Isotopic composition of ground ice, ebullition gases and thermokarst lake water, North America

Thermokarst lakes are thought to have been an important source of methane (CH4) during the last deglaciation when atmospheric CH4 concentrations increased rapidly. Here we demonstrate that meltwater from permafrost ice serves as an H source to CH4 production in thermokarst lakes, allowing for region-specific reconstructions of dD-CH4 emissions from Siberian and North American lakes. dD CH4 reflects regionally varying dD values of precipitation incorporated into ground ice at the time of its formation. Late Pleistocene-aged permafrost ground ice was the dominant H source to CH4 production in primary thermokarst lakes, whereas Holocene-aged permafrost ground ice contributed H to CH4 production in later generation lakes. We found that Alaskan thermokarst lake dD-CH4 was higher (-334 ± 17 per mil) than Siberian lake dD-CH4 (-381 ± 18 per mil). Weighted mean dD CH4 values for Beringian lakes ranged from -385 per mil to -382 per mil over the deglacial period. Bottom-up estimates suggest that Beringian thermokarst lakes contributed 15 ± 4 Tg CH4 /yr to the atmosphere during the Younger Dryas and 25 ± 5 Tg CH4 /yr during the Preboreal period. These estimates are supported by independent, top-down isotope mass balance calculations based on ice core dD-CH4 and d13C-CH4 records. Both approaches suggest that thermokarst lakes and boreal wetlands together were important sources of deglacial CH4.

Abundance of mesozooplankton in the western part of the Black Sea during the 2001 National Monitoring Programme of the Bulgarian Institute of Oceanography

The dataset is based on samples collected in the autumn of 2001 in the Western Black Sea in front of Bulgaria coast. The whole dataset is composed of 42 samples (from 19 stations of National Monitoring Grid) with data of mesozooplankton species composition abundance and biomass. Samples were collected in the layers 0-10, 0-20, 0-50, 10-25, 25-50, 50-100 and from bottom up to the surface at depths depending on water column stratification and the thermocline depth. Zooplankton samples were collected with vertical closing Juday net,diameter - 36cm, mesh size 150 µm. Tows were performed from surface down to bottom meters depths in discrete layers. Samples were preserved by a 4% formaldehyde sea water buffered solution. Sampling volume was estimated by multiplying the mouth area with the wire length. Mesozooplankton abundance: The collected material was analysed using the method of Domov (1959). Samples were brought to volume of 25-30 ml depending upon zooplankton density and mixed intensively until all organisms were distributed randomly in the sample volume. After that 5 ml of sample was taken and poured in the counting chamber which is a rectangle form for taxomomic identification and count. Large (> 1 mm body length) and not abundant species were calculated in whole sample. Counting and measuring of organisms were made in the Dimov chamber under the stereomicroscope to the lowest taxon possible. Taxonomic identification was done at the Institute of Oceanology by Kremena Stefanova using the relevant taxonomic literature (Mordukhay-Boltovskoy, F.D. (Ed.). 1968, 1969,1972). Taxon-specific abundance: The collected material was analysed using the method of Domov (1959). Samples were brought to volume of 25-30 ml depending upon zooplankton density and mixed intensively until all organisms were distributed randomly in the sample volume. After that 5 ml of sample was taken and poured in the counting chamber which is a rectangle form for taxomomic identification and count. Copepods and Cladoceras were identified and enumerated; the other mesozooplankters were identified and enumerated at higher taxonomic level (commonly named as mesozooplankton groups). Large (> 1 mm body length) and not abundant species were calculated in whole sample. Counting and measuring of organisms were made in the Dimov chamber under the stereomicroscope to the lowest taxon possible. Taxonomic identification was done at the Institute of Oceanology by Kremena Stefanova using the relevant taxonomic literature (Mordukhay-Boltovskoy, F.D. (Ed.). 1968, 1969,1972).

