Ecological investigations in the wadden area off Borkum, East Frisian Islands, Germany

Die Bodentiergemeinschaft des Wattenmeeres ist von Frühjahr bis Herbst eines jeden Jahres durch extrem hohe Dichten von Jungtieren charakterisiert. Die Kenntnisse über die Ansiedlung von fplanktischen Larven im Wattenmeer, sowie die Dynamik postlarvaler Stadien sind aufgrund der üblicherweise verwendeten, großen Siebmaschenweiten gering. Gerade aber diesen Altersstadien kommt möglicherweise eine besondere Stellung im Energiefluß des Wattenmeeres zu. An 5 Stationen (von NWL bis HWL, B1-B5) im Rückseitenwatt der ostfriesischen Insel Borkum wurden 1986 Ansiedlung, räumliche Verteilung, Wachstum, Mortalität und Produktion der Altersklasse 0 von Macoma balthica, Mya arenaria und Cerastoderma edule untersucht. Um die Ansiedlung der planktotrophen Larven dieser Arten zu beschreiben, wurden ihre Dichten in Plankton und Bodenproben miteinander verglichen. Die Untersuchungen zur Dynamik der benthischen Stadien wurden mit zwei in der Probenfläche und der Siebmaschenweite unterschiedlichen Probenserien durchgeführt. Die Drift postlarvaler Stadien wurde durch bodennahe Planktonfänge innerhalb des Eulitorals nachgewiesen. Parallel zu den Untersuchungen an der Endofauna wurden das Vorkommen und die Größe epibenthischer Räuber im Untersuchungsgebiet erfaßt. Die Hauptansiedlung von M. balthica- und M. arenaria-Larven erfolgte nahezu gleichzeitig Ende Mai/Anfang Juni. Die meisten Larven beider Arten gingen an der prielnächsten (tiefsten) Station (B1) zum Bodenleben über, gefolgt von der nächst höher gelegenen Station B2. Während frühe Bodenstadien von M. arenaria nicht im oberen Bereich des Watts (B3,B4) gefunden wurden, ist eine geringfügige Erstansiedlung von M. balthica in diesem Gebiet nicht auszuschließen. Ein die Ansiedlung limitierender Einfluß der relativ dichten Mya arenaria-Siedlung an den Stationen B1 und B2 sowie der Alttiere von M. balthica konnte nicht festgestellt werden. Die Ähnlichkeit des Ansiedlungsprozesses bei beiden Arten, die sich im Zahlenverhältnis Larvenangebot zu Anzahl der ersten Bodenstadien widerspiegelt, kann ein Hinweis auf eine überwiegend passive Ansiedlung der Larven am Boden sein. Der Ort der Hauptansiedlung von C. edule wurde durch den Transekt nicht erfaßt. Die Station B2 war zwar durch ein Herzmuschelfeld charakterisiert, dieses war aber nach zwei Eiswintern nahezu vollständig eliminiert. Der Abundanz der planktischen Larven zufolge war der Hauptansiedlungszeitraum ebenfalls Ende Mai/Anfang Juni. Zu dieser Zeit wurden nur vereinzelt frühe Bodenstadien an den Stationen B1 und B2 gefunden, keine an den Stationen B3 und B4. Während die frühen postlarvalen Stadien von M. arenaria überwiegend am Ort der Ansiedlung blieben, verbreiteten sich die von M. balthica bis in den oberen Bereich des Untersuchungsgebietes (B3-B5). Analog zu der Besiedlung dieser Gebiete durch postlarvale M. balthica wurde die im Verlauf des Untersuchungsjahres stattfindende Kolonisierung der Station B1 durch C. edule ebenfalls postlarvalem Transport zugeschrieben. Demzufolge spielt bei beiden Muschelarten postlarvaler Transport eine wichtige Rolle bei der Besiedlung von Habitaten. Planktonfänge innerhalb der bodennahen Wasserschicht bestätigten, daß im Untersuchungsgebiet M. balthica die am stärksten verdriftende Muschelart war, gefolgt von C. edule. Mortalität, Wachstum, mittlere Biomasse, Produktion und P/B-Verhältnis wurden für M. balthica an den Stationen B1, B3 und B4 sowie für M. arenaria an der Station B1 bestimmt. Wachstum und damit auch Produktion beider Arten erwiesen sich hier - wie an den höher gelegen Stationen (nur M. balthica) - als durch größenselektiven Feinddruck beeinflußt. Der Effekt postlarvalen Transports auf Wachstum wird diskutiert. Übergreifend über die auf Artebene diskutierten Ergebnisse wird die Bedeutung der Dispersion postlarvaler Stadien und die Wirkung epibenthischen Feinddrucks im Wattenmeer erörtert. Der Vergleich postlarvalen Transportes mit der Dispersion planktischer Larvenstadien, der Dispersion von Meiofauna und der Mobilität adulter Stadien der Makrofauna verdeutlicht, daß es sich hierbei um eine Strategie handeln kann, innerhalb eines unvorhersagbaren Biotops freiwerdende Ressourcen zu nutzen und dadurch Konkurrenz zu vermeiden. Es wird die Hypothese aufgestellt, daß Initialansiedlung und Immigration einerseits sowie Feinddruck und Emigration andererseits einen Regelkreis darstellen, der in verschiedenen Teilbereichen des Watts mit unterschiedlicher Geschwindigkeit abläuft.

