Oxygen and carbon stable isotopic profiles of the fan mussel, Pinna nobilis, and reconstruction of sea surface temperatures in the Mediterranean

Stable oxygen and carbon isotope profiles ('¹⁸O_skeletal and '¹³C_skeletal), taken along the direction of growth from the umbo to the shell margin in shells of the pinnid Pinna nobilis, were used to reconstruct sea surface temperatures (SST) in the south-east Mediterranean and ontogenetic records of metabolic CO₂ incorporation. Comparison of the seasonal cycle of SST, predicted from the '¹⁸O_skeletal record of a small (young) rapidly growing pinnid and temperature measured with a continuous in situ recorder showed that P. nobilis calcifies under isotopic equilibrium with surrounding seawater, thus indicating that P. nobilis shells can be used as a reliable predictor of SST. A 10-year SST record for the south-east Mediterranean was reconstructed from the shell profiles of four pinnid shells of different sizes and ages collected in 1995 and 1996. Reliable resolution of the seasonal SST could only be achieved during the first 4 years of shell growth. As the pinnids grew older, the temperature record was poorly resolved because the shell growth had diminished with age, resulting in time-averaging of the record. The amplitude of the generated seasonal temperature cycle compared favourably (DŽ°C) with a long-term temperature record from northern Mediterranean waters. Clear seasonal cycles in '¹³C_skeletal were observed with an amplitude of ~1.0‰, similar to the calculated seasonal changes in '¹³C of seawater (0.6‰) overlying seagrass meadows. An ontogenetic trend towards less positive '¹³C_skeletal values was too large to be attributed to any decrease in '¹³C in seawater resulting from the invasion of anthropogenic CO₂. It is suggested that the temporal changes of '¹³C_skeletal are due to incorporation of respiratory CO₂ into the extrapallial fluid and reflect changes in the metabolic activity of the pinnid rather than changes in the isotopic composition of dissolved inorganic carbon within the surrounding seawater.     

Oxygen and carbon isotope profiles from Nassa mutabilis shells (Gastropoda): accretion rates and biological behaviour

Six shells of the Gastropoda species Nassa mutabilis L. were collected in the Adriatic Sea at a depth of a few metres off the central Italian coast during 1996 and 1997. The oxygen and carbon isotope composition of spot samples collected from the shell apex to the top was measured according to well-established techniques. The results obtained allow the following conclusions: (1) the CaCO₃ of the N. mutabilis shells is essentially formed by aragonite as proved by X-ray diffraction measurements; (2) the CaCO₃ is precipitated under isotope equilibrium conditions or very close to equilibrium with the environmental water; (3) shell accretion takes place all year round, although the warm period is preferred since the accretion rates are higher during the warm half of the year and considerably reduced in winter; (4) the carbon isotope composition changes according to the age of the specimen, probably in relation to changes in its nutritional regimen; and (5) the alternation of warm and cold periods recorded by the shells allows the evaluation of their age and of the season of their birth and death. Received: 3 December 1998 / Accepted: 7 September 1999     

Environmental controls on shell growth rates and δ<Superscript>18</Superscript>O of the shallow-marine bivalve mollusk <Emphasis Type="Italic">Phacosoma</Emphasis><Emphasis Type="Italic">japonicum</Emphasis> in Japan

Microgrowth patterns and the oxygen isotope composition of juvenile, shallow-marine bivalve mollusk shells of Phacosoma japonicum (Reeve) in Japan were analyzed and cross-calibrated with environmental parameters. Mark-and-recovery experiments indicate that a pair of two microgrowth lines and two microgrowth increments is produced every lunar day. This finding makes it possible to assign exact calendar dates to each portion of the shell. Average daily growth rates decrease by 61% from age two to three and 55% from age three to four. The length of the growing season and the growth rate are mainly controlled by temperature: shell growth ceases below 14.2°C (age two) and 16.8°C (age four) and is most rapid between 24.6°C and 27.2°C. Based on local temperature cycles, the growing season is longest in Seto Inland Sea, central Honshu (from May to November) and shortest at Hakodate Bay, North Japan (from June to October). The annual oxygen isotope profiles of the shells reflect the temperature cycle and the varying amounts of freshwater added to the seawater by precipitation. The most negative '¹⁸O values of -3.15‰ occur during the rainy season, i.e. during the monsoon and typhoon seasons. Growth rates are only slightly affected by salinity changes. Strongly reduced growth rates during the second half of the year at Seto Inland Sea and to a lesser extent at Tokyo Bay are explained by nutrient deprivation. Our study provides the basis for the use of P. japonicum in high-resolution ecological studies and environmental reconstructions.     