Biomass of mesozooplankton in the western part of the Black Sea in 1992 during the Cooperative Marine Science Program for the Black Sea (CoMSBlack)

The "CoMSBlack92" dataset is based on samples collected in the summer of 1992 along the Bulgarian coast including coastal and open sea areas. The whole dataset is composed of 79 samples (28 stations) with data of zooplankton species composition, abundance and biomass. Sampling for zooplankton was performed from bottom up to the surface at standard depths depending on water column stratification and the thermocline depth. Zooplankton samples were collected with vertical closing Juday net,diameter - 36cm, mesh size 150 ?m. Tows were performed from surface down to bottom meters depths in discrete layers. Samples were preserved by a 4% formaldehyde sea water buffered solution. Sampling volume was estimated by multiplying the mouth area with the wire length. Sampling volume was estimated by multiplying the mouth area with the wire length. The collected material was analysed using the method of Domov (1959). Samples were brought to volume of 25-30 ml depending upon zooplankton density and mixed intensively until all organisms were distributed randomly in the sample volume. After that 5 ml of sample was taken and poured in the counting chamber which is a rectangle form for taxomomic identification and count. Large (> 1 mm body length) and not abundant species were calculated in whole sample. Counting and measuring of organisms were made in the Dimov chamber under the stereomicroscope to the lowest taxon possible. Taxonomic identification was done at the Institute of Oceanology by Asen Konsulov using the relevant taxonomic literature (Mordukhay-Boltovskoy, F.D. (Ed.). 1968, 1969,1972 ). The biomass was estimated as wet weight by Petipa, 1959 (based on species specific wet weight). Wet weight values were transformed to dry weight using the equation DW=0.16*WW as suggested by Vinogradov & Shushkina, 1987. Copepods and Cladoceras were identified and enumerated; the other mesozooplankters were identified and enumerated at higher taxonomic level (commonly named as mesozooplankton groups). Large (> 1 mm body length) and not abundant species were calculated in whole sample. The biomass was estimated as wet weight by Petipa, 1959 ussing standard average weight of each species in mg/m**3.

Biomass of mesozooplankton in the western part of the Black Sea in summer 1991 during the NATO Programme Hydroblack

The "Hydroblack91" dataset is based on samples collected in the summer of 1991 and covers part of North-Western in front of Romanian coast and Western Black Sea (Bulgarian coasts) (between 43°30' - 42°10' N latitude and 28°40'- 31°45' E longitude). Mesozooplankton sampling was undertaken at 20 stations. The whole dataset is composed of 72 samples with data of zooplankton species composition, abundance and biomass. Samples were collected in discrete layers 0-10, 0-20, 0-50, 10-25, 25-50, 50-100 and from bottom up to the surface at depths depending on water column stratification and the thermocline depth. Zooplankton samples were collected with vertical closing Juday net,diameter - 36cm, mesh size 150 µm. Tows were performed from surface down to bottom meters depths in discrete layers. Samples were preserved by a 4% formaldehyde sea water buffered solution. Sampling volume was estimated by multiplying the mouth area with the wire length. Mesozooplankton abundance: The collected materia was analysed using the method of Domov (1959). Samples were brought to volume of 25-30 ml depending upon zooplankton density and mixed intensively until all organisms were distributed randomly in the sample volume. After that 5 ml of sample was taken and poured in the counting chamber which is a rectangle form for taxomomic identification and count. Large (> 1 mm body length) and not abundant species were calculated in whole sample. Counting and measuring of organisms were made in the Dimov chamber under the stereomicroscope to the lowest taxon possible. Taxonomic identification was done at the Institute of Oceanology by Asen Konsulov using the relevant taxonomic literature (Mordukhay-Boltovskoy, F.D. (Ed.). 1968, 1969,1972). The biomass was estimated as wet weight by Petipa, 1959 (based on species specific wet weight). Wet weight values were transformed to dry weight using the equation DW=0.16*WW as suggested by Vinogradov & Shushkina, 1987. Taxon-specific abundance: The collected material was analysed using the method of Domov (1959). Samples were brought to volume of 25-30 ml depending upon zooplankton density and mixed intensively until all organisms were distributed randomly in the sample volume. After that 5 ml of sample was taken and poured in the counting chamber which is a rectangle form for taxomomic identification and count. Copepods and Cladoceras were identified and enumerated; the other mesozooplankters were identified and enumerated at higher taxonomic level (commonly named as mesozooplankton groups). Large (> 1 mm body length) and not abundant species were calculated in whole sample. Counting and measuring of organisms were made in the Dimov chamber under the stereomicroscope to the lowest taxon possible. Taxonomic identification was done at the Institute of Oceanology by Asen Konsulov using the relevant taxonomic literature (Mordukhay-Boltovskoy, F.D. (Ed.). 1968, 1969,1972). The biomass was estimated as wet weight by Petipa, 1959 ussing standard average weight of each species in mg/m3. WW were converted to DW by equation DW=0.16*WW (Vinogradov ME, Sushkina EA, 1987).

