Further observations on the genus Pappomonas Manton et Oates with special reference to P. virgulosa sp. nov. from West Greenland

Electron microscopy of dried preparations of Arctic nanoplankton has permitted a second species of the recently erected genus Pappomonas Manton et Oates to be described and named. P. virgulosa sp. nov. is characterised by coccolith appendages composed of a central shaft ending in a cluster of four finger-like rods. There are also calcified plates of characteristic construction, without appendages. Detailed observations on the coccoliths, especially on the shapes and arrangement of component calcite crystallites and the presence of unmineralised components, have permitted further comparisons to be made with equivalent structures in Papposphaera lepida Tangen, recently investigated in the southern hemisphere, as well as with Pappomonas flabellifera Manton et Oates. As a working hypothesis, an interpretation of coccolith morphology in all three taxa has been formulated in terms of known structural components of uncalcified scales in Chrysochromulina spp. The need for fuller information on certain details, notably the coccolith bases in Pappomonas spp. is stressed.     

Description of diatoms from the Southwest to West Greenland coastal and open marine waters

Detailed studies on sub-Arctic and Arctic marine diatom assemblages contribute to the understanding of spatial distribution patterns and their physical drivers. In this study, diatom species were analyzed from water samples collected with a Niskin bottle rosette combined with a CTD along the West Greenland coast (63°58′N–71°08′N and 50°49′W–59°06′W) during summer 2007. Diatom community was represented mainly by three genera Thalassiosira, Fragilariopsis, and Chaetoceros and linked to observed hydrographic and environmental conditions. Thalassiosira spp. were common in coastal waters (particularly Godthåbsfjord) and linked to increased surface water temperature, typical of summer water stratification in West Greenland. Fragilariopsis spp., along with other dominant species associated with higher geographic latitudes, dominated in Arctic fjords (Uummannaq Fjord-Qaumarujuk Fjord). These species generally characterized coastal waters influenced by melting sea ice and/or glacial ice. Chaetoceros spp. were linked to more saline open marine waters, particularly in the Davis Strait south of 70°N, probably corresponding to weaker water stratification and the influence of the West Greenland Current. The present paper provides new knowledge on diatom assemblages along a south–north climate gradient in West Greenland, which is necessary in order to understand how observed ocean-climate changes influence Arctic marine ecosystems. This study provides a reference for palaeoenvironmental reconstructions using diatom microfossils deposited in the West Greenland marine sediments.     

Contribution to the rotifer fauna of subarctic Greenland (Kangerlussuaq and Ammassalik area)

Six water bodies in the region of Kangerlussuaq, West Greenland (67° 30 N, 50° 46 W), and four near Ammassalik, East Greenland (65° 36 N, 37° 38 W) were sampled. Sixty-nine taxa (2 Bdelloidea, 67 Monogononta) are reported, forty-six of which represent new records for Greenland. Proales pejleri n. sp. is described.     

Heterotrophic amoebae,flagellates and heliozoa from Arctic marine waters (North West Territories,Canada and West Greenland)

Summary Thirty eight taxa of heterotrophic protists (amoebae, flagellates and heliozoa) were encountered in cultures established from marine samples from Disko Bay (West Greenland) and Foxe Basin (N.W.T. Canada). Most species are flagellates representing the choanoflagellates, the euglenids, the kinetoplastids, the bicosoecids, the chrysomonads, the pedinellids and a variety of taxa of uncertain affinities (Protista incertae sedis). A new species of thecate choanoflagellate, Salpingoeca intermedia sp. nov., is described, and 4 new types of triangular scales, representing the genus Thaumatomastix Lauterborn, 1899, are presented.     

A review of the genetic relationships of Atlantic walrus (Odobenus rosmarus rosmarus) east and west of Greenland

