Stable isotope profiles from subtropical marine gastropods of the family Fasciolariidae: growth histories and relationships to local environmental conditions

Oxygen and carbon stable isotope profiles were constructed for two species of large subtropical gastropods of the family Fasciolariidae—Triplofusus giganteus and Fasciolaria tulipa—from the Florida Keys and the Bahamas, to evaluate their life history and to assess their potential as paleoenvironmental proxies. Oxygen isotope profiles revealed T. giganteus and F. tulipa grew their shells for 6 and 3 years, respectively. Both mollusks show faster growth rates during the first half of their lifespan. Mean annual temperatures (MAT) derived from oxygen isotopes for T. giganteus were 26.5 °C and for F. tulipa were 26.7 °C, both matching instrumental MATs of 26.7 and 26.5 °C for the Florida Keys. Both shells, however, failed to record entire mean annual temperature ranges (MART). Fasciolaria tulipa yielded a calculated MART of 5.6 °C compared with a measured MART of 9.3 °C, and T. giganteus showed a calculated MART of 6.9 °C compared with a measured MART of 9.4 °C. Carbon isotopes of T. giganteus were ambiguous and reveal no significant relationships with trends in nutrient concentrations (N and P), dissolved oxygen, and dissolved organic carbon, although they did exhibit more negative values concomitant with landfall of Hurricane Irene and trended to increasing values with ontogeny that could reflect migration. Carbon isotopes in F. tulipa were lower during winters, possibly reflecting seasonal upwelling or seagrass-mediated carbon cycling.     

Osmoregulation, nutritional effects and buoyancy of marine larval fish: a bioassay for assessing density changes during the earliest life-history stages

It has been hypothesized that marine fish larvae in the advanced stages of starvation would show increased density (ρ = mass volume⁻¹) from water loss due to osmoregulation failure. Changes in larval buoyancy are currently attributed to swim bladder regulation and protein synthesis or catabolism. Osmoregulation-related changes in density is an alternative mechanism, the importance of which remains untested in the laboratory and the influence of which on vertical distributions is unknown. We provide evidence that loss of osmotic control is a plausible mechanism for increased density of larval cod (Gadus morhua L.). Furthermore, our results show that this mechanism is not restricted to larvae in the advanced stages of starvation. “Relative” larval densities are estimated using a modified density gradient. We use a gravimetric method to separate the effects of nutrition from osmoregulation failure. We assessed the importance of sampling strata on estimates of larval density. Proportional sampling within three depth strata (stratified sample) produced the least biased method for determining the “average” density of a population of larvae in laboratory culture. Larvae sampled from the bottom third of the culture tank were significantly more dense then those sampled from the surface. This was true for larvae of all ages. The average change in density from hatching till death from starvation for larvae sampled in the surface stratum was nominal (Δρ = 5.0 × 10⁻⁴ g cm⁻³), while the change for those sampled from the bottom stratum was large (Δρ = 3.8 × 10⁻³ g cm⁻³). These large density differences suggest that larvae sampled from the bottom stratum were either osmotically stressed or were facultatively changing their density via regulatory pathways. Preliminary observations suggest that vitality is lower amongst those larvae which are sampled near the bottom. The small change in average density of larvae sampled from the surface stratum was due to starvation. The density differences we observed between “osmotically stressed” and “starving” larvae could readily have been misconstrued as differences in feeding and growth experienced by individual larvae. The potential bias of increased density from osmoregulation failure must be considered as a factor in experimental designs developed to assess the effect of fed and starved treatments on buoyancy for larvae of all ages. The simple bioassay we describe may prove useful both as a means of assessing larval condition and as a mechanism for evaluating factors affecting larval vertical distributions in the field. Received: 13 January 1997 / Accepted: 3 February 1997     

Zooplankton growth rates: the influence of size in nauplii of tropical marine copepods

Growth rates and development times were determined for nauplii of the genera: Acartia, Centropages, Corycaeus, Oithona, Paracalanus, Parvocalanus and Temora in nearshore waters of Jamaica from in situ microcosm incubations. At these high local temperatures (∼28 °C), total naupliar development time was short: 3 to 4 d inshore and 4 to 5 d offshore. Mean instantaneous growth rates (g) ranged from as high as 0.90 d⁻¹ for Parvocalanuscrassirostris to as low as 0.41 d⁻¹ for Corycaeus spp. In general, nauplii of cyclopoid copepods appeared to grow more slowly than those of calanoids of the same size. Naupliar growth rates were significantly related to body size (r ²= 0.43 to 0.50), but were unrelated to chlorophyll concentration in any measured size-fraction. This suggests that nauplii are generally not limited by resources, but are growing at their maximum temperature and size-dependent rates. Received: 30 May 1997 / Accepted: 13 May 1998     

