Characterizing dietary variability and trophic positions of coastal calanoid copepods: insight from stable isotopes and fatty acids

The spring zooplankton community in the Strait of Georgia (British Columbia, Canada) is characterized by the presence of several calanoid copepod species which collectively make up ~90% of the mezozooplankton biomass. Here, we investigate interspecific, interannual, and geographic variability in the diets and trophic positions of these copepods using a combination of fatty acids and stable isotopes. To characterize geographic variability in diet, we compare our findings from the Strait of Georgia with similar data from Ocean Station P in the subarctic northeast Pacific. Both fatty acid and stable isotope signatures indicate the existence of three trophic levels, even within the limited size range of these copepods: Neocalanus plumchrus and Calanus marshallae are primarily omnivorous, while Euchaeta elongata is carnivorous and Eucalanus bungii is herbivorous. Fatty acid markers of trophic position (e.g., DHA/EPA, 18:1n-9/18:1n-7) correlate significantly with δ¹⁵N, while markers indicating the proportion of diatoms to flagellates in the diet (e.g., 16PUFA/18PUFA and DHA/EPA) correlate significantly with δ¹³C, after the effect of lipid concentration on δ¹³C is accounted for. Despite the general correlation between stable isotopes and fatty acids, the former are not sensitive enough to capture the range of interannual variability observed in the latter, and can only capture substantial shifts in the diet over geographic scales. However, regardless of variability in food quality, the relative trophic positions of these copepods do not change significantly either spatially or temporally.     

Movement and environmental preferences of Greenland sharks (<Emphasis Type="Italic">Somniosus microcephalus</Emphasis>) electronically tagged in the St. Lawrence Estuary,Canada

Three Greenland sharks (Somniosus microcephalus) were tagged with electronic tags, in Baie St. Pancrace, St. Lawrence Estuary, Quebec, Canada. One shark was tagged on 23 July 2004, with an acoustic telemetry tag. Two sharks were each tagged with a pop-up satellite archival tag (PSAT) on 27 August 2004. Two of the sharks remained in or close to the bay, one for 47 days and the other for at least 66 days. The third shark left the bay immediately after tagging on 27 August 2004. This shark entered the main channel of the St. Lawrence Estuary, and had moved 114.9 km upstream by 1 November 2004 when the tag reported to ARGOS satellites. The tags provided a total of 179 days of data on the movement and environmental preferences of Greenland sharks in the St. Lawrence Estuary. Sharks that reported depth and ambient water temperature data from the bay showed significant diel differences in depth preferences and corresponding ambient temperatures. The sharks remained near the bottom of the water column during the day and displayed increased vertical movements at night. The shark that resided in the main channel did not show this pattern, but generally remained at depths between 325 and 352 m. Sharks in the bay experienced water temperatures that ranged from −1.1 to 8.6°C at depths from 0 to 67 m. In the main channel the shark experienced temperatures that ranged from 1.0 to 5.4°C at depths from 132 to 352 m. This is the first report of numerous Greenland sharks inhabiting shallow near shore bays during summer and autumn.     

Fine-scale population structure of estuarine bottlenose dolphins (Tursiops truncatus) assessed using stable isotope ratios and fatty acid signature analyses

Stable isotope ratios and fatty acid signature analyses were employed to examine the fine-scale population structure of a year-round resident population of 600–800 bottlenose dolphins (Tursiops truncatus) in the Indian River Lagoon (IRL), Florida. The IRL, a 250-km-long estuary running along the central east coast of Florida (28.0°N, 80.6°W), is comprised of the northern and southern IRL, Mosquito Lagoon (ML), Banana River (BR), and St. Lucie Estuary. Samples of skin and blubber were collected from dead stranded (n = 61, 1994–2004) and live dolphins (n = 153, 2002–2007, 2010, 2011) from throughout the IRL and surrounding environs. Using stable isotopes (SI), dolphins could be assigned to a ML subpopulation, a St. Lucie Estuary subpopulation, and an IRL subpopulation. Fatty acid signature analysis (FASA) allowed for finer resolution, detecting ML and BR subpopulations, a separation of northern and southern IRL subpopulations, and a St. Lucie Estuary subpopulation. Differences between sexes were detectable within subpopulations using FASA, but not using SI. This may indicate that males and females are foraging in similar locations at a similar trophic level (detected using SI), but are varying in the types or proportions of specific prey (indicated by FASA). The combination of these complementary analyses results in a powerful tool for assessing fine-scale population substructure.     