Ostracod species-specific utilisation of sediment detritus and newly settled cyanobacteria, Aphanizomenon sp., in the Baltic Sea: evidence from stable carbon isotopes

The three meiobenthic ostracod species Candona neglecta, Paracyprideis fennica and Heterocyprideis sorbyana are commonly encountered in the deep (20-40 m) soft bottoms of the Baltic proper, and may contribute more to the total meiobenthic biomass than any other group. Experimental data indicate substantial differences in their utilisation of settling phytodetritus, with C. neglecta able to exploit newly settled organic material to a larger extent than P. fennica and H. sorbyana. Ostracod species composition, as well as ostracod and sediment carbon isotope contents, were studied in the field from an area with local differences in potential food resources. The study was performed during the late period of summer blooms of the cyanobacteria Aphanizomenon sp. Results showed that when all samples were taken together, adult C. neglecta (-22.4‰) was significantly more depleted in '¹³C than adult P. fennica (-21.0‰) and H. sorbyana (-20.3‰), indicating differences in food selection among the species. The flocculent sediment layer had, in all instances, lighter carbon ratios than did the lower layers. This trend was mirrored in most cases in the slight enrichment in C. neglecta compared to generally greater enrichment in the other two species. Carbon signatures of C. neglecta also varied significantly between stations, indicating that this species fed on different resources depending on location. Juvenile C. neglecta were far more depleted in '¹³C than adults and reflected the carbon signature of the cyanobacteria Aphanizomenon sp. The latter is known to be the most '¹³C-depleted phytoplankton member in the area.     

Effects of exposure to toxic and non-toxic dinoflagellates on oxygen consumption and locomotion in stage 1 larvae of the crabs Cancer oregonensis and C. magister

Sublethal effects on larval crabs upon exposure to toxic dinoflagellates were examined in the laboratory in early 1999. Specifically, oxygen consumption rates and geotaxis responses were determined for stage 1 larvae of the crabs Cancer oregonensis (Dana) and C. magister Dana that were exposed to non-toxic (Alexandrium tamarense, strain 115) or toxic (A. fundyense, strain 1719) dinoflagellates or to freshly hatched nauplii of the brine shrimp Artemia sp. In C. oregonensis, larvae exposed to the toxic dinoflagellate showed reduced rates of oxygen consumption compared to those exposed to non-toxic dinoflagellates or brine shrimp nauplii. Larvae exposed to a filtrate of the non-toxic dinoflagellate showed no change in oxygen consumption, but a reduced rate when exposed to filtrate from the toxic alga at densities >5᎒² cells ml^-1. In C. magister, larvae exposed to the non-toxic A. tamarense or the toxic A. fundyense had reduced oxygen consumption rates. Larvae exposed to filtrates of non-toxic and toxic dinoflagellates had no change in oxygen consumption. In geotaxis tests, C. oregonensis larvae exposed for 1 day to the toxic A. fundyense reduced their level of locomotion compared to those exposed to non-toxic A. tamarense or to brine shrimp nauplii. C. magister larvae showed no change in activity after a 1-day exposure to the toxic A. fundyense. After a 4-day exposure to A. fundyense, C. magister larvae had much reduced locomotion. Reduced locomotory activity in larvae exposed to toxic algae is consistent with the changes in oxygen consumption rates. Responding to exposure to toxic algae by reducing locomotion may affect vertical migration in these negatively buoyant crab larvae, resulting in sinking below a toxic alga bloom, at least temporarily.     