Pollen and spores record, and paleotemperature reconstructions for mid-Holocene sediments of the Northwest Pacific

Sannai-Maruyama is one of the most famous and best-researched mid-Holocene (mid-Jomon) archaeological sites in Japan, because of a large community of people for a long period. Archaeological studies have shown that the Jomon people inhabited the Sannai-Maruyama site from 5.9-4.2 +/- 0.1 cal. kyr B.P. However, a continuous record of the terrestrial and marine environments around the site has not been available. Core KT05-7 PC-02, was recovered from Mutsu Bay, only 20 km from the site, for the reconstruction of high-resolution time series of environmental records, including sea surface temperature (SST). C37 alkenone SSTs showed clear fluctuations, with four periods of high (8.4-7.9, 7.0-5.9, 5.1-4.1, and 2.3-1.4 cal. kyr B.P.) and four of low (-8.4, 7.9-7.0, 5.9-5.1, and 4.1-2.3 cal. kyr B.P.) SST. Thus, each SST cycle lasted 1.0-2.0 kyr, and the amplitude of fluctuation was about 1.5-2.0 °C. Total organic carbon (TOC) and C37 alkenone contents, and the TOC/total nitrogen ratio indicate that marine biogenic production was low before 7.0 cal. kyr B.P., but was clearly increased between 5.9 and 4.0 cal. kyr B.P., because of stronger vertical mixing. During the period when the community at the site prospered (between 5.9 and 4.2 +/- 0.1 cal. kyr B.P.), the terrestrial climate was relatively warm. The high relative abundance of pollen of both Castanea and Quercus subgen. Cyclobalanopsis supports the interpretation that the local climate was optimal for human habitation. Between 5.9 and 5.1 cal. kyr B.P., in spite of warm terrestrial climates, the C37 alkenone SST was low; this apparent discrepancy may be attributed to the water column structure in the Tsugaru Strait, which differed from the modern condition. The evidence suggests that at about 5.9 cal. kyr B.P, high productivity of marine resources such as fish and shellfish and a warm terrestrial climate led to the establishment of a human community at the Sannai-Maruyama site. Then, at about 4.1 +/- 0.1 cal. kyr B.P., abrupt marine and terrestrial cooling, indicated by a decrease of about 2 °C in the C37 alkenone SST and an increase in pollen of taxa of cooler climates, led to a reduced terrestrial food supply, causing the people to abandon the site. The timing of the abandonment is consistent with the timing (around 4.0-4.3 cal. kyr B.P.) of the decline of civilizations in north Mesopotamia and along the Yangtze River. These findings suggest that a temperature rise of ~2 °C in this century as a result of global warming could have a great impact on the human community and especially on agriculture, despite the advances of contemporary society.

Radiocarbon ages and average physical properties of sediment cores obtained during Icebreaker Oden cruise OSO0910

To date, understanding of ice sheet retreat within Pine Island Bay (PIB) following the Last Glacial Maximum (LGM) was based on seven radiocarbon dates and only fragmentary seafloor geomorphic evidence. During the austral summer 2009-2010, restricted sea ice cover allowed for the collection of 27 sediment cores from the outer PIB trough region. Combining these cores with data from prior cruises, over 133 cores have been used to conduct a detailed sedimentological facies analysis. These results, augmented by 23 new radiocarbon dates, are used to reconstruct the post-LGM deglacial history of PIB. Our results record a clear retreat stratigraphy in PIB composed of, from top to base; terrigenous sandy silt (distal glacimarine), pebbly sandy mud (ice-proximal glacimarine), and till. Initial retreat from the outer-continental shelf began shortly after the LGM and before 16.4 k cal yr BP, as a likely response to rising sea level. Bedforms in outer PIB document episodic retreat in the form of back-stepping grounding zone wedges and are associated with proximal glacimarine sediments. A sub-ice shelf facies is observed in central PIB and spans ~12.3-10.6 k cal yr BP. It is possible that widespread impingement of warm water onto the continental shelf caused an abrupt and widespread change from sub-ice shelf sedimentation to distal glacimarine sedimentation dominated by widespread dispersal of terrigenous silt between 7.8 and 7.0 k cal yr BP. The final phase of retreat ended before ~1.3 k cal yr BP, when the grounding line migrated to a location near the current ice margin.

