Individual relationship between aneuploidy of gill cells and growth rate in the cupped oysters Crassostrea angulata, C. gigas and their reciprocal hybrids

The Portuguese oyster, Crassostrea angulata, is taxonomically close to the Pacific oyster, C. gigas, but there are clear genetic and phenotypic differences between these taxa. Among those differences, the faster growth of C. gigas compared with C. angulata has often been observed in the field. Crosses between C. angulata and C. gigas were performed to investigate the relationship between growth variation and somatic aneuploidy at the individual level in the two taxa and their reciprocal hybrids. The different progenies were reared in Ria Formosa (Portugal) under standard farming conditions. Growth rate and survival were significantly higher in C. gigas than in C. angulata, and the hybrids showed intermediate performances. Significant differences were also observed in the proportion of aneuploid cells (PAC) and of missing chromosomes (PMC) between the two taxa, C. angulata showing the highest values. Intermediate values of PAC and PMC were observed in the hybrids, supporting additive genetic bases of these parameters. Our results also confirm the negative correlation between somatic aneuploidy and growth rate at the individual level, as previously reported in C. gigas.     

Miocene bristlemouths (Teleostei: Stomiiformes: Gonostomatidae) from the Makrilia Formation,Ierapetra, Crete

Bristlemouths of the genus Cyclothone are currently regarded as the most abundant vertebrates on Earth. The fossil record seems to suggest that these fishes diversified during the Miocene in the Pacific Ocean, but there is no evidence of their presence in the Miocene of the Atlantic Ocean and Mediterranean basin. A new bristlemouth, Cyclothone gaudanti sp. nov. (Teleostei, Stomiiformes, Gonostomatidae), is described herein based on 16 specimens from the Upper Miocene Makrilia Formation (late Tortonian of Crete, Greece). The small sized species is characterized by light pigmentation, 30–31 (14–15 + 15–16) vertebrae, dorsal fin with 10–13 rays, anal fin with 10–14 rays, premaxilla bearing seven closely spaced teeth, maxilla with 42–55 teeth, epipleurals, and autogenous parhypural. The presence of epipleurals appears to be unique of this Miocene species, and the re-establishment of this ancestral character state may be possibly interpreted as related to a phylogenetic character reversal. Morphological and paleoecological considerations suggest that this species possibly inhabited the upper mesopelagic layer, at depths ranging from 2–300 and 500 meters.     

Oldest record of the Great White Shark (Lamnidae,Carcharodon; Miocene) in the Southern Atlantic

Lamnid teeth close to but more plesiomorph than Carcharodon carcharias were collected in the late Miocene beds of the Paraná Formation in the central eastern Argentina. We propose, as other authors, that some species formerly assigned to Isurus or to Cosmopolitodus should be included in Carcharodon. Some workers suggested that C. carcharias originated by phyletic evolution in the Pacific basin. The teeth from Paraná could pertain to a new species of Carcharodon already known from Perú. In contrast with the Pacific basin, Carcharodon sp. was sympatric with the wide toothed species Carcharodon plicatilis in central Argentina. This is the oldest record of Carcharodon in the southern Atlantic.     

Ostracodes et stratigraphie du néogène et du quaternaire méditerranéens

A stratigraphical chart of marine ostracoda from Lower Miocene to Recent is established. Selected species (approximatively 220) are those morphologicaly well characterized and known from different parts of the Mediterranean area. It appears that: • lower Miocene ostracodes are still poorly known; • specific diversity is high during the Tortonian and the Lower Messinian before the complete disappearance of marine Mediterranean species during the Upper Messinian évaporitic episodes; • during the early Pliocene, about half of the Upper Miocene marine species are reintroduced with the Atlantic waters; other species migrate for the first time in the Mediterranean Sea by the same way; • at the end of the Pliocene or at the beginning of the Pleistocene several species known in Mediterranean since the Middle Miocene or before, such as Cytherella sp. gr. transversa and Ruggieria tetraptera, as well some “nordic guests” such as Hemicythere villosa and Cythere lutea, appear. This work is an opportunity to confirme a Late Miocene age for the Neogene of Skyros (Aegean Sea), to assign the “Upper Pliocene” of Terquem to the Lower Pleistocene and to refute the existence of a pliocene psychrosphere.     