Limits of the Southern Ocean

The knowledge about processes concerning perception and understanding is of paramount importance for designing means of communication like maps and charts. This is especially the case, if one does not want to lose sight of the map-user and if map-design is to be orientated along the map-users needs and preferences in order to improve the cartographic product's usability. A scientific approach to visualization can help to achieve useable results. The insights achieved by such an approach can lead to modes of visualization that are superior to those, which have seemingly proved their value in praxis - so-called "bestpractices" -, concerning their utility and efficiency. This thesis shows this by using the example of visualizing the limits of bodies of waters in the Southern Ocean. After making some introductorily remarks on the chosen mode of problem-solution in chapter one, which simultaneously illustrate the flow of work while working on the problem, in chapter two the relevant information concerning the drawing of limits in the Southern Ocean is outlined. Chapter 3 builds the theoretical framework, which is a multidisciplinary approach to representation. This theoretical framework is based on "How Maps Work" by the American Cartographer MacEachren (1995/2004). His "scientific approach to visualization" is amended and adjusted by the knowledge gained from recent findings of the social sciences where necessary. So, the approach suggested in this thesis represents a synergy of psychology, sociology, semiotics, linguistics, communication theory and cartography. It follows the tradition of interdisciplinary research getting over the boundaries of a single scientific subject. The achieved holistic approach can help to improve the usability of cartographic products. It illustrates on the one hand those processes taking place while perceiving and recognizing cartographic information - so-called bottom-up-processes. On the other hand it illuminates the processes which happen during understanding this information in so-called top-down-processes. Bottom-up- and top-down-processes are interdependent and inseparably interrelated and therefore cannot be understood without each other. Regarding aspects of usability the approach suggested in this thesis strongly focuses on the map-user. This is the reason why the phenomenon of communication gains more weight than in MacEachren's map-centered approach. Because of this, in chapter 4 a holistic approach to communication is developed. This approach makes clear that only the map-user can evaluate the usability of a cartographic product. Only if he can extract the information relevant for him from the cartographical product, it is really useable. The concept of communication is well suited to conceive that. In case of the visualization of limits of bodies of water in the Southern Ocean, which is not complex enough to illustrate all results of the theoretical considerations, it is suggested to visualize the limits with red lines. This suggestion deviates from the commonly used mode of visualization. So, this thesis shows how theory is able to ameliorate praxis. Chapter 5 leads back to the task of fixing limits of the bodies of water in the area of concern. A convention by the International Hydrographic Organization (IHO) states that those limits should be drawn by using meridians, parallels, rhumb lines and bathymetric data. Based on the available bathymetric data both a representation and a process model are calculated, which should support the drawing of the limits. The quality of both models, which depends on the quality of the bathymetric data at hand, leads to the decision that the representation model is better suited to support the drawing of limits.

Seawater carbonate chemistry, thickness and carbonate elemental composition of the test of juvenile sea urchins in a laboratory experiment

Continuous anthropogenic CO2 emissions to the atmosphere and uptake by the oceans will cause a reduction of seawater pH and saturation state (Omega) of CaCO3 minerals from which marine calcifiers build their shells and skeletons. Sea urchins use the most soluble form of calcium carbonate, high-magnesium calcite, to build their skeleton, spines and grazing apparatus. In order to highlight the effects of increased pCO2 on the test thickness and carbonate elemental composition of juvenile sea urchins and potential differences in their responses linked to the diet, we performed a laboratory experiment on juvenile Paracentrotus lividus, grazing on calcifying (Corallina elongata) and non-calcifying (Cystoseira amentacea, Dictyota dichotoma) macroalgae, under different pH (corresponding to pCO2 values of 390, 550, 750 and 1000 µatm). Results highlighted the importance of the diet in determining sea urchin size irrespectively of the pCO2 level, and the relevance of macroalgal diet in modulating urchin Mg/Ca ratio. The present study provides relevant clues both in terms of the mechanism of mineral incorporation and in terms of bottom-up processes (algal diet) affecting top-down ones (fish predation) in rocky subtidal communities

(Table A1) Annual biomass of plants and animals recorded on Bylot Island between 1993-2009

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator--rey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.