Iberian Margin 420 kyr geochemical and color variations record

Here we report 420 kyr long records of sediment geochemical and color variations from the southwestern Iberian Margin. We synchronized the Iberian Margin sediment record to Antarctic ice cores and speleothem records on millennial time scales and investigated the phase responses relative to orbital forcing of multiple proxy records available from these cores. Iberian Margin sediments contain strong precession power. Sediment "redness" (a* and 570-560 nm) and the ratio of long-chain alcohols to n-alkanes (C26OH/(C26OH + C29)) are highly coherent and in-phase with precession. Redder layers and more oxidizing conditions (low alcohol ratio) occur near precession minima (summer insolation maxima). We suggest these proxies respond rapidly to low-latitude insolation forcing by wind-driven processes (e.g., dust transport, upwelling, precipitation). Most Iberian Margin sediment parameters lag obliquity maxima by 7-8 ka, indicating a consistent linear response to insolation forcing at obliquity frequencies driven mainly by high-latitude processes. Although the lengths of the time series are short (420 ka) for detecting 100 kyr eccentricity cycles, the phase relationships support those obtained by Shackleton []. Antarctic temperature and the Iberian Margin alcohol ratios (C26OH/(C26OH + C29)) lead eccentricity maxima by 6 kyr, with lower ratios (increased oxygenation) occurring at eccentricity maxima. CO2, CH4, and Iberian SST are nearly in phase with eccentricity, and minimum ice volume (as inferred from Pacific d18Oseawater) lags eccentricity maxima by 10 kyr. The phase relationships derived in this study continue to support a potential role of the Earth's carbon cycle in contributing to the 100 kyr cycle.

Porphyrin geochemistry of DSDP Hole 71-511

Late Jurassic-early Cretaceous black shales and an overlying sequence of Albian-Campanian zeolitic claystones from the Falkland Plateau (DSDP/IPOD Leg 71, Site 511) were analyzed for tetrapyrrole pigment type and abundance. The "black shale" sequence was found to be rich in DPEP-series dominated free-base, nickel (Ni) and, to a lesser extent, vanadyl (V = 0) porphyrins. A low level of organic maturity (i.e. precatagenesis) is indicated for these strata as nickel chelation by free-base porphyrins is only 50-75% complete, proceeding down-hole to 627 meters sub-bottom. Electronic and mass spectral data reveal that the proposed benzo-DPEP (BD) and tetrahydrobenzo-DPEP (THBD) series are present in the free-base and Ni species, as well as the more usual occurrence in V = 0 porphyrin arrays. Highly reducing conditions are suggested by an abundance of the PAH perylene, substantial amounts of the THBD/BD series and a redox equilibrium between free-base DPEP and 7,8-dihydro-DPEP series, which exist in a 7:1 molar ratio. The Albian-Campanian claystone strata were found to be tetrapyrrolepoor, and those pigments present were typed as Cu/Ni highly dealkylated (C26 max.) etioporphyrins, thought to be derived via redeposition and oxidation of terrestrial organic matter (OM). Results from the present study are correlated to our past analyses of Jurassic-Cretaceous sediments from Atlantic margins in an effort to relate tetrapyrrole quality and quantity to basin evolution and OM sources in the proto-Atlantic.

Coccoliths and carbonate grain-size compositions in sediment samples from the Mid-Atlantic Ridge