The role of stable isotopes and mercury concentrations to describe seabird foraging ecology in tropical environments

Nitrogen (δ¹⁵N) and carbon (δ¹³C) stable isotopes and contaminants, such as mercury, have been widely used to characterise foraging ecology of temperate and polar seabirds. In this study, for the first time, we used isotopic signatures and mercury levels of feathers and blood of eight tropical seabird species, that forage in a range-gradient between inshore and offshore areas, to describe the foraging habits of a large tropical seabird community (from two neighboring islands of the Seychelles archipelago, western Indian Ocean) during both the breeding and inter-breeding periods. Overall, we found a high overlap in both δ¹⁵N and δ¹³C signatures among species. The high inter-specific overlap in δ¹⁵N values was expected, given the similarities in the diet of the species from this community. However, several unexpected results, such as (1) the consistently higher δ¹⁵N signatures of white terns (Gygis alba), (2) the large variation in inter-specific differences in δ¹⁵N signatures among the sampling groups (season, age, island and tissue) and (3) the consistent low δ¹⁵N values of breeding birds during the northwest monsoon (austral summer), suggest that δ¹⁵N signatures cannot be used as indicators of seabird trophic levels in this community. The high inter-specific overlap in δ¹³C signatures and the absence, during the breeding season, of a δ¹³C gradient that follows the inshore-offshore foraging gradient within the community can be explained by the habitat homogeneity of the Seychelles continental shelf and suggest that birds forage mostly within the limits of this “plateau”. On the other hand, the similarities in δ¹³C values between the breeding and inter-breeding periods in species that are known to show post-breeding dispersal, strongly support the hypothesis of a lack of latitudinal variation in δ¹³C signatures of POM in the central Indian Ocean, and the consequent inaccuracy of δ¹³C values to track seabird movements within this geographic area. Inter-specific differences in mercury levels seem to be related to prey size, while consistent higher mercury concentrations in one of the studied islands suggest different island mercury-backgrounds and possible segregation in foraging areas between the seabirds of the two islands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Under-ice amphipods in the Greenland Sea and Fram Strait (Arctic): environmental controls and seasonal patterns below the pack ice

During three "Polarstern" cruises to the ice-covered Greenland Sea (spring 1997, summer 1994, autumn 1995) studies on the under-ice habitat (morphology, hydrography, ice-algal biomass) and on the macrofaunal, autochthonous under-ice amphipods (species diversity, abundance) were carried out in order to describe environmental controls and seasonal patterns in this community. In spring, the ice underside was rather smooth and whitish, while in summer melting structures and sloughed-off ice-algal threads were observed, in autumn detritus clumps accumulated in depressions at the ice underside. Only in summer, a thin layer of warm (up to -0.6°C) and less saline (as low as S=6.3) water was found at the ice-water interface above Polar Water. Integrated ice-algal biomass was highest during autumn (2.6 mg chl a m^-2) and lowest during summer (1.2 mg chl a m^-2). Four species of under-ice amphipods occurred in spring and summer (Apherusa glacialis, Onisimus glacialis, O. nanseni, Gammarus wilkitzkii), but only the last species was observed at the ice underside in autumn. A. glacialis and G. wilkitzkii were equally abundant in spring; A. glacialis dominated in summer. The highest total abundance of amphipods occurred during summer (31.9 ind. m^-2), compared to lower abundances in spring and autumn (5.3 and 1.1 ind. m^-2, respectively). A factor analysis revealed seasonal patterns in the data set, which mainly influenced A. glacialis, and species-specific relations between several environmental factors and the distribution of under-ice amphipods. Abundance of A. glacialis was closely related to the under-ice hydrography and ice-algal biomass, whereas the other amphipod species were more influenced by the under-ice morphology. It is therefore stated that the observed thinning of the Arctic sea ice and the resulting increased meltwater input and change in morphology of floes will have a profoundly adverse effect on the under-ice amphipods.     