Upper Eocene to Middle Miocene ostracode faunas and paleo-oceanography of the North Coastal Belt,Jamaica, West Indies

The ostracode fauna of the Montpelier Group (Upper Eocene-Middle Miocene) exposed in the western part of the North Coastal Belt of Jamaica contains representatives of three paleo-environments. Allochthonous shelf and littoral species, characterized by abraded carapaces and eye tubercles, and a diverse group of archibenthal (slope) forms are present throughout. Species restricted to the World Ocean Psychrosphere (> 1000 m), notably Bradleya dictyon, Australoecia tipica, Agrenocythere hazelae and Macrocypris spp., first occur in sediments of earliest Miocene age (planktonic foraminiferal zone N4). Their presence establishes a minimum age for the entry into the Cayman Trench of frigid (<8–10°C) water masses drawn from the Atlantic thermohaline stratification.Upper Eocene and Oligocene assemblages are dominated by smooth, blind genera such as Krithe, Messinella and Bythocypris and lived in lower thermospheric conditions (10°C) at or near the base of the thermocline (700– 900 m). During the Early Miocene, downfaulting of the north island slope of Jamaica in response to southward tilting of the Nicaraguan Rise brought the Montpelier depositional site into contact with progressively deeper and colder water-masses. This subsidence culminated in the late Early Miocene; faunas of this age contain large numbers of Bradleya dictyon, Agrenocythere hazelae, Macrocypris sp. 1, Bairdia oarion and Bairdoppilata cassida. They suggest paleodepths of 1500–2000 m and the local presence of a 3–4° water-mass tentatively identified as the North Atlantic deep water. If this interpretation is correct, then the maximum sill-depth and water-mass configuration of the Miocene Cayman Trench have persisted to the present. Middle Miocene assemblages indicate a return to paleodepths of 1000–1500 m heralding the pronounced Upper Miocene to mid-Pleistocene tectonism which elevated Montpelier strata above sea level. An alternative view attributes the Miocene ostracode succession in the Montpelier to episodes in the development of the Atlantic circulation as recognized in the northeast Atlantic.Montpelier deposition took place either on a broad step intervening between the northern edge of the Clarendon Platform and the abyssal basins of the Cayman Trench or on a Tertiary analog of the existing north island slope. These results have emphasized the value of detailed taxonomic and morphologic studies of deep-sea ostracodes in geohistorical reconstruction.     

Zooplankton of the Patagonian slope and adjacent waters in the autumn

In autumn, from March until May 1988 in the Patagonian shelf and slope zones, the system of the water current was modified and water temperature decreased. The biomass of seston and abundance of zooplankton decreased simultaneously, especially those of the subantactric species Calanus simillimus. Zooplankton organisms started their descent to deeper waters. The Antarctic species Rhincalanus gigas was common in the upper layers in March. In May, R. gigas was rare and occurred in deeper layers. Subtropical zooplankton species dominated in the shelf zone, while subantartic species dominated in the slope zone within the Falkland current. In May, the abundance of shelf species in the slope zone increased due to weakening of the Falkland current and its dislocation to the east. The zooplankton biomass and abundance increased at the zone of water divergence in the east and at the slope frontal zone in the west.     

The autumn mesozooplankton community at South Georgia: biomass, population structure and vertical distribution

Mesozooplankton were sampled at shelf and oceanic stations close to South Georgia, South Atlantic during austral autumn 2004 with a Longhurst Hardy Plankton Recorder. Onshelf biomass ranged from 2.18 to 5.75 g DM m^?2 (0–200 m) and was dominated by the small euphausiid Thysanöessa spp. At the oceanic stations (10.57–14.71 g DM m^?2, 0–1,000 m) large calanoids, principally Rhincalanus gigas comprised ～47–52% of biomass. Here Calanus simillimus was still active and reproducing in surface waters (0–11.2 eggs fem day^?1) but R. gigas and Calanoides acutus were largely resident in the warm deep water and undergoing their seasonal descent. A comparison with spring and summer data indicated increased abundance and biomass from spring through to summer followed by a decline towards autumn particularly over the shelf. Autumn values in oceanic waters differed little from summer. Mesozooplankton biomass in the surface 200 m of the oceanic stations as a proportion of that found in the top 1,000 m ranged from 63 to 78% of the total in spring and 62–73% in summer, but was only 23–29% of the total in this study, following redistribution down the water column.     