Cardiac output during exposure to <Emphasis Type="Italic">Chattonella</Emphasis><Emphasis Type="Italic">marina</Emphasis> and environmental hypoxia in yellowtail (<Emphasis Type="Italic">Seriola</Emphasis><Emphasis Type="Italic">quinqueradiata</Emphasis>)

The experiments were undertaken to measure, for the first time, cardiac output in yellowtail (Seriola quinqueradiata) during exposure to a harmful red tide flagellate (Chattonella marina). The responses were compared with those during exposure to environmental hypoxia to evaluate the significance of the drop of arterial oxygen partial pressure (PaO₂) in the fish-kill mechanisms by C. marina. PaO₂ immediately decreased, whereas heart rate (HR) was maintained until shortly before death during exposure to C. marina. Suffocation developing during the exposure resulted from a decrease in blood oxygen content, but not from lowered blood flow to the tissue. Although exposure to both C. marina and hypoxia immediately decreased PaO₂, arterial oxygen content (CaO₂) and pH (pHa) were significantly lower, whereas HR and cardiac output (&Qdot;) remained significantly higher, for the C. marina-exposed fish than for hypoxia-exposed fish. Although the drop in PaO₂ appears to play a pivotal role in the mechanisms of fish death by C. marina, other physiological response(s) should also be considered.     

Measurement of calcium carbonate deposition in molluscs by controlled etching of radioactively labeled shells

Rates of calcium carbonate removal from shell pieces of Argopecten irradians (Say) and Mercenaria mercenaria (L.) were dependent on the type of etching fluid used and not shell origin. Etching was uniform over the entire shell surface, but surface morphology differed with etching fluids. Peak radioactivity was in early eluant fractions of shells etched immediately after radioactive labeling, and in later fractions when individuals were placed in isotope-free seawater after labeling. The etching technique can measure growth during the labeling period and subsequently by estimating the amount of calcium in fractions prior to the radioactive peak. Geometry of shell layers influenced the pattern of radioactivity seen in fractions. Peak location varied inside and outside the pallial line of individual M. mercenaria. A significant portion of the inorganic carbon used in shell formation by M. mercenaria was derived from metabolic CO₂.     

Feeding responses of the bivalves Crassostrea gigas and Mytilus trossulus to chemical composition of fresh and aged kelp detritus

The chemical composition of kelps (e.g. polyphenolics) deters grazing by herbivores, but kelp detritus is potentially a source of nutrition for suspension feeders. The effects of kelp detritus derived from two species [Agarum fimbriatum Harvey and Costaria costata (Turner) Saunders] on feeding of oysters, Crassostrea gigas Thunberg, and mussels, Mytilus trossulus Gould, were examined in feeding experiments. Fresh and aged kelp particles were sequentially presented in combination with the microalga Rhodomonas lens at an initial total concentration of 5᎒^-4 ml^-1. Aging of kelp particles for 4 days in seawater significantly reduced the concentration of polyphenolics without changing the total carbon or nitrogen content. Clearance rates of both mussels and oysters were significantly lower in the presence of fresh versus aged kelp particles, and clearance rates declined overall with declining polyphenolic concentrations. Video endoscopy was used to examine feeding selectivity at the level of the gill in oysters in the same food treatments used in the clearance rate experiments. Comparison of particle composition in the water versus the pseudofeces in both oysters and mussels was also used as a measure of feeding selectivity. When presented with R. lens in combination with fresh and aged kelp particles selectivity for R. lens tended to be greater against fresh than aged particles, and there was some indication that this was stronger for A. fimbriatum than for C. costata particles. The ability to select was lower at very high polyphenolic concentrations, which may reflect poisoning of sensory binding sites. These data suggest that bivalves distinguish among particles of varying chemical composition and respond by changing their clearance rates and their selectivity.     

Energy budgets and feeding rates of Coryphaenoides acrolepis and C. armatus

This study develops energy budgets and estimates feeding rates for two macrourid fishes, Coryphaenoides acrolepis, dominant in the bathyal eastern North Pacific, and the abyssal cosmopolitan species, Coryphaenoides armatus. Daily energy expenditure by C. acrolepis was nearly twice that of C. armatus. C. acrolepis allocated nearly equal amounts of energy to metabolism and growth. Once sexual maturity was reached reproduction became the dominant energetic cost. Either these costs are necessary to retain adequate numbers of eggs and larvae on the continental slopes, or this fish does not reproduce on an annual basis and the calculated costs are an overestimate. C. armatus allocated relatively more energy to metabolism than growth. It may be semelparous, and this strategy would be of great energetic savings in its food-poor but stable environment. Individual daily ration for C. acrolepis decreased from 0.31% to 0.07% of body weight (BW) and for C. armatus from 0.12% to 0.02% BW with increasing fish length. These rates are substantially lower than those for fishes living in cold waters on the continental shelves. The population feeding rates for C. acrolepis ranged from 0.8 to 15 kg km^-2 day^-1 and for C. armatus from 5 to 2,800 g km^-2 day^-1. The scavenging behaviour of C. acrolepis was used to investigate the role of carrion as a food supply to the deep-sea benthos. It was estimated that the carrion eaten by C. acrolepis is equivalent to 0.04 mg C m^-2 day^-1 or only 0.2-0.4% of the average small particulate flux. Carrion consumption is important for scavengers like C. acrolepis, but it is not an important component of the carbon flux into the deep-sea benthic environment.     