Interactions between nutritional status and long-term responses to ultraviolet-B radiation stress in a marine diatom

Influences of nutritional status on the photoinhibitory effects of ultraviolet-B radiation (UVBR: 290 to 320 nm) on the specific growth rates (_obs) and biomass of Phaeodactylum tricornutum were determined using nutrient-replete batch cultures and nutrient-limited continuous cultures. P. tricornutum cultures were exposed to UVBR doses representative of current mid-latitude and ozone-depletion intensities. Specific growth rates and biomass were inhibited from 2 to 16% by UVBR during nutrient-replete growth. However, no effect of UVBR on _obs or biomass was detectable when nutrient limitation exceeded the potential for limitation by UVBR. Thus, a competitive interaction appears to occur between macronutrient stress and UVBR stress, such that _obs and biomass will be determined by the most limiting factor. Our results suggest that measurable decreases in phytoplankton _obs and biomass from UVBR are most likely in nutrient-rich areas of the ocean, while these parameters may not be appropriate for measuring UVBR stress in regions of nutrient limitation.     

Zooplankton growth rates: the influence of size and resources in tropical marine copepodites

Growth rates were determined for copepodites of the genera: Acartia, Centropages, Corycaeus, Oithona, Paracalanus, Parvocalanus and Temora in nearshore waters of Jamaica from in situ microcosm incubations. At these high local temperatures (∼28 °C), total copepodite development time was as short as 4 to 5 d. Mean instantaneous growth rates (g) ranged from as high as 1.2 d⁻¹ to as low as 0.1 d⁻¹. In general, cyclopoid copepods appeared to grow more slowly than calanoids of the same size. Enhancement of resources by nutrient addition caused a 32% increase in growth rates in experiments from a mesotrophic site, but only a 17% increase at a more eutrophic site. Additionally, copepodites at both sites showed faster development and generally larger size at stage in response to nutrient addition. Growth rates were positively related to chlorophyll concentration in the >2 μm size-fraction. A significant relationship of growth rate to body size (r ² = 0.45) emerged across a wide range of trophic status, but it was confounded with resource availability. It appears that growth in tropical copepod copepodites may be frequently limited by resources in a size-dependent manner. Received: 30 May 1997 / Accepted: 13 May 1998     

Stable carbon isotope ratios in Red Sea barnacles (Cirripedia) as an indicator of their food source

The stable isotope ratio ¹²C/¹³C was used to investigate the source of carbon in free-living barnacles and in coral-inhabiting barnacles from the Red Sea. The δ¹³C of most of the barnacles collected on the open shore ranges between −17.5 and −19.7‰, indicating relative enrichment of light carbon originating from the open-sea phytoplankton. Those collected in closed habitats showed heavier isotopic composition. The δ¹³C of the coral-inhabiting barnacles ranges from −14.1 to −16.7‰, suggesting that the carbon contribution of open-sea plankton to these barnacles is less important than it is to free-living barnacles. We hypothesize that coral organic matter and zooxanthellae expelled by the host coral contribute carbon to the barnacle, and that a mixture of this relatively heavy carbon with carbon from other sources is responsible for the high values of δ¹³C in coral barnacles. Received: 28 February 1997 / Accepted: 16 September 1997     

GPS tracking a marine predator: the effects of precision,resolution and sampling rate on foraging tracks of African Penguins

We used a prototype GPS logger to track the movements of breeding African Penguins (Spheniscus demersus). The loggers also recorded temperature and water depth, which allowed us to reconstruct foraging tracks in three dimensions, although GPS signals are interrupted when the birds dive. Here we report the loggers performance in the field and assess the effects of GPS error, resolution and sampling rate on estimates of foraging track length and speed. There is a trade-off between sampling rate and battery lifespan. We tested loggers at sampling intervals of 1 s, 10 s, 1 min, 2 min and 10 min. Sampling less frequently increases the chance of tracking an entire foraging trip, but it slows uplink times, slightly decreases the accuracy of positional fixes, and significantly reduces the ability to measure fine-scale aspects of foraging behaviour. Compared with radio or satellite tracking, GPS loggers offer unprecedented detail about animal movements. The results of our analysis suggest that techniques that sample relatively infrequently, such as satellite tracking, underestimate actual track lengths by up to 50%. However, caution is needed when interpreting fine-scale sampling for relatively slow-moving organisms. Re-sampling 1-s tracks suggests that c. 35% of apparent movements at this scale are due to measurement error and, more importantly, the limited spatial resolution of GPS (1.85×1.54 m at the study area). We recommend that researchers use a 1-s sampling rate for fine-scale studies, but resample at less frequent intervals to remove spurious noise for slow-moving animals. At current levels of resolution, animals should move at least 4 m per sampling interval. We provide empirical correction factors to compare inferred track length sampled at different rates, but caution that these are idiosyncratic and strongly dependent on the animals behaviour. Overall, GPS loggers offer a significant advance for studies of fine-scale animal movement patterns.Communicated by O. Kinne, Oldendorf/Luhe     