Interspecific hybridization in oysters: Restriction Enzyme Digestion Chromosome Banding confirms Crassostrea angulata × Crassostrea gigas F1 hybrids

The taxonomic status of the two commercially important cupped oysters, Crassostrea angulata, the Portuguese oyster (Lamarck, 1819) and Crassostrea gigas, the Japanese oyster (Thunberg, 1793) has long been in question. The recent observation of the hybridization between C. gigas and C. angulata and the production of fertile F1s led us to search for cytogenetic evidence of both parental genomes in the interspecific hybrids. The cytogenetic characterization of the hybrids was performed by the use of restriction endonuclease treatments. This technique has recently shown the potential for individual chromosome identification by banding in oysters. Chromosomes of C. gigas, C. angulata and their hybrids were treated with two different restriction enzymes (ApaI and HaeIII), stained with Giemsa, and examined for banding patterns. These chromosome markers allowed the parental haploid sets to be identified in the hybrids. The analysis of the banded karyotypes of the interspecific hybrids showed that for each chromosome pair, one of the homologues presented a banding pattern consistent with that of C. gigas and the other homologue presented a banding pattern consistent with that of C. angulata. These cytogenetic results substantiate the reported interspecific hybridization between C. gigas and C. angulata. In view of these results and taking into account the present expansion of C. gigas aquaculture in southern Europe, the question of the need for preservation of pure C. angulata stocks should be raised as only a few populations remain in the south of Spain and Portugal. Recently, changes in the genetic composition of populations in southern Portugal have indeed been observed, showing that human activities have created contact zones between the two taxa while no natural sympatric zones exist in Europe.     

Depth distribution of the percophidMatsubaraea fusiforme in Fukiagehama beach,Kyushu

Collections with a small beam trawl and push-net at Fukiagehama beach, Kyushu, revealed that individuals of the percophid,Matsubaraea fusiforme, usually inhabit water not exceeding 5 m deep along sandy beaches during warm seasons. They are especially abundant along the low water line of spring tide regardless of the tidal phase. They may over winter in the deeper waters. Their length composition suggests protandry in the species.     

Response of deep-sea benthic foraminifera to Miocene paleoclimatic events, DSDP Site 289

Changes in the Miocene deep-sea benthic foraminifera at DSDP Site 289 closely correlate to the climatically induced variations in deep and bottom waters in the Pacific Ocean. In early Miocene time, oxygen and carbon isotopes indicate that bottom waters were relatively warm and poorly oxygenated. Benthic foraminiferal assemblages are characterized by various species inherited from the Oligocene. Expansion of the Antarctic icecap in the early middle Miocene, 14–16 m.y. ago, increased oxygen isotope values, produced cold, more oxygenated bottom waters and lead to a turnover in the benthic foraminifera. An Oligocene—early Miocene assemblage was replaced by a cibicidoid-dominated assemblage. Some species became extinct and benthic faunas became more bathymetrically restricted with the increased stratification of deep waters in the ocean. In mid-Miocene time, Epistominella exigua and E. umbonifera, indicative of young, oxygenated bottom waters, are relatively common at DSDP Site 289. Further glacial expansion 5–9 m.y. ago lowered sealevel, increased oceanic upwelling and associated biological productivity and intensified the oxygen minima. Abundant hispid and costate uvigerines become a dominant faunal element at shallow depths above 2500 m as E. umbonifera becomes common to abundant below 2500 m. By late Miocene time, benthic faunas similar in species composition and proportion to modern faunas on the Ontong-Java plateau, had become established.     

Co-occurring Synechococcus ecotypes occupy four major oceanic regimes defined by temperature,macronutrients and iron

Marine picocyanobacteria, comprised of the genera Synechococcus and Prochlorococcus, are the most abundant and widespread primary producers in the ocean. More than 20 genetically distinct clades of marine Synechococcus have been identified, but their physiology and biogeography are not as thoroughly characterized as those of Prochlorococcus. Using clade-specific qPCR primers, we measured the abundance of 10 Synechococcus clades at 92 locations in surface waters of the Atlantic and Pacific Oceans. We found that Synechococcus partition the ocean into four distinct regimes distinguished by temperature, macronutrients and iron availability. Clades I and IV were prevalent in colder, mesotrophic waters; clades II, III and X dominated in the warm, oligotrophic open ocean; clades CRD1 and CRD2 were restricted to sites with low iron availability; and clades XV and XVI were only found in transitional waters at the edges of the other biomes. Overall, clade II was the most ubiquitous clade investigated and was the dominant clade in the largest biome, the oligotrophic open ocean. Co-occurring clades that occupy the same regime belong to distinct evolutionary lineages within Synechococcus, indicating that multiple ecotypes have evolved independently to occupy similar niches and represent examples of parallel evolution. We speculate that parallel evolution of ecotypes may be a common feature of diverse marine microbial communities that contributes to functional redundancy and the potential for resiliency.     