The emergence of stable isotopes in environmental and forensic geochemistry studies: a review

In the past decade, environmental forensics has emerged as a discipline directed toward determining parties liable for causing spills of contaminants into the environment. Such investigations, while geared toward determining the guilty parties in order to recover costs of the cleanup and remediation, require various questions to be addressed. These include determination of the nature of the product; Where did it come from?; Extent of weathering, if any; How long has it been there?; and Is it degrading naturally? Traditionally, these studies have been addressed through utilization of techniques such as gas chromatography (GC) and gas chromatography–mass spectrometry (GCMS). However, in recent years, stable isotopes, primarily determined through the use of combined gas chromatography–isotope ratio mass spectrometry (GCIRMS), have emerged as an equally important tool in environmental forensics. For relatively low molecular, volatile compounds such as MTBE, BTEX, or chlorinated solvents, the isotopes, primarily carbon and hydrogen, have been used extensively for evaluating the onset of natural attenuation. For larger molecules such as PCBs or PAHs, in which the effects of biodegradation on the isotope composition of these molecules is minimal, the isotopic fingerprints of the individual compounds can be used for correlation purposes. In this paper, a brief introduction to isotope geochemistry will be given, followed by a review of applications of stable isotopes to a variety of environmental problems. While the review may not necessarily be exhaustive, it will provide a comprehensive overview of areas where isotopes have been used and potential applications for the future. Most of the review is concerned with carbon and hydrogen isotopes, although a brief overview of the emerging area of chlorine isotopes will also be discussed.     

Oxygen and carbon stable isotopes in the otoliths of wild and laboratory-reared Australian salmon (Arripis trutta)

Australian salmon,Arripis trutta, collected from the east coast of Tasmania, Australia, in 1987, were weighed and measured and their otoliths marked by immersing fish in an oxytetracycline hydrochloride/seawater solution before placement in constant-temperature aquaria. Individual somatic and otolith growth rates were determined for input into mass balance models. Mass balance models were used to determine the oxygen and carbon isotopic composition of otolith material produced during captivity. There was a significant relationship between ¹⁸O measured in the otolith aragonite and ambient temperature (r ² = 0.77). The linear relationship between these data, where ¹⁸O = 6.69 – 0.326 (T, °C), was not significantly different from a relationship indicative of equilibrium deposition of oxygen isotopes in aragonite. Otolith carbon was significantly depleted in¹³C relative to equilibrium deposition, with depletions >6.0 at all temperatures. There was no relationship between ¹³C and temperature. It was estimated that >30% of the otolith carbon was from metabolically derived sources. Significant differences in otolith carbon isotopes among wild juvenile Australian salmon were hypothesised to be attributable to differences in diet. Levels of variability for both oxygen and carbon isotopes in laboratory-maintained and wild fish were similar to that found by other researchers for foraminifera and these results highlight the importance of large sample sizes when estimating environmental temperatures from oxygen isotopes measured in fish otoliths.     

Potential correlation between perfluorinated acids and liver morphology in East Greenland polar bears (Ursus maritimus)

In this study, residual concentrations of chlorpyrifos (CPF) were determined in feed (40) and fodder (25) samples collected from various locations of Tarai region of Uttarakhand. For extracting residues, liquid–liquid partition followed by alumina column clean up was used and the detection and quantification of residues was undertaken with the help of high-performance liquid chromatography using C18 column and diode array detector at 220?nm. Of the total 40 feed samples analyzed, 7 (17.5%) samples were found positive for CPF with the mean residual concentration of 0.058?µg?g^?1; while out of 25 fodder samples, CPF residues were detected in a single (4%) sample with residual concentration of 0.39?µg?g^?1. However, none of the feed or fodder samples contained CPF residues above the prescribed limit.     