Stable isotope and biochemical fractionation in the marine pelagic food chain: the jellyfish Pelagia noctiluca and net zooplankton

In our field study we analyzed the C and H isotopic and biochemical (C, N, P, protein, lipid, carbohydrate) composition of the jellyfish Pelagia noctiluca (collected from the Gulf of Trieste in 1985 to 1986) and its presumed diet-net zooplankton. The mean ¹³C (-18.8) and D (-58.4) ratios of P. noctiluca showed enrichment in heavy isotopes relative to net zooplankton (2 for carbon and 30 for hydrogen). Both the jellyfish and net zooplankton were characterized by a linear correlation between ¹³C and D. C. N, P, protein, lipid, and carbohydrate contents of P. noctiluca were low on a dry weight basis as compared to net zooplankton. Significantly lower C:N and C:P ratios were found in jellyfish indicating a greater loss of carbon relative to nitrogen and phosphorus along the passage to a higher trophic level. Isotopic and biochemical evidence indicate that, though collected in nearshore waters, P. noctiluca depended on autochthonous marine organic matter.     

Stable isotope study of sedimentary carbon utilization by Capitella spp.: effects of two carbon sources and geochemical conditions during their diagenesis

Two experiments were conducted to determine the effect of the type of sediment organic matter and geochemical conditions during diagenesis on the stable carbon isotope ratio ¹³C of the deposit-feeding polychaete worms Capitella spp. Laboratory experiments showed ¹³C values of-13.5 to-13.9%. for worms grown on fresh and aged kelp, Macrocystis pyrifera. Field experiments on worms in the Santa Barbara Channel, California (USA), in 1986 revealed more negative ¹³C values (to-19%.) when oil and kelp were added to sediments. The more negative values suggest the incorporation of oil carbon into worm tissues, supporting previous indications of the importance of hydrocarbons in food webs near oil seeps.     

Ecology of loggerhead marine turtles Caretta caretta in a neritic foraging habitat: movements, sex ratios and growth rates

Much is still to be learned about the spatial ecology of foraging marine turtles, especially for juveniles and adult males which have received comparatively little attention. Additionally, there is a paucity of ecological information on growth rates, size and age at maturity, and sex ratios at different life stages; data vital for successful population modelling. Here, we present results of a long-term (2002–2011) study on the movements, residency, growth and sex ratio of loggerhead turtles (Caretta caretta) in Amvrakikos Gulf (39°0′N 21°0′E), Greece, using satellite telemetry (N = 8) and ongoing capture–mark–recapture (CMR; N = 300 individuals). Individuals encountered at sea ranged from large juvenile to adult (46.2–91.5 cm straight carapace length) and demonstrated growth rates within published norms (<2.7 cm yr^?1) that slowed with increasing body size. We revealed that an unexpectedly high proportion of animals were male (>44 % of captures above 65 cm straight carapace length), compared to region-wide female-biased hatchling production, indicating sex-biased survival or possible behavioural drivers for likelihood of capture in the region. Satellite tracking confirmed that some turtles establish discrete, protracted periods of residency spanning more than 1 year, whilst others migrated away from the site. These findings are underlined by CMR results with individual capture histories spanning up to 7 years, and only 18 % of individuals being recaptured.     

Nephtys hombergii, a free-living predator in marine sediments: energy production under environmental stress

Nephtys hombergii is a free-living, burrowing predator in marine sediments. The worm is, therefore, exposed to various environmental conditions which tube-dwelling polychaetes of the same habitat most likely do not encounter. The worms have to survive periods of severe hypoxia and sulphide exposure, while at the same time, they have to maintain agility in order to feed on other invertebrates. N. hombergii is adapted to these conditions by utilising several strategies. The species has a remarkably high content of phosphagen (phosphoglycocyamine), which is the primary energy source during periods of environmental stress. With increasing hypoxia, energy is also provided via anaerobic glycolysis (pO₂<7 kPa), with strombine as the main end-product. Energy production via the succinate pathway becomes important only under severe hypoxia (<2 kPa), suggesting a biphasic response to low oxygen conditions which probably is related to the worm's mode of life. The presence of sulphide resulted in a higher anaerobic energy flux and a more pronounced energy production via glycolysis than in anoxia alone. Nevertheless, after sulphide exposure under anaerobic conditions of <24 h, N. hombergii is able to recover completely. Although N. hombergii appears to be well adapted to a habitat with short-term fluctuations in oxygen and appearance of hydrogen sulphide, its high energy demand as a predator renders it likely to limit its survival in an environment with longer lasting anoxia and concomitant sulphide exposure. Received: 28 May 1997 / Accepted: 21 June 1997     