Miocene spores and pollen from Pelitçik Basin,Turkey–environmental and climatic implications

The palynological investigation of the early Middle Miocene fluviolacustrine sedimentary rocks interfingering with volcanics of the Galatean Volcanic Province at Pelitçik Basin (Central Turkey) have yielded palynomorphs belonging to 51 spore and pollen taxa. The pollen record is dominated by Ulmus, Pinaceae, Quercus, Carpinus and Carya, and appears to reflect climatic conditions. Two pollen zones were established based on changing abundances of plant taxa. Zone 1 is characterized by dominance of Ulmus and Pinaceae. Zone 2 is differentiated and characterized by a dramatic increase in, and predominance of, Ulmus, Carya, deciduous Quercus, Carpinus, Salix and Pinaceae. Mixed mesophytic forests were widespread in the basin suggesting warm and temperate climate. The decreasing relative percentage of thermophilous taxa, with Engelhardia as the main component, and warm temperate taxa, such as Carya, at the upper part of Zone 2 generally indicates a climatic deterioration, probably related to the Middle Miocene cooling.     

Miocene Shark and Batoid Fauna from Nosy Makamby (Mahajanga Basin,Northwestern Madagascar)

Madagascar is well known for producing exceptional fossils. However, the record for selachians remains relatively poorly known. Paleontological reconnaissance on the island of Nosy Makamby, off northwest Madagascar, has produced a previously undescribed assemblage of Miocene fossils. Based on isolated teeth, ten taxonomic groups are identified: Otodus, Carcharhinus, Galeocerdo, Rhizoprionodon, Sphyrna, Hemipristis, Squatina, Rostroraja, Himantura and Myliobatidae. Six are newly described from Madagascar for the Cenozoic (Galeocerdo, Rhizoprionodon, Sphyrna, Squatina, Rostroraja and Himantura). In association with these specimens, remains of both invertebrates (e.g., corals, gastropods, bivalves) and vertebrates (e.g., bony fish, turtles, crocodylians, and sirenian mammals) were also recovered. The sedimentary facies are highly suggestive of a near-shore/coastal plain depositional environment. This faunal association shares similarities to contemporaneous sites reported from North America and Europe and gives a glimpse into the paleoenvironment of Madagascar’s Miocene, suggesting that this region was warm, tropical shallow-water marine.     

Isolation of Pandemic Vibrio parahaemolyticus from UK Water and Shellfish Produce

Vibrio parahaemolyticus is a Gram-negative, halophilic bacterium found commonly in temperate and warm estuarine waters worldwide. V. parahaemolyticus is considered an emerging bacterial pathogen in Europe and has been responsible for several recent seafood-associated outbreaks. During ad hoc testing of raw shellfish produce in May 2012, pandemic group (O3:K6) V. parahaemolyticus was isolated from Pacific oysters (Crassostrea gigas), harvested in Southern England. Follow-on testing of water and shellfish, encompassing a small number geographically diverse sites, also retrieved pandemic group isolates. These strains are amongst the most northerly pandemic strains described to date and represent the first instance of pandemic V. parahaemolyticus isolated in the UK, highlighting the expanding geographical distribution of these foodborne pathogens in the environment.     