Bat-cave catchment areas: using stable isotopes (deltaD) to determine the probable origins of hibernating bats

The application of stable hydrogen isotope (deltaD) techniques has swiftly advanced our understanding of animal movements, but this progression is dominated by studies of birds and relatively long-distance, north-south migrants. This dominance reflects the challenge of incorporating multiple sources of error into geographic assignments and the nature of spatially explicit deltaD models, which possess greater latitudinal than longitudinal resolution. However, recent progress in likelihood-based assignments that incorporate multiple sources of isotopic error and Bayesian approaches that include additional sources of information may advance finer-scale understanding of animal movements. We develop a stable-isotope method for determining probable origins of bats within hibernacula and show that this method produces spatially explicit, continuous assignments with regional resolution. We outline how these assignments can be used to infer hibernacula connectivity, an application that could inform spatial modeling of white-nose syndrome. Additionally, estimates of seasonal and annual flight distances for many cave-dwelling bat species can be derived from this approach. We also discuss how this application can be used in general to provide insights into variable migratory and foraging strategies within bat populations.     

The role of water in the information exchange between the components of an ecosystem

Living organisms and ecosystems have been shown to be sensitive to very weak signals originating very far away. The dynamics governing these phenomena is discussed in the framework of Quantum Field Theory. This phenomenon gives an indication on the dynamics responsible for the exchange of information in ecosystems. The peculiar role of coherent water is stressed. It is shown that energy is able to travel in a coherent medium in form of solitons, without any losses.     

Movements of blue sharks (Prionace glauca) in depth and course

Acoustic telemetry was used to follow 22 blue sharks,Prionace glauca (Linnaeus), over the continental shelf and slope in the region between George's Bank and Cape Hatteras between 1979 and 1986. The sharks frequently made vertical excursions between the surface and depths of several hundred meters. The oscillations, which were repeated every few hours, were largest in the daytime and were smaller in amplitude and confined to depths near the thermocline at night. This behavior was prominent in trials from August through March, but was not seen from June through July. Diving is discussed in terms of a hunting tactic and behavioral thermoregulation. Most of the sharks moved in a southeasterly direction from the release point and many maintained a constant course day and night for several days. The sharks may orient to the earth's magnetic field, or to the ocean's electric fields, allowing them to swim on a constant heading in the absence of celestial cues. These possibilities are discussed in the appendix.     

Satellite tagging of blue sharks (Prionace glauca) and other pelagic sharks off eastern Australia: depth behaviour, temperature experience and movements

Satellite telemetry was used to study the movements and behaviour of ten blue sharks and one individual each of shortfin mako, thresher and bigeye thresher off eastern Australia. The tracks showed latitudinal movements of up to 1,900 km, but none of the sharks travelled away from the eastern Australian region. Tracking periods did not exceed 177 days. All species showed oscillatory dive behaviour between the surface layers to as deep as 560–1,000 m. Blue sharks spent 35–58% of their time in <50 m depths and 10–16% of their time in >300 m. Of these four species, the bigeye thresher spent the least time in the surface layers and the most time at >300 m depth. All four species showed clear diel behaviour generally occupying shallower depths at night than during the day. Blue sharks were mainly in 17.5–20.0°C water, while the thresher sharks showed a more bimodal temperature distribution.     