This study presents a differentiated carbonate budget for marine surface sediments from the Mid-Atlantic Ridge of the South Atlantic, with results based on carbonate grain-size composition. Upon separation into sand, silt, and clay sub-fractions, the silt grain-size distribution was measured using a SediGraph 5100. We found regionally characteristic grain-size distributions with an overall minimum at 8 µm equivalent spherical diameter (ESD). SEM observations reveal that the coarse particles (>8 µm ESD) are attributed to planktic foraminifers and their fragments, and the fine particles (<8 µm ESD) to coccoliths. On the basis of this division, the regional variation of the contribution of foraminifers and coccoliths to the carbonate budget of the sediments are calculated. Foraminifer carbonate dominates the sediments in mesotropic regions whereas coccoliths contribute most carbonate in oligotrophic regions. The grain size of the coccolith share is constant over water depth, indicating a lower susceptibility for carbonate dissolution compared to foraminifers. Finally, the characteristic grain-size distribution in fine silt (<8 µm ESD) is set into context with the coccolith assemblage counted and biometrically measured using a SEM. The coccoliths present in the silt fraction are predominantly large species (length > 4 µm). Smaller species (length < 4 µm) belong to the clay fraction (<2 µm ESD). The average length of most frequent coccolith species is connected to prominent peaks in grain-size distributions (ESD) with a shape factor. The area below Gaussian distributions fitted to these peaks is suggested as a way to quantitatively estimate the carbonate contribution of single coccolith species more precisely compared to conventional volume estimates. The quantitative division of carbonate into the fraction produced by coccoliths and that secreted by foraminifers enables a more precise estimate for source/sink relations of consumed and released CO2 in the carbon cycle. The allocation of coccolith length and grain size (ESD) suggests size windows for the separation or accumulation of distinct coccolith species in investigations that depend on non to slightly-mixed signals (e.g., isotopic studies).

Seawater carbonate chemistry, mass, size and regenerative capacity of Luidia clathrata during experiments, 2011

The present study examines sublethal effects of near-future (year 2100) ocean acidification (OA) on regenerative capacity, biochemical composition, and behavior of the sea star Luidia clathrata, a predominant predator in sub-tropical soft-bottom habitats. Two groups of sea stars, each with two arms excised, were maintained on a formulated diet in seawater bubbled with air alone (pH 8.2, approximating a pCO2 of 380 µatm) or with a controlled mixture of air/C02 (pH 7.8, approximating a pCO2 of 780 µatm). Arm length, total body wet weight, and righting responses were measured weekly. After 97 days, a period of time sufficient for 80% arm regeneration, pyloric caecal indices, and protein, carbohydrate, lipid, and ash levels were determined for body wall and pyloric caecal tissues of intact and regenerating arms of individuals held in both seawater pH treatments. The present study indicates that predicted near-term levels of ocean acidification (seawater pH 7.8) do not significantly impact whole animal growth, arm regeneration rates, biochemical composition, or righting behavior in this common soft bottom sea star.

CO2-driven ocean acidification alters and weakens integrity of the calcareous tubes produced by the serpulid tubeworm, Hydroides elegans

As a consequence of anthropogenic CO2-driven ocean acidification (OA), coastal waters are becoming increasingly challenging for calcifiers due to reductions in saturation states of calcium carbonate (CaCO3) minerals. The response of calcification rate is one of the most frequently investigated symptoms of OA. However, OA may also result in poor quality calcareous products through impaired calcification processes despite there being no observed change in calcification rate. The mineralogy and ultrastructure of the calcareous products under OA conditions may be altered, resulting in changes to the mechanical properties of calcified structures. Here, the warm water biofouling tubeworm, Hydroides elegans, was reared from larva to early juvenile stage at the aragonite saturation state (Omega A) for the current pCO2 level (ambient) and those predicted for the years 2050, 2100 and 2300. Composition, ultrastructure and mechanical strength of the calcareous tubes produced by those early juvenile tubeworms were examined using X-ray diffraction (XRD), Fourier transform infrared spectroscopy (FT-IR), scanning electron microscopy (SEM) and nanoindentation. Juvenile tubes were composed primarily of the highly soluble CaCO3 mineral form, aragonite. Tubes produced in seawater with aragonite saturation states near or below one had significantly higher proportions of the crystalline precursor, amorphous calcium carbonate (ACC) and the calcite/aragonite ratio dramatically increased. These alterations in tube mineralogy resulted in a holistic deterioration of the tube hardness and elasticity. Thus, in conditions where Omega A is near or below one, the aragonite-producing juvenile tubeworms may no longer be able to maintain the integrity of their calcification products, and may result in reduced survivorship due to the weakened tube protection.