Studies of the genetic variation involving allozymes, mitochondrial and nuclear DNA (microsatellites) in walruses (Odobenus rosmarus) were reviewed. In addition, the genetic relationships of a total of 211 Atlantic walruses, O. r. rosmarus, from 5 sampling areas west and east of Greenland were studied using 12 nuclear DNA-microsatellite loci and restriction fragment length polymorphism obtained from the ND1, ND2 and ND3/4 segments of the mitochondrial DNA (mtDNA). At the mtDNA level, no divergence was observed among the three sampling areas east of Greenland (i.e. East Greenland, Svalbard and Franz Josef Land), whereas areas west of Greenland (i.e. Northwest and West Greenland) showed some differentiation. The genetic variation at the microsatellite loci grouped the individuals into four sub-populations: Northwest Greenland, West Greenland, East Greenland and a common Svalbard-Franz Josef Land sub-population. A significant correlation between genetic distance and geographic distance between the sampling areas (isolation-by-distance effect) was detected, especially at the mtDNA level. At a small-scale phylo-geographical level, the mtDNA data indicated that Atlantic walruses have diverged into two major groups: one northwest (i.e. in the North Water) and one east of Greenland (i.e. an East Greenland-Svalbard-Franz Josef Land group), whereas the haplotype distribution in the West Greenland sample reflected a mixture of both these groups. The microsatellite data supported a general grouping of walruses to the west and east of Greenland.     

EFFECTS OF NITROGEN AND PHOSPHORUS AVAILABILITY ON THE EXPRESSION OF THE COCCOLITH‐VESICLE V‐ATPASE (SUBUNIT C) Of PLEUROCHRYSIS (HAPTOPHYTA)1

The coccolithophores Emiliania and Pleurochrysis demonstrate increased coccolith production when growth is reduced by nitrate or phosphate limitation. The function of enhanced coccolith production under these conditions and its regulation have not been resolved. Studies at the molecular level are ideally suited to determine the exact relationship between calcification and other cellular functions. In a previous study we provided evidence for the presence of a vacuolar H⁺‐ATPase on coccolith vesicle membranes of P. carterae. These trans‐Golgi–derived vesicles are the sites of coccolith production, with each vesicle containing one coccolith. In this study, expression of the vap gene, encoding subunit c of the vacuolar H⁺‐ATPase, was investigated. Our objective was to explore potential relationships between vap expression, nutrient‐dependent growth, and calcification. Specifically, we monitored vap expression relative to two genes, fcp and pcna, whose expression was previously shown to vary with growth conditions; fcp encodes a fucoxanthin chl a/c‐binding protein, and pcna encodes the proliferating cell nuclear antigen. Relative to the expression of pcna and fcp, vap expression was highest at nutrient concentrations where growth curves and chl a patterns indicated arrest of cell division. Our results indicate that the level of vap expression does not decrease when cell growth diminishes.     

Collaborative Moments. Expanding the Anthropological Field through Cross-Disciplinary Practice

ABSTRACT

This article explores some implications of cross-disciplinarity, as experienced in practice. Anthropologists are used to fuse different styles of reasoning by integrating the points of view and unspoken certainties of their partners in the field into their analysis. Fieldwork can be seen as an experiment in real time, where insights gained intersubjectively gradually shape up as knowledge through analysis. This line of thought is brought to bear on a discussion of collaboration between anthropologists, archaeologists, and biologists in North West Greenland. Through actual experiences from the field, this article shows how knowledge generated on the edge of one’s familiar disciplinary territory may both expand and intensify the anthropological field. Collaborative moments are seen to make new anthropological insights emerge through the co-presence of several analytical perspectives in the field.     

Genetic population structure of Littorina brevicula around Korean waters

Originally described from Spitzbergen, Greenland, Iceland and Scandinavia, Chaetozone setosa is now widely recognised to be a species complex by workers involved in analysing survey material in the northeast Atlantic. Chaetozone species were examined from wide-scale surveys of the Faroe-Shetland Channel and nearby areas, which included water depths ranging from 100 to 2000 m and diverse hydrographic conditions. Comparisons were made with other deep-water material from around Iceland. There appear to be at least five Chaetozone species occurring in the cold, deeper waters of the area. One of the commonest of these, from the lower West Shetland Continental Slope (Faroe-Shetland Channel) and the Iceland-Faroe Rise, is described as a new species.     