Distribution of foraging habitats of male loggerhead turtles (Caretta caretta) as revealed by stable isotopes and satellite telemetry

Most studies on the foraging ecology of loggerhead turtles (Caretta caretta) have focused on adult females and juveniles. Little is known about the foraging patterns of adult male loggerheads. We analyzed tissues for carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N) from 29 adult male loggerheads tracked with satellite transmitters from one breeding area in Florida, USA, to evaluate their foraging habitats in the Northwest Atlantic (NWA). Our study revealed large variations in δ¹³C and δ¹⁵N and a correlation between both δ¹³C and δ¹⁵N and the latitude to which the loggerheads traveled after the mating season, thus reflecting a geographic pattern in the isotopic signatures. Variation in δ¹³C and δ¹⁵N can be explained by differences in food web baseline isotopic signatures rather than differences in loggerhead trophic levels. Stable isotope analysis may help elucidate residency and migration patterns and identify foraging sea turtle subpopulations in the NWA due to the isotopically distinct habitats used by these highly migratory organisms.     

Movements and swimming behaviour of white sharks (Carcharodon carcharias) in Australian waters

We used a combination of satellite telemetry, archival and conventional tags to show that white sharks made broad-scale movements consistent with mixing of the population across their entire Australasian range. The capture of one of these sharks in New Zealand, some 3,550 km from the point of tagging in South Australia, provides further confirmation that white sharks sometimes move into open ocean waters and cross deep ocean basins. However, most movements were confined to shelf waters, generally in areas of less than 100 m depth and in some cases into waters of less than 5 m depth. Sharks showed considerable plasticity in swimming patterns, which included many of the behaviours reported for other species. One of the archival-tagged sharks showed separate periods of distinct swimming behaviour as it moved into different habitats and travelled between them. The changes in swimming behaviour were abrupt and suggested rapid switching of hunting strategies for different prey types in these habitats. All tracked sharks showed both prolonged periods of directional swimming in coastal waters at swimming speeds of 2–3 km h⁻¹ as well as temporary residency in particular regions. Movements of tagged white sharks, together with data from shark control programs and bycatch records, suggest a seasonal movement northward along the east coast of Australia during the autumn–winter months and south in spring–early summer. The consistency of paths taken by white sharks in Australian waters suggests that they may follow common routes or “highways” in some areas. If so, identifying such areas may assist in reducing interactions with fishing operations and thus reduce bycatch.     

Influence of nutrients in the feeding ecology of seagrass (Posidonia oceanica L.) consumers: a stable isotopes approach

Nitrogen inputs to coastal environments can considerably alter the abundance of primary producers. However, how herbivores modify their trophic signatures and adjust to changes in food resource conditions remains controversial. Here, we assess the effect of nutrient availability on the diet shifts of the two main Mediterranean herbivores, the Sparid fish Sarpa salpa L. and the sea urchin Paracentrotus lividus (Lmk.) that feed mostly on the seagrass Posidonia oceanica L. (Delile), epiphytes and benthic macroalgae. To do this, we (1) investigate the patterns of isotopic composition (δ¹³C and δ¹⁵N signatures) of the two herbivores and their potential food sources in three areas of contrasting nutrient conditions and, (2) we assess the diet shift along this nutrient gradient by estimating the isotopic nutrient enrichment (i.e., the contribution of δ¹³C and δ¹⁵N signatures in consumers’ tissues relative to potential food sources). Food web signatures of δ¹³C were similar among the three study sites, and no patterns of δ¹³C shift were observed in their diets. In contrast, there was a consistent increase in N contents and δ¹⁵N along the nutrient gradient for all primary producers and their consumers. The rate of δ¹⁵N enrichment was also clearly distinctive between the two herbivores: in P. lividus it increased by 61% along the nutrient gradient, while in S. salpa it remained constant. Our results suggest that sea urchins behave as facultative omnivores and feed on vegetable or mixed diets depending on the trophic status of the system. It is unclear, however, if this modification is behavioral or the consequence of mere changes in the availability of food items, as animal epiphytes (e.g., hydrozoans, bryozoans and ascidians) can also became more abundant on seagrass leaves under increased nutrient conditions. In contrast, adult fish appear to feed on vegetal material independent of nutrient availability in the ecosystem.     