Studies on pigments,proteins and photosynthetic rates in some mangroves and mangrove associates from Bhitarkanika,Orissa

Pigment contents, proteins and net photosynthesis were investigated in fully developed leaf of 1-year-old seedlings of six mangroves (Bruguiera gymnorrhiza, Rhizophora apiculata) and mangrove associates (Caesalpinia bonduc, Cerbera manghas, Derris heterophylla, Thespesia populnea), collected from Bhitarkanika, located on the east coast of India. Large variations in the photosynthetic rates (P_N) among the six species were observed, ranging from 10.16 µmol CO₂ m^-2 s^-1 in C. bonduc to 15.28 µmol CO₂ m^-2 s^-1 in R. apiculata. The total leaf protein content ranged from 12.09 mg g^-1 dry wt in T. populnea to 51.89 mg g^-1 dry wt in B. gymnorrhiza. The chlorophyll a/b ratio was typically about 3.0 in all the studied species, except C. bonduc (2.8). Photosynthetic rates and chl a/b ratio in the leaves were found to be correlated. Analysis of chlorophyll and xanthophyll spectra suggested: (1) variations in different forms and amounts of carotenes as well as xanthophylls and (2) the presence of high amounts of near-UV-absorbing substances in leaves, particularly in the two mangroves (B. gymnorrhiza, R. apiculata) and a mangrove associate (T. populnea), which appears to be an adaptive feature. Estimation of the chl a/b ratios in isolated thylakoids yielded a low value of 1.8 for R. apiculata and >2.6 for other species. The total protein/chlorophyll ratios in thylakoids varied considerably from 3.14 (D. heterophylla) to 10.88 (T. populnea) among the mangrove associates and from 16.09 to 18.88 between the members of the Rhizophoraceae. The chlorophyll/carotenoid ratios in thylakoids of the six species were more or less similar. The absorption spectra for washed thylakoids of C. manghas and D. heterophylla exhibited absorption characteristics typical for C₃-plant thylakoids. However, thylakoids isolated from R. apiculata, B. gymnorrhiza, C. bonduc and T. populnea exhibited an unusual increase in absorption in the blue region (380-410 nm) of the absorption spectrum. The presence of high-absorbing (in the short-wavelength, near-UV region) pigments appears to be closely associated with the thylakoids in R. apiculata and T. populnea. Our results, therefore, suggest a wide range of variation, not only in protein and pigment contents of photosynthetic tissues, but also in the spectral characteristics and composition of the pigments in mangrove species. An understanding of the nature of these pigments in mangroves and their associates, under their natural conditions and especially in relation to eco-physiological adaptations, is necessary, not only in relation to conservation, but also to allow propagation under different salinity conditions.     

Extension rate: A primary control on the isotopic composition of West Indian (Jamaican) scleractinian reef coral skeletons

Large variations in skeletal microarchitecture, and in the carbon and oxygen isotopic composition of the skeleton, exist within single calices and over the surfaces of single colonies of various species of West Indian (Jamaican) scleractinian reef corals. Rapidly extending parts of individual colonies are depleted in both C¹³ and O¹⁸ regardless of the abundance of zooxanthellae in the overlying tissues. We suggest that this relationship is due to active translocation of organic compounds to sites of rapid calcification. Considerable variation in isotopic composition is found in skeletons of different specimens of the same species even when the skeletons are sampled in a consistent manner, come from the same locality and depth, and have a comparable growth history. The isotopic composition of the scleractinian skeleton is vastly more complex than has heretofore been realized.     