Temperature Dependent Effects of Elevated CO2 on Shell Composition and Mechanical Properties of Hydroides elegans: Insights from a Multiple Stressor Experiment

The majority of marine benthic invertebrates protect themselves from predators by producing calcareous tubes or shells that have remarkable mechanical strength. An elevation of CO2 or a decrease in pH in the environment can reduce intracellular pH at the site of calcification and thus interfere with animal's ability to accrete CaCO3. In nature, decreased pH in combination with stressors associated with climate change may result in the animal producing severely damaged and mechanically weak tubes. This study investigated how the interaction of environmental drivers affects production of calcareous tubes by the serpulid tubeworm, Hydroides elegans. In a factorial manipulative experiment, we analyzed the effects of pH (8.1 and 7.8), salinity (34 and 27), and temperature (23°C and 29°C) on the biomineral composition, ultrastructure and mechanical properties of the tubes. At an elevated temperature of 29°C, the tube calcite/aragonite ratio and Mg/Ca ratio were both increased, the Sr/Ca ratio was decreased, and the amorphous CaCO3 content was reduced. Notably, at elevated temperature with decreased pH and reduced salinity, the constructed tubes had a more compact ultrastructure with enhanced hardness and elasticity compared to decreased pH at ambient temperature. Thus, elevated temperature rescued the decreased pH-induced tube impairments. This indicates that tubeworms are likely to thrive in early subtropical summer climate. In the context of climate change, tubeworms could be resilient to the projected near-future decreased pH or salinity as long as surface seawater temperature rise at least by 4°C.

Multiple stressor effects of near-future elevated seawater temperature and decreased pH on righting and escape behaviors of two common Antarctic gastropods

Warming seawater temperatures and ocean acidification on the coastal western Antarctic Peninsula pose unique challenges to stenothermal marine invertebrates. The present study examines prospective sub-lethal effects of elevated temperature, pCO2, and resultant decrease in seawater pH, on righting behavior and maximal escape speeds for two common gastropods, the limpet Nacella concinna (Strebel) and mesogastropod snail Margarella antarctica (Lamy). Replicate individuals held in individual containers were exposed to four combinations of seawater temperature (1.5 °C-current average, 3.5 °C-projected average by 2100) and pH (pH 8.0-current average, pH 7.8-projected average by 2100 as a result of elevated pCO2 levels) for a period of 6 weeks. Following this chronic exposure, righting behavior, determined for the limpets as proportion to right over 24 h and for snails as time to right, as well as maximum escape speed following contact with a sea star predator were measured. We found no significant differences in proportions of limpets displaying the capacity to right among the four temperature-pH treatments. However, there was a significant temperature-pH interaction effect for mean righting times in snails, indicating that the effect of pH on the time to right is dependent on temperature. We found no significant effects of temperature or pH on mean maximal escape speed in limpets. Additionally, we observed a significant temperature-pH interaction effect for mean maximal escape speed in snails. These interactive effects make it difficult to make clear predictions about how these environmental factors may impact behavioral responses.

Biogeochemical processes in anoxic sediment from Lake Plußsee, Germany

In anoxic environments, volatile methylated sulfides like methanethiol (MT) and dimethyl sulfide (DMS) link the pools of inorganic and organic carbon with the sulfur cycle. However, direct formation of methylated sulfides from reduction of dissolved inorganic carbon has previously not been demonstrated. When studying the effect of temperature on hydrogenotrophic microbial activity, we observed formation of DMS in anoxic sediment of Lake Plußsee at 55 °C. Subsequent experiments strongly suggested that the formation of DMS involves fixation of bicarbonate via a reductive pathway in analogy to methanogenesis and engages methylation of MT. DMS formation was enhanced by addition of bicarbonate and further increased when both bicarbonate and H2 were supplemented. Inhibition of DMS formation by 2-bromoethanesulfonate points to the involvement of methanogens. Compared to the accumulation of DMS, MT showed the opposite trend but there was no apparent 1:1 stoichiometric ratio between both compounds. Both DMS and MT had negative d13C values of -62 per mil and -55 per mil, respectively. Labeling with NaH**13CO3 showed more rapid incorporation of bicarbonate into DMS than into MT. The stable carbon isotopic evidence implies that bicarbonate was fixed via a reductive pathway of methanogenesis, and the generated methyl coenzyme M became the methyl donor for MT methylation. Neither DMS nor MT accumulation were stimulated by addition of the methyl-group donors methanol and syringic acid or by the methyl-group acceptor hydrogen sulphide. The source of MT was further investigated in a H2**35S labeling experiment, which demonstrated a microbially-mediated process of hydrogen sulfide methylation to MT that accounted for only <10% of the accumulation rates of DMS. Therefore, the major source of the 13C-depleted MT was neither bicarbonate nor methoxylated aromatic compounds. Other possibilities for isotopically depleted MT, such as other organic precursors like methionine, are discussed. This DMS-forming pathway may be relevant for anoxic environments such as hydrothermally influenced sediments and fluids and sulfate-methane transition zones in marine sediments.