Habitat use and foraging behavior of tiger sharks (<Emphasis Type="Italic">Galeocerdo cuvier</Emphasis>) in a seagrass ecosystem

Understanding the foraging behavior and spatial distribution of top predators is crucial to gaining a complete understanding of communities. However, studies of top predators are often logistically difficult and it is important to develop appropriate methods for identifying factors influencing their spatial distribution. Sharks are top predators in many marine communities, yet no studies have quantified the habitat use of large predatory sharks or determined the factors that might influence shark spatial distributions. We used acoustic telemetry and animal-borne video cameras ("Crittercam") to test the hypothesis that tiger shark (Galeocerdo cuvier) habitat use is determined by the availability of their prey. We also used Crittercam to conduct the first investigation of foraging behavior of tiger sharks. To test for habitat preferences of sharks, the observed proportion of time in each habitat for each individual was compared to the predicted values for that individual based on correlated random walk and track randomization methods. Although there was individual variation in habitat use, tiger sharks preferred shallow seagrass habitats, where their prey is most abundant. Despite multiple encounters with potential prey, sharks rarely engaged in prolonged high-speed chases, and did not attack prey that were vigilant. We propose that the tiger sharks' foraging tactic is one of stealth, and sharks rely upon close approaches to prey in order to be successful. This study shows that using appropriate analysis techniques and a variety of field methods it is possible to elucidate the factors influencing habitat use and gain insights into the foraging behavior of elusive top predators.     

Chemotaxonomy ofGyrocotyle (Platyhelminthes: Cercomeria) species,parasites of chimaerid fish (Holocephali), by chemometry of their fatty acids

The validity of some morphological gyrocotylid species, exclusive gut parasites of ratfishes (Holocephali), is contested. Non-morphological characters, such as fatty acid profiles, may be used to resolve taxonomic problems. FiveGyrocotyle species were studied: three inChimaera monstrosa collected between 1985 and 1987 from the north-east Atlantic (on the Norwegian coast); and two inHydrolagus colliei, collected in 1987 from the north-east Pacific (in Nanaimo, British Columbia, Canada). Their fatty acids were obtained by methanolytic extraction of worm tissue samples, followed by gas chromatography of the fatty acid methyl esters, and finally multivariate data treatment (principal component analysis) of gas chromatographic results. Complete separation of the five species was obtained; classification agrees well with that arrived at by morpholog and enzyme electrophoresis.     

Essential fatty acids influence metabolic rate and tolerance of hypoxia in Dover sole (Solea solea) larvae and juveniles

Dover sole (Solea solea, Linneaus 1758) were raised from first feeding on brine shrimp (Artemia sp.) with different contents and compositions of the essential fatty acids (EFA) arachidonic acid (ARA, 20:4n − 6); eicosapentaenoic acid (EPA, 20:5n − 3), and docosahexaenoic acid (DHA, 22:6n − 3), and their metabolic rate and tolerance to hypoxia measured prior to and following metamorphosis and settlement. Four dietary Artemia preparations were compared: (1) un-enriched; (2) enriched with a commercial EFA mixture (Easy DHA SELCO Emulsion); (3) enriched with a marine fish oil combination (VEVODAR and Incromega DHA) to provide a high ratio of ARA to DHA, and (4) enriched with these fish oils to provide a low ratio of ARA to DHA. Sole fed un-enriched Artemia were significantly less tolerant to hypoxia than the other dietary groups. Larvae from this group had significantly higher routine metabolic rate (RMR) in normoxia, and significantly higher O₂ partial pressure (PO₂) thresholds in progressive hypoxia for their regulation of RMR (P _crit) and for the onset of agitation, respiratory distress and loss of equilibrium. Metamorphosis was associated with an overall decline in RMR and increase in P _crit, but juveniles fed on un-enriched Artemia still exhibited higher P _crit and agitation thresholds than the other groups. Sole fed un-enriched Artemia had significantly lower contents of EFA in their tissues, both before and after settlement. Thus, enriching live feeds with EFA has significant effects on the respiratory physiology of sole early life stages and improves their in vivo tolerance to hypoxia. We found no evidence, however, for any effect of the ratio of ARA to DHA.