DNA sequence evidence for speciation,paraphyly and a Mesozoic dispersal of cancellothyridid articulate brachiopods

Because the classification of extant and fossil articulate brachiopods is based largely upon shell characters observable in fossils, it identifies morphotaxa whose biological status can, in practice, best be inferred from estimates of genetic divergence. Allozyme polymorphism and restriction fragment length polymorphism of mitochondrial DNA (mtDNA RFLP) have been used to show that nuclear and mitochondrial genetic divergence between samples of the cancellothyridid brachiopods Terebratulina septentrionalis from Canada and T. retusa from Europe is compatible with biological speciation, but the genetic distances obtained were biased by methodological limitations. Here, we report estimates of divergence in 12S rDNA mitochondrial sequences within and between samples of these brachiopods. The sequence-based genetic distance between these samples (5.98ǂ.07% SE) is at least 10 times greater than within them and, since they also differ in a complex life-history trait, their species status is considered to be securely established. Divergence levels between 12S rDNA genes of three other cancellothyridids, T. unguicula from Alaska, T. crossei from near Japan, and Cancellothyris hedleyi from near Australia are higher than between the two North Atlantic species, and the mean nucleotide distance between all these cancellothyrids is similar to the mean distance between species of Littorina (Mollusca: Gastropoda). Sequences of both 12S and 16S genes from cancellothyridids and other short-looped brachiopod species show neither saturation nor lineage-specific rate differences and, when analysed with different outgroups, either separately or together, yield one unexpected, but well-supported, tree with Alaskan T. unguicula basal and C. hedleyi nested within Terebratulina, i.e. these genera are paraphyletic. A geologically dated divergence between Antarctic and New Zealand species of the short-looped brachiopod Liothyrella is used to calibrate the rate of 12S divergence at ca. 0.1% per million years (MY), and this rate is used to infer that T. septentrionalis and T. retusa have been diverging for ca. 60 MY and that they and T. unguicula have been diverging from their last common ancestor for ca. 100 MY. This indicates a Mesozoic origin for the present-day distribution of cancellothyridids and the basal position of T. unguicula suggests a possible North Pacific centre of origin, with separate Atlantic and Pacific radiations. The inclusion of Cancellothyris within Terebratulina also shows that adult shell characters such as umbo, foramen and symphytium shape, whilst probably indispensible for the practical classification of fossils, are not reliable guides to genealogy.     

Reproductive cycles of three dominant Calanus species in Disko Bay, West Greenland

This study describes the annual reproductive cycles of the three dominant Calanus species, C. finmarchicus, C. glacialis and C. hyperboreus, in Disko Bay (West Greenland) in relation to seasonal phytoplankton development. Relative abundance of females, copepodite stage V (CV) and males, and the developmental stage of the female gonad were examined from plankton samples collected at weekly to monthly intervals from May 1996 to June 1997 with a WP2 net or a pump. During spring and summer, egg production rates were determined. Females of all three species were present year round. Maximum relative abundance was reached by C. hyperboreus females at the beginning of February, by C. glacialis in mid-February, and by C. finmarchicus in April. All three species reproduced successfully in Disko Bay. Their reproductive cycles were considerably different with respect to the timing of final gonad maturation and spawning, and hence in their relation to seasonal phytoplankton development. In all three species, early gonad development took place during winter, before living food became plentiful, suggesting that these processes were largely food independent. Final gonad maturation and spawning in C. finmarchicus was related to the phytoplankton concentration, reflecting that final gonad maturation processes are food dependent in this species. C. glacialis females matured and spawned prior to the spring bloom. Our results indicate that first internal lipid stores and later ice alga grazing supplied final gonad maturation and egg production. Maximum egg production rates of C. glacialis were found in spring and summer, when the chlorophyll a concentration was high. Mature female C. hyperboreus were found from February until mid-April, when the chlorophyll a concentration was still low. In this species, reproductive activity was decoupled from phytoplankton development, and final maturation processes and spawning were solely fuelled by internal energy stores.     

Identification of tenuis of four French Polynesian Carapini (Carapidae: Teleostei)

Four species of adult Carapini (Carapidae) occur on Polynesian coral reefs: Encheliophis gracilis, Carapus boraborensis, C. homei and C. mourlani. Samples collected in Rangiroa and Moorea allowed us to obtain different tenuis (larvae) during their settlement phases or directly inside their hosts. These were separated into four lots on the basis of a combination of pigmentation, meristic, morphological, dental and otolith (sagittae) features. Comparison of these characters with those of the adults allows, for the first time, taxonomic identification of these tenuis-stage larvae.     

