habits of silky sharks captured in sets on floating objects, primarily drifting fish-aggregating devices (FADs). A strong spatial shift in diet was identified by the tree analysis, with different foraging patterns in the eastern (inshore) and western (offshore) regions. Greater proportions of FAD-associated prey than non-FAD-associated prey were observed in the diet throughout the EPO, with the greatest proportion in the offshore region. Thus, silky sharks appear to take advantage of the associative behavior of prey fishes to increase their probability of encountering and capturing prey. Evaluation of prey–predator size relationships showed that maximum prey size increased with increasing silky shark size, but minimum prey size remained relatively constant across the range of shark sizes. Results such as these from spatially oriented analyses of predator feeding habits are essential for populating ecosystem models with space-based food webs, which otherwise suffer from generic representations of food webs.

Global-scale studies of marine food webs are rare, despite their necessity for examining and understanding ecosystem level effects of climate variability. Here we review the progress of an international collaboration that compiled regional diet datasets of multiple top predator fishes from the Indian, Pacific and Atlantic Oceans and developed new statistical methods that can be used to obtain a comprehensive ocean-scale understanding of food webs and climate impacts on marine top predators. We loosely define top predators not as species at the apex of the food web, but rather a guild of large predators near the top of the food web. Specifically, we present a framework for world-wide compilation and analysis of global stomach-contents and stable-isotope data of tunas and other large pelagic predatory fishes. To illustrate the utility of the statistical methods, we show an example using yellowfin tuna in a “test” area in the Pacific Ocean. Stomach-contents data were analyzed using a modified (bagged) classification tree approach, which is being prepared as an R statistical software package. Bulk δ15N values of yellowfin tuna muscle tissue were examined using a Generalized Additive Model, after adjusting for spatial differences in the δ15N values of the baseline primary producers predicted by a global coupled ocean circulation-biogeochemical-isotope model. Both techniques in tandem demonstrated the capacity of this approach to elucidate spatial patterns of variations in both forage species and predator trophic positions and have the potential to predict responses to climate change. We believe this methodology could be extended to all marine top predators. Our results emphasize the necessity for quantitative investigations of global-scale datasets when evaluating changes to the food webs underpinning top ocean predators under long-term climatic variability.

We review present understanding of the spatial and temporal diet variability (trophodynamics) of a range of pelagic marine top predators, at both early and adult life history stages. We begin with a review of methodologies used to advance our understanding of the trophodynamics of marine top predators, particularly in relation to climate change. We then explore how these developments are informing our understanding of the major trophic groups in food webs leading to, and including, marine top predators. We examine through specific examples how the impacts of ocean warming may affect pelagic food web relationships from both top-down and bottom-up perspectives. We examine the potential, in the absence of long-term data sets, of using large-scale spatial studies to examine how potential changes in biological oceanography could impact the biomass and composition of prey species, particularly the role of phytoplankton size spectra. We focus on examples from regions where biotic change with respect to climate change is likely. In particular, we detail the effects of climate change on oceanographic and bathymetric “hotspots” and provide the example involving seabirds in the Benguela Current system. We end by urging the development of international collaborations and databases to facilitate comprehensive ocean-scale understanding of climate impacts on marine top predators.

Widespread climate-induced habitat compression and reductions in biological production are beginning to alter food webs in tropical and subtropical oceans, but the effects on midtrophic level micronekton communities that support commercially important pelagic fishes are unclear. The predation habits of yellowfin tuna Thunnus albacares, a wide-ranging generalist predator with high energy requirements, provide rare insights into the distribution and availability of prey communities in pelagic regions. We used a modified classification tree approach to analyze spatial, temporal, environmental, and biological covariates explaining the predation patterns of 3362 yellowfin sampled across the eastern tropical Pacific Ocean (ETP) during two 2 yr periods occurring a decade apart. Persistent zoogeographical assemblages of prey were important only in relatively small subtropical regions at the extreme northern and southern ranges of the purseseine fishery for tunas. Prey biomass patterns for the majority of the samples over most of the ETP (6 degrees S to 17 degrees N, coast to 150 degrees W) were best explained by a tree partition distinguishing samples by sampling period, 1992 to 1994 and 2003 to 2005. The classification tree revealed that a major diet shift had transpired in the heart of the ETP. Yellowfin predation had changed from primarily larger epipelagic fish prey in the 1990s to a diverse array of smaller mesopelagic species and a crustacean that apparently had expanded its range in the 2000s. Partial dependence plots from the tree model showed range expansions previously described for some prey and unknown for other prey. Diet analysis of selected marine predators offers a practical means of monitoring prey communities poorly sampled by conventional methods.

Assessment of isotopic compositions at the base of food webs is a prerequisite for using stable isotope analysis to assess foraging locations and trophic positions of marine organisms. Our study represents a unique application of stable-isotope analyses across multiple trophic levels (primary producer, primary consumer and tertiary consumer) and over a large spatial scale in two pelagic marine ecosystems. We found that δ15N values of particulate organic matter (POM), barnacles and phenylalanine from the muscle tissue of yellowfin tuna all showed similar spatial patterns. This consistency suggests that isotopic analysis of any of these can provide a reasonable proxy for isotopic variability at the base of the food web. Secondly, variations in the δ15N values of yellowfin tuna bulk-muscle tissues paralleled the spatial trends observed in all of these isotopic baseline proxies. Variation in isotopic composition at the base of the food web, rather than differences in tuna diet, explained the 11‰ variability observed in the bulk-tissue δ15N values of yellowfin tuna. Evaluating the trophic position of yellowfin tuna using amino-acid isotopic compositions across the western Indian and equatorial Pacific Oceans strongly suggests these tuna occupy similar trophic positions, albeit absolute trophic positions estimated by this method were lower than expected. This study reinforces the importance of considering isotopic baseline variability for diet studies, and provides new insights into methods that can be applied to generate nitrogen isoscapes for worldwide comparisons of top predators in marine ecosystems.

Cephalopods are an important prey resource for fishes, seabirds, and marine mammals, and are also voracious predators on crustaceans, fishes, squid and zooplankton. Because of their high feeding rates and abundance, squids have the potential to exert control on the recruitment of commercially important fishes. In this review, we synthesize the available information for two intrinsic markers (δ15N and δ13C isotopic values) in squids for all oceans and several types of ecosystems to obtain a global view of the trophic niches of squids in marine ecosystems. In particular, we aimed to examine whether the trophic positions and trophic widths of squid species vary among oceans and ecosystem types. To correctly compare across systems, we adjusted squid δ15N values for the isotopic variability of phytoplankton at the base of the food web provided by an ocean circulation–biogeochemistry–isotope model. Studies that focused on the trophic ecology of squids using isotopic techniques were few, and most of the information on squids was from studies on their predators. Our results showed that squids occupy a large range of trophic positions and exploit a large range of trophic resources, reflecting the versatility of their feeding behavior and confirming conclusions from food-web models. Clear differences in both trophic position and trophic width were found among oceans and ecosystem types. The study also reinforces the importance of considering the natural variation in isotopic values when comparing the isotopic values of consumers inhabiting different ecosystems.