Formation and mosaicity of coccolith segment calcite of the marine algae Emiliania huxleyi

Coccolithophores belong to the most abundant calcium carbonate mineralizing organisms. Coccolithophore biomineralization is a complex and highly regulated process, resulting in a product that strongly differs in its intricate morphology from the abiogenically produced mineral equivalent. Moreover, unlike extracellularly formed biological carbonate hard tissues, coccolith calcite is neither a hybrid composite, nor is it distinguished by a hierarchical microstructure. This is remarkable as the key to optimizing crystalline biomaterials for mechanical strength and toughness lies in the composite nature of the biological hard tissue and the utilization of specific microstructures. To obtain insight into the pathway of biomineralization of Emiliania huxleyi coccoliths, we examine intracrystalline nanostructural features of the coccolith calcite in combination with cell ultrastructural observations related to the formation of the calcite in the coccolith vesicle within the cell. With TEM diffraction and annular dark‐field imaging, we prove the presence of planar imperfections in the calcite crystals such as planar mosaic block boundaries. As only minor misorientations occur, we attribute them to dislocation networks creating small‐angle boundaries. Intracrystalline occluded biopolymers are not observed. Hence, in E. huxleyi calcite mosaicity is not caused by occluded biopolymers, as it is the case in extracellularly formed hard tissues of marine invertebrates, but by planar defects and dislocations which are typical for crystals formed by classical ion‐by‐ion growth mechanisms. Using cryo‐preparation techniques for SEM and TEM, we found that the membrane of the coccolith vesicle and the outer membrane of the nuclear envelope are in tight proximity, with a well‐controlled constant gap of ~4 nm between them. We describe this conspicuous connection as a not yet described interorganelle junction, the “nuclear envelope junction”. The narrow gap of this junction likely facilitates transport of Ca²⁺ ions from the nuclear envelope to the coccolith vesicle. On the basis of our observations, we propose that formation of the coccolith utilizes the nuclear envelope–endoplasmic reticulum Ca²⁺‐store of the cell for the transport of Ca²⁺ ions from the external medium to the coccolith vesicle and that E. huxleyi calcite forms by ion‐by‐ion growth rather than by a nanoparticle accretion mechanism.     

Ultrastructure and taxonomy ofJomonlithus littoralis gen. et sp. nov. (Class prymnesiophyceae), a coccolithophorid from the Northwest Pacific

A new coccolithophorid genusJomonlithus withJ. littoralis as the type species, is described based on specimens isolated from a sand sample collected at the mouth of Nakagawa river, Ibaraki, Japan. This genus is characterized by the coccolith which is composed of an organic base-plate scale and calcified rim elements made up of two different subelements.J. littoralis has been found in several localities along the coast of Japan. Culture and ultrastructure studies gave special attention to the cell cycle, coccolith and scale morphology and the ultrastructure of the cellular organelles. It was confirmed thatJomonlithus is similar toWigwamma Mantonet al., Papposphaera Tangen andPappomonas Manton et Oates on the basis of coccolith morphology and toCricosphaera Braarud andHymenomonas Stein in cellular structure.     

Three Types of Acidic Polysaccharides Associated with Coccolith of Pleurochrysis haptonemofera: Comparison with Pleurochrysis carterae and Analysis Using Fluorescein-Isothiocyanate-Labeled Lectins

In the coccolithophorid microalgae acidic polysaccharides are considered to be involved in the formation of the calcified scale, coccolith. Characteristics of the acidic polysaccharides extracted from the cell surface of the coccolithophorid Pleurochrysis haptonemofera were analyzed. The acidic polysaccharides on the cell surface can be detected by measuring fluorescence of cells after fluorescein-isothiocyanate-labeled lectin staining by flow cytometry. Flow cytometric analyses revealed that the acidic polysaccharides remained on the cell surface even after CaCO₃ in the coccolith was dissolved by lowering pH, but they were extracted by subsequent EDTA or EGTA treatment, suggesting that they are bound not into the CaCO₃ crystals of the coccolith, but onto the surface via Ca²⁺. Analyses of the acidic polysaccharides by anion exchange chromatography, colloidal precipitation with divalent cations, and polyacrylamide gel electrophoresis (PAGE) revealed that P. haptonemofera has 3 types of acidic polysaccharides (Ph-PS-l, -2, and -3). The PAGE patterns suggested that Ph-PS-2 has a repeated structure with a broad range of molecular weight, as in Pleurochrysis carterae, while Ph-PS-1 and -3 contain several minor components in addition to a major component, respectively. The minor components in Ph-PS-1 and -3 that have not been found in P. carterae might be characteristic of P. haptonemofera. Analyses of both the cell surface treated by various concentrations of EDTA and EGTA and the extracts suggested that Ph-PS-2, which is distinguishable by a higher affinity to concanavalin A, is bound onto the coccolith surface more intensely than the other two types of acidic polysaccharides.     