Late Miocene ostracod assemblages from eastern Mediterranean coral reef complexes (central Crete,Greece)

Ostracods from ten Late Miocene coral reef complexes built by Siderastrea, Tarbellastrea and Porites, cropping out in the Messara Plain (southern Iraklion basin, central Crete), have been investigated and five assemblages have been recognised, which point to different marine environments: (1) assemblage from the basal sandy silts, dominated by very shallow inner-infralittoral species, such as Cyamocytheridea meniscus, Cyamocytheridea obstipa, Cyamocytheridea dertonensis, Cytheretta semiornata and Nonurocythereis seminulum; (2) assemblage from the coral reef complexes within which Grinioneis haidingeri, Aurila cicatricosa, Cimbaurila diecii, Tenedocythere cruciata, Pokornyella italica and Callistocythere quadrangula are dominant and point to a stable inner-infralittoral environment characterised by warm, quiet and well-oxygenated waters; (3) assemblage from the silts intercalated among the coral reef complexes, mainly characterised by Neomonoceratina laskarevi, Cytheridea acuminata, Phlyctenophora farkasi and Aurila albicans together with Callistocythere spp., Xestoleberis communis and Xestoleberis dispar, which points to a very shallow marine environment rich in aquatic vegetation; (4) assemblage from the upper silts, which records the absolute dominance of Xestoleberis species, reflecting a very shallow and highly-vegetated environment and (5) assemblage from the uppermost silty clays, dominated by Hemicytherura defiorei, Xestoleberis spp. and Palmoconcha dertobrevis, accompanied by Acanthocythereis hystrix, Cytherella scutulum, Bairdoppilata conformis, Semicytherura spp., Krithe sp., Cytheropteron alatum, Bythocypris sp. and Pseudocythere caudata, which suggest deeper marine environments probably located in the outer infralittoral/inner-circalittoral zones. The studied section has been dated by means of calcareous nannoplankton to be not younger than Zone MNN9 (Early Tortonian), which is the biostratigraphical datum recorded in the fine-grained deposits that overlie the coral reef complexes. An age not older than Tortonian can be inferred by the stratigraphical distribution of the recognized ostracods. Thus, the coral reef complexes have been tentatively referred to the Early Tortonian.     

Facing the Heat: Thermoregulation and Behaviour of Lowland Species of a Cold-Dwelling Butterfly Genus,Erebia

Understanding the potential of animals to immediately respond to changing temperatures is imperative for predicting the effects of climate change on biodiversity. Ectothermic animals, such as insects, use behavioural thermoregulation to keep their body temperature within suitable limits. It may be particularly important at warm margins of species occurrence, where populations are sensitive to increasing air temperatures. In the field, we studied thermal requirements and behavioural thermoregulation in low-altitude populations of the Satyrinae butterflies Erebia aethiops, E. euryale and E. medusa. We compared the relationship of individual body temperature with air and microhabitat temperatures for the low-altitude Erebia species to our data on seven mountain species, including a high-altitude population of E. euryale, studied in the Alps. We found that the grassland butterfly E. medusa was well adapted to the warm lowland climate and it was active under the highest air temperatures and kept the highest body temperature of all species. Contrarily, the woodland species, E. aethiops and a low-altitude population of E. euryale, kept lower body temperatures and did not search for warm microclimates as much as other species. Furthermore, temperature-dependence of daily activities also differed between the three low-altitude and the mountain species. Lastly, the different responses to ambient temperature between the low- and high-altitude populations of E. euryale suggest possible local adaptations to different climates. We highlight the importance of habitat heterogeneity for long-term species survival, because it is expected to buffer climate change consequences by providing a variety of microclimates, which can be actively explored by adults. Alpine species can take advantage of warm microclimates, while low-altitude grassland species may retreat to colder microhabitats to escape heat, if needed. However, we conclude that lowland populations of woodland species may be more severely threatened by climate warming because of the unavailability of relatively colder microclimates.     

Palaeoenvironmental significance of Miocene larger benthic foraminifera from the Xisha Islands,South China Sea

We studied 344 samples from Well XK-1 in Xisha Islands, South China Sea, and identified 66 species of larger benthic foraminifera, providing critical evidence for biostratigraphy and palaeoenvironmental interpretation of the Miocene reef carbonate sequence. Three assemblages are recognized, namely, Spiroclypeus higginsi–Borelis pygmaeus Assemblage (Letter Stage Te5, Early Miocene, 1256.28–1180.15 m), Nephrolepidina–Miogypsina Assemblage (Tf, Middle Miocene, 1031.10–577.04 m), and Cycloclypeus–Heterostegina Assemblage (Tg, Late Miocene, 468.13–380.42 m). On the basis of the palaeoecological preference of the larger foraminifera, we interpret that the Miocene carbonate sequence was deposited mainly in a warm tropical shallow water environment, characterized by five stages of continuous long-term evolution: backreef lagoon to shelf in the Early Miocene, normal to frontal reef in the early Middle Miocene, backreef lagoon to shelf in the later Middle Miocene, normal to frontal reef in the early Late Miocene, and proximal forereef shelf in the later Late Miocene.     