Uptake of cadmium in tropical marine lamellibranchs,and effects on physiological behaviour

The effect of cadmium introduced as inorganic salts (chloride, nitrate, sulfate, carbonate, acetate and iodide) or organic chelates (with EDTA, NTA and humic acid) on the physiological behaviour of six species of tropical lamellibranchs — arcid blood clamsAnadara granosa andA. rhombea, venerid clamsMeretrix casta andKatelysia opima, and mytilid musselsPerna viridis andP. indica, collected from various intertidal areas in southern India from 1986 to 1989 — was evaluated. Inorganic salts of Cd are reported to completely ionize upon introduction to full-strength sea water and form predominantly (97%) chloride complexes. Some inorganic Cd salts, therefore, would hardly affect physiological parameters. However, bioaccumulation of Cd was greatest upon exposure to Cd introduced as sulfate, followed by iodide, acetate, chloride, nitrate and carbonate; 96-h LC₅₀'s also varied, but in a different order, being highest for chloride (3.5µg ml^–1), followed by nitrate, acetate, iodide and sulfate (1.8µg ml^–1). Furthermore, Cd levels increased linearly with time of exposure and levels in medium, and were species-specific, being highest in the arcid clams, followed by the venerid clams and mytilid mussels. The presence of other metals such as Zn reduced bioaccumulation of Cd significantly (P < 0.001), but the reverse was not true. Upon exposure to Cd in the presence of Cu, on the other hand, accumulation of both metals was reduced by 15 to 20%. In the presence of organic chelators — EDTA, NTA and reduced glutathione (GSH) — Cd accumulation was significantly inhibited, by 1/2 to 1/3. Humic acid, on the other hand, had no effect. Cd levels in soft tissues were inversely related to ambient salinity in the range of 12 to 32. However, both Zn and Cu significantly inhibited Cd accumulation irrespective of salinity. Cd increased the rate of filtration inAnadara granosa over the first 24 h by 28%, but on prolonged exposure (96 h) that rate decreased significantly, by 50%. Zn alone and in the presence of Cd, on the contrary, initially reduced filtration activity, which subsequently increased to control levels upon continued exposure. Cu alone and in combination with Cd and Zn also significantly (P < 0.001) inhibited filtration relative to controls. This combination also progressively reduced respiratory activity with time, the maximum decrease being 80% after 96 h of exposure. Furthermore, impact of all three metals on both filtration rate and oxygen consumption was initially synergistic, but changed to antagonistic upon prolonged exposure up to 96 h.Deceased. Please send all correspondence and requests for reprints to Dr. S. Patel at the same address     

Large nonlethal effects of an invasive invertebrate predator on zooplankton population growth rate

We conducted a study to determine the contribution of lethal and nonlethal effects to a predator's net effect on a prey's population growth rate in a natural setting. We focused on the effects of an invasive invertebrate predator, Bythotrephes longimanus, on zooplankton prey populations in Lakes Michigan and Erie. Field data taken at multiple dates and locations in both systems indicated that the prey species Daphnia mendotae, Daphnia retrocurva, and Bosmina longirostris inhabited deeper portions of the water column as Bythotrephes biomass increased, possibly as an avoidance response to predation. This induced migration reduces predation risk but also can reduce birth rate due to exposure to cooler temperatures. We estimated the nonlethal (i.e., resulting from reduced birth rate) and lethal (i.e., consumptive) effects of Bythotrephes on D. mendotae and Bosmina longirostris. These estimates used diel field survey data of the vertical gradient of zooplankton prey density, Bythotrephes density, light intensity, and temperature with growth and predation rate models derived from laboratory studies. Results indicate that nonlethal effects played a substantial role in the net effect of Bythotrephes on several prey population growth rates in the field, with nonlethal effects on the same order of magnitude as or greater (up to 10-fold) than lethal effects. Our results further indicate that invasive species can have strong nonlethal, behaviorally based effects, despite short evolutionary coexistence with prey species.     