Polyanion-mediated mineralization — assembly and reorganization of acidic polysaccharides in the Golgi system of a coccolithophorid alga during mineral deposition

Summary Immunolocalization of two highly acidic polysaccharides (PS-1 and PS-2) in a calcifying algaPleurochrysis carterae is described throughout the mineralization process, from before crystal nucleation through the cessation of crystal growth. This unicellular coccolithophorid alga is a useful model for mineralization because it produces calcified scales known as coccoliths in homogeneous cell culture. PS-1 and PS-2 were localized in the crystal coats of mature coccoliths and in electron dense Golgi particles. The polyanions are synthesized in medial Golgi cisternae and co-aggregate with calcium ions into discrete 25 nm particles. Particle-laden vesicles bud from cisternal margins and fuse with a coccolith-forming saccule containing an organic oval-shaped scale which forms the base of the future coccolith. The particles are localized on the base before the onset of mineral deposition and are present in the coccolith saccule throughout the period of crystal (CaCO₃) nucleation and growth. During the final phase of coccolith formation, the particles disappear, and the mature crystals acquire an amorphous coat containing PS-1 and PS-2 polysaccharides which remain with the mineral phase after the coccoliths are extruded from the cell. Postulated mechanisms of polyanion-mediated mineralization are reviewed and their relevance to the calcification of coccoliths is addressed.Abbreviations PS-1 polysaccharide one - PS-2 polysaccharide two - BSA bovine serum albumin - SDS sodium dodecyl sulfate - MES 2-(N-morpholino)-ethanesulfonic acid - EDTA ethylenediaminetetraacetic acid - DHA 3-deoxy-lyxo-2-heptulosaric acid - TCA trichloroacetic acid     

Global diversity of two widespread,colony‐forming diatoms of the marine plankton,Chaetoceros socialis (syn. C. radians) and Chaetoceros gelidus sp. nov.

Marine phytoplankton samples containing diatoms of the Chaetoceros socialis group were collected from Thailand, China, Denmark, and Greenland, and cells were isolated into culture for light and electron microscopy and DNA sequencing of D1–D3 of the LSU rDNA. Species of this lineage are characterized by three short and one long setae extending from each cell, the long setae from several cells joining into a common center to form large colonies, which are sometimes visible with the naked eye. Phylogenetic analyses including sequences from other parts of the world revealed segregation into three groups. Most sequences fell into two large clades, one comprising material from cold waters, whereas the other contained material from warmer waters. Strain CCMP 172 from the Strait of Georgia, Washington State, USA, formed a separate group. The warm‐water species included Chinese and Thai material and therefore probably also material from the type locality of C. socialis, Hong Kong. It is characterized by all setae being covered by spines and the setae extending from the valve at some distance from the margin. In the resting spores, both valves are ornamented with spines. The cold‐water material is characterized by three spiny and one mostly smooth long setae, and the setae extend from the valve near the margin. Both valves of the resting spore are smooth. This material is described as C. gelidus sp. nov. C. radians, described from the Baltic in 1894, is considered a synonym of C. socialis. CCMP172 is in many ways intermediate and probably constitutes a separate species. The published evidence on this globally distributed and sometimes bloom‐forming group of species indicates higher species diversity than presently thought.     

Seasonal occurrence of coccoliths in sediment traps from West Caroline Basin, equatorial West Pacific Ocean