Acoustical identification of the concentration layer of a copepod species,Calanus euxinus

Swimming trajectories of Calanus euxinus Hulsemann in the Black Sea were studied using an echosounder at 120 and 200 kHz. C. euxinus were acoustically discriminated with respect to vertical migration and swimming speed, according to dissolved oxygen (DO) concentration and the timing of migrations. Species became torpid in water with DO values <0.5 mg l^у. The time spent swimming under DO conditions between 2 and 5 mg l^у was insignificant, and varied greatly from the 10% to 25% of total time spent swimming under normoxic conditions (5-10 mg l^у). C. euxinus formed a concentration layer in the water of 1-3 m thickness. Upward migration was completed in about 3.5 h, starting 2.5 h before and ending 1 h after sunset (average rate: 0.95 cm s^у) in summer. Species ascended discretely from the suboxic to the lower boundry of the cold intermediate layer (CIL) at 0.82 cm s^у, and passed up the CIL and thermocline fast (2.3 cm s^у). Downward migration took less time (2 h), starting ~1 h before and ending ~1 h after sunrise. Swimming speed within the thermocline and CIL was 2.7 cm s^у; copepods subsequently returned to daylight depth at a sinking speed of 0.57 cm s^у. Total time for C. euxinus to settle to their nocturnal depth layer was about 5 h.     

Mesozooplankton community structure, abundance and biomass in the central Arctic Ocean

During the "International Arctic Ocean Expedition 1991" (20 August-21 September 1991) mesozooplankton was sampled at six stations in the Nansen, Amundsen and Makarov Basins of the central Arctic Ocean from 1,500 m depth to the surface by multiple opening/closing net hauls. Total mesozooplankton abundance decreased from 268 ind. m^-3 in the surface layer (0-50 m) to <25 ind. m^-3 below 200 m depth. The small copepods Oithona similis and Microcalanus pygmaeus, as well as copepod nauplii, were most abundant close to the surface, while Oncaea borealis and Spinocalanus spp. frequently occurred at greater depth. Mesozooplankton dry mass (DM) integrated over the upper 1,500 m of the water column was surprisingly stable throughout the investigation area and measured 2.0ǂ.3 g DM m^-2. Dry mass in the upper 50 m measured 20.9 mg m^-3 and was dominated by Calanus hyperboreus (57.4%) and C. glacialis (21.1%). C. finmarchicus was very abundant only in the Nansen Basin. Below 200 m the calanoid copepods Metridia longa, Microcalanus pygmaeus and Pareuchaeta spp., the decapod Hymenodora glacialis and chaetognaths of the genus Eukrohnia were the principal contributors to biomass values of <1 mg DM m^-3. Hence, vertical changes in abundance, biomass and species composition were much more pronounced than regional differences between the basins. Three different mesozooplankton communities were differentiated according to their faunistic composition and are discussed in context with the major water masses: Polar Surface Water, Atlantic Layer and Arctic Deep Water.     

Seasonal variations in the stable carbon and nitrogen isotope ratios (13C/12C and 15N/14N) of primary producers and consumers in a western Mediterranean coastal lagoon

Stable carbon and nitrogen isotope ratios (¹³C/¹²C and ¹⁵N/¹⁴N) of primary producers and consumers were investigated seasonally throughout 1999, in order to describe the food web in a western Mediterranean coastal lagoon (Lake of Sabaudia, central Italy). Particulate organic matter and algal material (seagrass epiphytes and macroalgae) seem to constitute the main food sources for primary consumers (zooplankton and small benthic invertebrates, respectively) throughout the sampling year, while the seagrass Cymodocea nodosa appears to play a negligible trophic role. As regards the ichthyofauna, carbon stable isotopes differentiated between planktivore and benthivore fish species. However, a benthic-pelagic coupling seems to occur, with some fish of higher trophic levels feeding both on benthic and pelagic materials. Analysis of variance showed that the interaction between the three main factors (species2size2season) significantly affects the isotopic composition of fish, suggesting the presence of intra- and inter-specific resource partitioning. Wide seasonal variations in the isotopic composition were observed in organic matter sources, invertebrates and fish, with a general trend towards depleted values in winter and enriched values in summer. The winter depletion of organic matter sources may be due to several environmental factors and seems to be mirrored in the upper trophic levels. Primary producers and invertebrates are known to have shorter time-integrated isotopic signatures than vertebrates, yet fish also exhibited seasonal isotopic differences. We concluded that the examined fish species can assume a new muscle isotopic signature relatively quickly in response to changes in the isotopic composition of their diet and/or diet shifts.     