Biological and environmental time series obtained from Point B, Bay of Villefranche

Ecological succession provides a widely accepted description of seasonal changes in phytoplankton and mesozooplankton assemblages in the natural environment, but concurrent changes in smaller (i.e. microbes) and larger (i.e. macroplankton) organisms are not included in the model because plankton ranging from bacteria to jellies are seldom sampled and analyzed simultaneously. Here we studied, for the first time in the aquatic literature, the succession of marine plankton in the whole-plankton assemblage that spanned 5 orders of magnitude in size from microbes to macroplankton predators (not including fish or fish larvae, for which no consistent data were available). Samples were collected in the northwestern Mediterranean Sea (Bay of Villefranche) weekly during 10 months. Simultaneously collected samples were analyzed by flow cytometry, inverse microscopy, FlowCam, and ZooScan. The whole-plankton assemblage underwent sharp reorganizations that corresponded to bottom-up events of vertical mixing in the water-column, and its development was top-down controlled by large gelatinous filter feeders and predators. Based on the results provided by our novel whole-plankton assemblage approach, we propose a new comprehensive conceptual model of the annual plankton succession (i.e. whole plankton model) characterized by both stepwise stacking of four broad trophic communities from early spring through summer, which is a new concept, and progressive replacement of ecological plankton categories within the different trophic communities, as recognised traditionally.

Table 1: Station list with macrobenthos biomass results from the ocean floor

Among the Siberian shelf seas the Kara Sea is most strongly influenced by riverine runoff with nearly 1500 km fresh water discharge per year. This fresh water, discharged mainly by Ob and Yenisei, contains about 3.1 * 106 and 4.6 * 106 tons of total organic carbon per year, respectively (Gordeev et al. 1996). Little is known about the relevance of this organic material for biological communities, neither for the Kara Sea nor for the adjacent deep basins of the central Arctic Ocean. Aiming at elucidating the fate of fluvial matter transported from the rivers via estuaries into the central Arctic Ocean and the relative importance of marine organic matter being produced such information is crucial. Here we present calculations on the organic carbon demand of the Kara Sea macrozoobenthos based on measured biomass (total wet weight [ww] per 0.25 m ) from quantitative box corer samples and empirical relationships between biomass, annual production, annual respiration, and carbon remineralisation. This bottom-up approach may serve as a first estimate of the carbon remineralization potential of a given zoobenthos community (or area) as long as no data on in situ respiration rates are available. Our data basis comprises 54 stations sampled in summer seasons 1997, 1999 and 2000 in the Kara Sea at water depths between 10 and 68 m. The geographical area represented by stations analysed covers roughly 178 000 km**2, which is about one fifth of the total Kara Sea area. In this area, 290 species of invertebrate macrozoobenthos were identified with polychaeta, Crustacea, mollusca and echinodermata being the most abundant. For all stations analysed, mean biomass values ranged between 4.3 and 778.1 g ww/m**2 with organic carbon demands between 3.5 and 43.2 mg C/m**2/d. For the area of 178 000 km2 a preliminary total consumption of 1.4 * 10**6t Corg/y (equivalent to 21.5 mg C/m**2/d) was calculated for the macrozoobenthos. An extrapolation of our data would lead to an annual carbon demand of about 5-7 * 106 t for the whole Kara Sea macrozoobenthos (or 15.5-21.7 mg C/m2/d).

Ocean acidification affects the phyto-zoo plankton trophic transfer efficiency

The critical role played by copepods in ocean ecology and biogeochemistry warrants an understanding of how these animals may respond to ocean acidification (OA). Whilst an appreciation of the potential direct effects of OA, due to elevated pCO2, on copepods is improving, little is known about the indirect impacts acting via bottom-up(food quality) effects. We assessed, for the first time, the chronic effects of direct and/or indirect exposures to elevated pCO2 on the behaviour, vital rates, chemical and biochemical stoichiometry of the calanoid copepod Acartia tonsa. Bottom-up effects of elevated pCO2 caused species-specific biochemical changes to the phytoplanktonic feed, which adversely affected copepod population structure and decreased recruitment by 30 %. The direct impact of elevated pCO2 caused gender-specific respiratory responses in A.tonsa adults, stimulating an enhanced respiration rate in males (> 2-fold), and a suppressed respiratory response in females when coupled with indirect elevated pCO2 exposures. Under the combined indirect+direct exposure, carbon trophic transfer efficiency from phytoplankton-to-zooplankton declined to < 50 % of control populations, with a commensurate decrease in recruitment. For the first time an explicit role was demonstrated for biochemical stoichiometry in shaping copepod trophic dynamics. The altered biochemical composition of the CO2-exposed prey affected the biochemical stoichiometry of the copepods, which could have ramifications for production of higher tropic levels, notably fisheries. Our work indicates that the control of phytoplankton and the support of higher trophic levels involving copepods have clear potential to be adversely affected under future OA scenarios.