Coccolith fluxes were investigated by sediment trap studies in the West Caroline Basin, which is located in the equatorial western Pacific. The investigation was conducted from June 1991 to March 1992 at two water depths, 1592 and 3902 m, as part of the Northwest Pacific Carbon Cycle Study (NOPACCS) program. Two seasonal maxima of coccolith fluxes were observed during September–early October and late December–January. The average coccolith and coccosphere fluxes at the depth of the shallow trap were 1800×10⁶ coccoliths m⁻² day⁻¹ and 1.9×10⁶ coccospheres m⁻² day⁻¹, respectively. The flux of coccoliths followed the same trend as the total flux, and was closely correlated with the flux of organic matter flux. Florisphaera profunda, Gladiolithus flabellatus, Gephyrocapsa oceanica, Umbilicosphaera sibogae var. sibogae, Emiliania huxleyi, and Oolithotus fragilis were the most abundant species together comprising more than 85% of the total flora. Observed seasonal changes of the species composition of coccolith flora, as well as analysis of the R-mode cluster, revealed that during the summer, the assemblage was marked by the dominance of G. oceanica and U. sibogae. However, during the winter, the assemblage was dominated by E. huxleyi and O. fragilis. These assemblage changes were influenced by monsoonal events, which were observed off the New Guinea coast. F. profunda dominated the community in the shallow trap throughout most of the year; peak values of this species were recorded during the winter. The coccosphere assemblage was dominated by G. oceanica at both water depths. In the deep trap, the sedimentation pattern was similar to that observed at the shallow depth. Mean coccolith and coccosphere fluxes at the deep trap were 2000×10⁶ coccolith m⁻² day⁻¹ and 0.08×10⁶ coccospheres m⁻² day⁻¹, respectively. The increase in coccolith flux with water depth suggests a lateral influx. The estimated average daily mass of CaCO₃ flux in coccoliths and coccospheres was 16.6 mg m⁻² day⁻¹ at the 1592 m trap and 17.9 mg m⁻² day⁻¹ at the 3902 m trap, respectively. These calculated values contributed only 23.3% to the total CaCO₃ flux at the shallow trap and 27.9% at the deep trap.     

Coccolithophores in the equatorial Atlantic Ocean: response to seasonal and Late Quaternary surface water variability

The present study was initiated to ascertain the significance of coccolithophores as a proxy for paleoceanographic and paleoproductivity studies in the equatorial Atlantic. Data from a range of different samples, from the plankton, surface sediments as well as sediment cores are shown and compared with each other.In general, the living coccolithophores in the surface and subsurface waters show considerable variation in cell numbers and distribution patterns. Cell densities reached a maximum of up to 300×10³ coccospheres/l in the upwelling area of the equatorial Atlantic. Here, Emiliania huxleyi is the dominant species with relatively high cell numbers, whereas Umbellosphaera irregularis and Umbellosphaera tenuis are characteristic for oligotrophic surface waters. Although they are observed in high relative abundances, these species only occur in low absolute numbers. The lower photic zone is dominated by high abundances and considerable cell numbers of Florisphaera profunda.The geographical distribution pattern of coccoliths in surface sediments reflects the conditions of the overlying surface water masses. However, abundances of the oligotrophic species Umbellosphaera irregularis and Umbellosphaera tenuis are strongly diminished, causing an increase in relative abundance of the lower photic zone taxa Florisphaera profunda and Gladiolithus flabellatus.During the past 140,000 years the surface water circulation of the equatorial Atlantic has changed drastically, as can be seen from changes in the coccolithophore species composition, absolute coccolith numbers, as well as coccolith accumulation rates. Significant increases in coccolith numbers and accumulation rates is observed in the southern equatorial Atlantic during the last glacial interval (oxygen isotope stages 2–4), which we attribute to enhanced upwelling intensities and advection of cool nutrient rich waters at this site. In the western equatorial Atlantic we observe an opposite trend with decreasing numbers of coccoliths during glacial periods, which probably is caused by a deepening of the thermocline.     

Pelagic primary production during summer along 65 to 72°N off West Greenland

The distribution of phytoplankton biomass and primary production were studied during summer 1993 at 16 stations from 65 to 72°N off West Greenland, ranging more than 900 km. Hydrography, nutrients and chlorophyll a profiles revealed a significant change in structure from south to north. Nitrate was depleted in the euphotic zone at most stations except close to the ice edge (West Ice) or close to outflow from large glaciers. The vertical distribution of phosphate followed that of nitrate, but was never depleted. Despite two stations with relatively high surface concentrations, silica showed the same distribution as the other two nutrients. In the south, chlorophyll a concentration and primary production were lower than north of Disko Bay (69°N), associated with a well-mixed versus a salinity-generated stratification, respectively. In Vaigat, a high-production station was identified, (st. 910, 69°52′69N–51°30′61W) with a chlorophyll a concentration in the euphotic zone of >13 μg l⁻¹ and an area primary production of 3.2 g C m⁻² day⁻¹. This is seldom encountered in arctic waters and was presumably due to nutrient-rich melt-water originating from the Iluliíssat Glacier. The overall primary production for the studied area was 67–3207 mg C m⁻² day⁻¹ (mean ± SD=341± 743 mg C m⁻² day⁻¹), which is within the range of the few results published for West Greenland and eastern Canadian Arctic waters. Accepted: 24 October 1998     