In situ mapping of growth lines in the calcitic prismatic layers of mollusc shells using X-ray absorption near-edge structure (XANES) spectroscopy at the sulphur K-edge

The microstructure and composition, including chemical speciation of sulphur (S), of two mollusc shells were investigated using a combination of scanning electron microscopy, X-ray absorption near-edge structure spectroscopy (XANES) and electron probe microanalysis (EPMA). The shell of Pinna is composed of monocrystalline, and Pinctada, of polycrystalline, calcite prisms separated by organic-rich walls. Sulphur speciation information from XANES spectra using a scanning X-ray microscope showed that the protein S content of the interprismatic walls is higher than the SO₄ content, whereas the reverse is true for the intraprismatic structures. High-spatial-resolution XANES maps for the different S species across adjacent calcite prisms confirm their distinctive distributions in the molluscan shells and illustrate the presence of narrow, submicron transverse growth features. On a larger scale, a series of wider growth zones, incorporating the submicron zones, are aligned parallel to each other and cross-cut many calcite prisms. EPMA element maps for magnesium (Mg) and S demonstrate that these growth increments are compositionally zoned, comprising alternating layers of high mineral (Mg-rich) and high organic (S-rich) components. Additionally, these maps confirm that the organic interprismatic walls have lower Mg and higher S than the intraprismatic structures.     

Precipitation of aragonite by calcitic bivalves in Mg-enriched marine waters

To understand the relative importance of biological versus physicochemical control over biomineralization, we have tested if the chemical composition of the medium (i.e., the Mg/Ca ratio) can change the mineralogy of mollusk shells. The shells of mollusks are made of calcite and/or aragonite, which are by far the most common CaCO₃ polymorphs. Several species of bivalves with predominantly calcitic shells have been cultivated in artificial seawater with a Mg/Ca molar ratio within the range of 8.3–9.2, well above the present value for seawater (5.2). Four out of six species used (the scallop Chlamys varia, the oyster Ostrea edulis, the saddle oyster Anomia ephippium and the mussel Mytilus edulis) survived long enough to secrete significant amounts of calcium carbonate. The deposits (sometimes extensive) formed on the interior shell surfaces were predominantly aragonitic. Three individuals of C. varia also increased their length by adding new shell at the margin. Contrary to the internal shell deposits, these margins were high-Mg calcite. This implies that the marginal mantle is able to exert a more strict control on the secreted mineral phase than the mantle facing the internal shell surface. This is the first report on an in vivo experimentally forced switch in bivalve shell mineralogy, from calcite to aragonite due to a change in water chemistry.     

Stable isotopic investigation of physiological and environmental changes recorded in shell carbonate from the giant clam Tridacna maxima

The aragonitic shell of the photosymbiont-bearing bivalve Tridacna maxima contains a record of the physiological and environmental changes the organism has experienced during its lifetime. This record is preserved as chemical and microstructural variations throughout the shell. Stable isotopic analyses of oxygen (¹⁸O/¹⁶O) and carbon (¹³C/¹²C) in shell carbonate were combined with growth increment studies to interpret the shell record of specimens collected from the Rose Atoll (Lat. 14°31S; Long. 168°10W) in April 1982. The seasonal water temperature cycle is recorded in the oxygen isotopic signature of the clams, permitting the recognition of annual cycles in the ¹⁸O profile. The total number of these cycles corresponds to the age of a specimen, while the cycle length is a measure of the yearly growth rate. Large-amplitude cycles, reflecting year-round calcification, characterize the early portion of the growth record. With the onset of sexual maturity and slower growth at an age of approximately ten years, the cycles decrease in amplitude and become more erratic. During this later growth phase calcification is limited to the cooler months of the year, perhaps in response to a re-ordering of energy priorities between growth and gametogenesis. A growth curve developed from the ¹⁸O profile indicates rapid juvenile shell growth followed by slower growth thereafter producing a lifespan of several decades. Carbon isotopic analyses of T. maxima were compared to analyses of the symbiont-barren gastropod Terebra areolata collected from the same locality in April 1984. A 2 depletion in the ¹³C composition of T. maxima shell carbonate is attributed to a symbiontenhanced metabolic rate and an increased flow of isotopically light, respired CO₂ into the carbon pool used in calcification. Such a depletion may prove useful in identifying the presence of photosymbionts in extinct species of fossil mollusks.