Clonal integration facilitates invasiveness of the alien aquatic plant Myriophyllum aquaticum L. under heterogeneous water availability

The avoidance of visually feeding fish has long been considered as the primary driver of diel vertical migration of zooplankton. The diurnal vertical distribution of Cyclops gr. abyssorum, Arctodiaptomus alpinus, and Daphnia gr. longispina from 13 alpine lakes with fish (Salvelinus fontinalis) and without, was compared in order to understand whether fish in transparent lakes reduce the presence of large zooplankton from the irradiated zone. We used the light level at each sampling depth and the size of each specimen as proxies of predation risk, and we tested two predictions: (P1) the relative abundance of zooplankton in the well-lit surface waters vs. the darker waters will be greater in fishless lakes; (P2) the size of zooplankton in the well-lit surface waters vs. the deeper, darker waters will be greater in fishless lakes. We did not find any evidence of the validity of P1, but we confirmed P2 for Arctodiaptomus alpinus. These results support with new field data the Transparency Regulator Hypothesis, which argues that in transparent lakes, fish predation is less important for the vertical distribution of zooplankton than ultraviolet radiation, and further suggest that zooplankton size rather than vertical distribution may be more effective against visual predators in transparent lakes.     

Genetics, recruitment, and migration patterns of Arctic cisco (Coregonus autumnalis) in the Colville River, Alaska, and Mackenzie River, Canada

Arctic cisco Coregonus autumnalis have a complex anadromous life history, many aspects of which remain poorly understood. Some life history traits of Arctic cisco from the Colville River, Alaska, and Mackenzie River basin, Canada, were investigated using molecular genetics, harvest data, and otolith microchemistry. The Mackenzie hypothesis, which suggests that Arctic cisco found in Alaskan waters originate from the Mackenzie River system, was tested using 11 microsatellite loci and a single mitochondrial DNA gene. No genetic differentiation was found among sample collections from the Colville River and the Mackenzie River system using molecular markers (P > 0.19 in all comparisons). Model-based clustering methods also supported genetic admixture between sample collections from the Colville River and Mackenzie River basin. A reanalysis of recruitment patterns to Alaska, which included data from recent warm periods and suspected changes in atmospheric circulation patterns, still finds that recruitment is correlated to wind conditions. Otolith microchemistry (Sr/Ca ratios) confirmed repeated, annual movements of Arctic cisco between low-salinity habitats in winter and marine waters in summer.     

Stable isotopic study of Oligocene-Miocene sediments from DSDP Site 354, Equatorial Atlantic

The oxygen- and carbon-isotope compositions of planktic and benthic foraminifera and calcareous nannofossils from Middle Oligocene-Early Miocene Equatorial Atlantic sediments (DSDP Site 354) indicate two important paleoceanographic changes, in the Late Oligocene (foraminiferal Zone P.21) and in the Early Miocene (foraminiferal Zone N.5). The first change, reflected by a δ¹⁸O increase of 1.45‰ inGlobigerina venezuelana, affected only intermediate pelagic and not surface, deep or bottom waters. The second change affected surface and intermediate waters, whereas deep and bottom waters showed only minor fluctuations. In the case of the former the isotope effect of the moderate ice accumulation on the Antarctic continent is amplified in the Equatorial Atlantic by changes in the circulation pattern. The latter paleoceanographic change, reflected by a significant increase in¹⁸O in both planktic and benthic forms (about 1.0‰ and 0.5‰, respectively), may have been caused by ice volume increase and temperature decrease. Both oxygen- and carbon-isotope compositions indicate a marked depth-habitat stratification for planktic foraminifera and calcareous nannofossils. Three different dwelling groups are recognized: shallowGlobigerinoides, Globoquadrina dehiscens, Globorotalia mayeri and nannofossils; intermediateGlobigerina venezuelana; and deepCatapsydrax dissimilis. The comparison of foraminifera and calcareous nannofossils suggests that the isotopic compositions of nannofossils are generally controlled by the same parameters which control the isotopic composition of shallow-dwelling foraminifera, but the former are more enriched in¹⁸O.