Structural Studies on Diurodrilus Remane (Diurodrilidae fam.n.), with Description of Diurodrilus westheidei sp.n. from the Arctic Interstitial Meiobenthos,W. Greenland*

Diurodrilus westheidei sp.n. is described in terms of the location of the sensoria, ventral ciliated cells, ciliophores, adhesive toes and a big anal cone. Specimens were collected in mid winter at Disko Island, West Greenland. The ultrastructure of D. subterraneus Remane, collected in the summer at Disko Island and Ystad, Sweden, is discussed with new information on the systematic status of this aberrant genus as indicated by the cuticle, pharyngeal area, ciliophores and the spermatozoa. Diurodrilidae fam.n. is introduced for the genus Diurodrilus, here placed in Polychaeta.     

Effect of Coccolith Polysaccharides Isolated from the Coccolithophorid, <Emphasis Type="Italic">Emiliania huxleyi</Emphasis>, on Calcite Crystal Formation in In Vitro CaCO<Subscript>3</Subscript> Crystallization

Marine coccolithophorids (Haptophyceae) produce calcified scales “coccoliths” which are composed of CaCO₃ and coccolith polysaccharides (CP) in the coccolith vesicles. CP was previously reported to be composed of uronic acids and sulfated residues, etc. attached to the polymannose main chain. Although anionic polymers are generally known to play key roles in biomineralization process, there is no experimental data how CP contributes to calcite crystal formation in the coccolithophorids. CP used was isolated from the most abundant coccolithophorid, Emiliania huxleyi. CaCO₃ crystallization experiment was performed on agar template layered onto a plastic plate that was dipped in the CaCO₃ crystallization solution. The typical rhombohedral calcite crystals were formed in the absence of CP. CaCO₃ crystals formed on the naked plastic plate were obviously changed to stick-like shapes when CP was present in the solution. EBSD analysis proved that the crystal is calcite of which c-axis was elongated. CP in the solution stimulated the formation of tabular crystals with flat edge in the agarose gel. SEM and FIB-TEM observations showed that the calcite crystals were formed in the gel. The formation of crystals without flat edge was stimulated when CP was preliminarily added in the gel. These observations suggest that CP has two functions: namely, one is to elongate the calcite crystal along c-axis and another is to induce tabular calcite crystal formation in the agarose gel. Thus, CP may function for the formation of highly elaborate species-specific structures of coccoliths in coccolithophorids.     

Coccolithogenesis In Scyphosphaera apsteinii (Prymnesiophyceae)

Coccolithophores are the most significant producers of marine biogenic calcite, although the intracellular calcification process is poorly understood. In the case of Scyphosphaera apsteinii Lohmann 1902, flat ovoid muroliths and bulky, vase‐shaped lopadoliths with a range of intermediate morphologies may be produced by a single cell. This polymorphic species is within the Zygodiscales, a group that remains understudied with respect to ultrastructure and coccolith ontogeny. We therefore undertook an analysis of cell ultrastructure, morphology, and coccolithogenesis. The cell ultrastructure showed many typical haptophyte features, with calcification following a similar pattern to that described for other heterococcolith bearing species including Emiliania huxleyi. Of particular significance was the reticular body role in governing fine‐scale morphology, specifically the central pore formation of the coccolith. Our observations also highlighted the essential role of the inter‐ and intracrystalline organic matrix in growth and arrangement of the coccolith calcite. S. apsteinii secreted mature coccoliths that attached to the plasma membrane via fibrillar material. Time‐lapse light microscopy demonstrated secretion of lopadoliths occurred base first before being actively repositioned at the cell surface. Significantly, growth irradiance influenced the coccosphere composition with fewer lopadoliths being formed relative to muroliths at higher light intensities. Overall, our observations support dynamic metabolic (i.e., in response to growth irradiance), sensory and cytoskeletal control over the morphology and secretion of polymorphic heterococcoliths. With a basic understanding of calcification established, S. apsteinii could be a valuable model to further study coccolithophore calcification and cell physiological responses to ocean acidification.