Carbon isotope effects during precipitation of barium carbonate: implications for environmental studies

Here we demonstrate that precipitation of dissolved inorganic carbon (DIC) to at pH 10.26, 11.38 and 12.11 does not cause measurable carbon isotope fractionation. However, NaOH solution prepared under standard laboratory procedure and used to increase pH value for precipitation of BaCO₃ can be contaminated by atmospheric CO₂. In our experiment, δ¹³C value in the contaminant DIC was −24.30‰, which resulted in decrease in the δ¹³C(DIC) value by 0.03, 0.07 and 0.22 at pH 10.26, 11.38 and 12.11, respectively.     

Testing isosource: stable isotope analysis of a tropical fishery with diverse organic matter sources

We sampled consumers and organic matter sources (mangrove litter, freshwater swamp-forest litter, seagrasses, seagrass epiphytes, and marine particulate organic matter [MPOM]) from four estuaries on Kosrae, Federated States of Micronesia for stable isotope (sigma13C and sigma34S) analysis. Unique mixing solutions cannot be calculated in a dual-isotope, five-endmember scenario, so we tested IsoSource, a recently developed statistical procedure that calculates ranges in source contributions (i.e., minimum and maximum possible). Relatively high minimum contributions indicate significant sources, while low maxima indicate otherwise. Litter from the two forest types was isotopically distinguishable but had low average minimum contributions (0-8% for mangrove litter and 0% for swamp-forest litter among estuaries). Minimum contribution of MPOM was also low, averaging 0-13% among estuaries. Instead, local marine sources dominated contributions to consumers. Minimum contributions of seagrasses averaged 8-47% among estuaries (range 0-88% among species). Minimum contributions of seagrass epiphytes averaged 5-27% among estuaries (range 0-69% among species). IsoSource enabled inclusion of five organic matter sources in our dual-isotope analysis, ranking trophic importance as follows: seagrasses > seagrass epiphytes > MPOM > mangrove forest > freshwater swamp-forest. IsoSource is thus a useful step toward understanding which of multiple organic matter sources support food webs; more detailed work is necessary to identify unique solutions.     

Zooplankton growth rates: the influence of female size and resources on egg production of tropical marine copepods

Egg production was measured in 17 species of copepods from the genera Acartia, Calanopia, Centropages, Clausocalanus, Corycaeus, Eucheata, Euterpina, Oithona, Oncaea, Paracalanus, Parvocalanus, Temora and Undinula in Jamaican waters. At the high local temperatures (∼28 °C), mean egg production ranged from 3.2 to 88 eggs female^–1 d^–1, and instantaneous female growth (g, as egg production) ranged from 0.04 to 0.87 d^–1. Female growth was positively related to ambient chlorophyll concentration (r ² = 0.44) and negatively to female body size (r ² = 0.29). Together these two variables explained 60% of the variation in growth. When quadratic terms for chlorophyll and a term for interaction of body size and chlorophyll were introduced, 82% of the variance in growth rate was explained. Egg production rates represent an extension of the resource and size-dependent relationship established for copepodites. In smaller species (<3.5 μg), egg production was comparable to prior copepodite somatic growth; in larger species (>3.5 μg), egg production is compromised at lower resource concentrations than copepodite somatic growth. Thus, it appears that egg production in tropical copepods may be frequently limited by resources in a size-dependent manner. Under conditions where growth is resource limited, we caution against the application of egg production rates for the calculation of total copepod production. Received: 30 May 1997 / Accepted: 13 May 1998     

Stable isotope ratio variations in non-scleractinian coelenterate carbonates as a function of temperature

Some coelenterates of the class Hydrozoa and some anthozoan coelenterates from the subclass Octocorallia secrete skeletons of calcium carbonate. Skeletal carbonates of three hydrozoans and of two octocorals were analyzed for the stable isotopes of carbon and oxygen. The results suggest that each of these coelenterates deposits CaCO₃ in oxygen isotopic equilibrium with seawater, and that at least one octocoral, Heliopora, has skeletal carbon in apparent isotopic equilibrium with atmospheric CO₂. Two of these coelenterates, Millepora and Helipora, are significant contributors to the construction of coral reefs. Whereas ¹⁸O of these corals is temperature dependent, ¹³C is not obviously related to temperature. The ¹⁸O-temperature relationship is not significantly different from the oxygen isotope paleotemperature scale developed by Epstein et al. (1953). These findings contrast with numerous analyses of the carbonate in scleractinian coelenterates, which have long been reported to deposit CaCO₃ skeletons whose carbon and oxygen isotopic compositions are not in equilibrium with the external sea-water environment.