Faecal pellet production of copepoda collected in water depth up to 30 meters in the eastern Mediterranean Sea in Oktober 2008 during SES_GR2

Mesozooplankton is collected by vertical tows within the Black sea water body mass layer in the NE Aegean, using a WP-2 200 µm net equipped with a large non-filtering cod-end (10 l). Macrozooplankton organisms are removed using a 2000 µm net. A few unsorted animals (approximately 100) are placed inside several glass beaker of 250 ml filled with GF/F or 0.2 µm Nucleopore filtered seawater and with a 100 µm net placed 1 cm above the beaker bottom. Beakers are then placed in an incubator at natural light and maintaining the in situ temperature. After 1 hour pellets are separated from animals and placed in separated flasks and preserved with formalin. Pellets are counted and measured using an inverted microscope. Animals are scanned and counted using an image analysis system. Carbon- Specific faecal pellet production is calculated from a) faecal pellet production, b) individual carbon: Animals are scanned and their body area is measured using an image analysis system. Body volume is then calculated as an ellipsoid using the major and minor axis of an ellipse of same area as the body. Individual carbon is calculated from a carbon- total body volume of organisms (relationship obtained for the Mediterranean Sea by Alcaraz et al. (2003) divided by the total number of individuals scanned and c) faecal pellet carbon: Faecal pellet length and width is measured using an inverted microscope. Faecal pellet volume is calculated from length and width assuming cylindrical shape. Conversion of faecal pellet volume to carbon is done using values obtained in the Mediterranean from: a) faecal pellet density 1,29 g cm**3 (or pg µm**3) from Komar et al. (1981); b) faecal pellet DW/WW=0,23 from Elder and Fowler (1977) and c) faecal pellet C%DW=25,5 Marty et al. (1994).

Iron and manganese in bottom sediments and interstitial waters sampled in the White Sea on August 21-30, 2003

Iron and manganese in bottom sediments studied along the sublatitudinal transect from Kandalaksha to Arkhangelsk are characterized by various contents and speciations depending on sedimentation environment, grain size of sediments, and diagenetic processes. The latter include redistribution of reactive forms leading to enrichment in Fe and Mn of surface sediments, formation of films, incrustations, and ferromanganese nodules. Variations in total Fe content (2-8%) are accompanied by changes in concentration of its reactive forms (acid extraction) and concentration of dissolved Fe in interstitial waters (1-14 µM). Variations in Mn content in bottom sediments (0.03-3.7%) and interstitial waters (up to 500 µM) correspond to high diagenetic mobility of this element. Changes in oxidation degree of chemical elements result in redox stratification of sediment strata with maximum concentrations of Fe, Mn, and sulfides. Organic matter of bottom sediments with considerable terrestrial constituent is oxidized by bottom water oxygen mainly at the sediment surface or in anaerobic conditions within the sediment strata. The role of inorganic components in organic matter oxidation changes from surface layer bottom sediments (where manganese oxyhydroxide dominates among oxidants) to deeper layers (where sulfate of interstitial water serves as the main oxidant). Differences in river runoff and hydrodynamics are responsible for geochemical asymmetry of the transect. The deep Kandalaksha Bay serves as a sediment trap for manganese (Mn content in sediments varies within 0.5-0.7%), whereas the sedimentary environment in the Dvina Bay promotes its removal from bottom sediments (Mn 0.05%).

Abundance and biomass of benthic foraminifera in bottom sediments from the Sea of Okhotsk and the Pacific Ocean

A study of distribution of live individuals of benthic foraminifera in sediments of the Sea of Okhotsk and of the Northwestern Basin of the Pacific Ocean shows that they can be present in sediments up to depth of 30 cm and probably can live there for long periods, sometimes forming high concentrations. Living individuals in the subsurface layer often account for more than 50% of total biomass, which varies from 1 to 21 g/m**2 in different morphological structures. The largest biomass values are attained in underwater rises embedded in relatively warm, oxygen-saturated Pacific waters. Minimum total biomass concentrations occur in deep-water depressions where stagnation phenomena are observed. Foraminifera biomass everywhere decreases gradually with increasing depth from the surface of sediments regardless of relief, depth, and nature of sediments.

Chemical composition of bottom sediments from the northern part of the Deryugin Trough, Sea of Okhotsk

Distribution trace element contents in the upper (up to 5 m) Holocene-Upper Pleistocene sediment layer along the northern and southern sublatitudinal profiles in the northern part of the Deryugin Trough is discussed. Mathematical processing of chemical analysis data has been made. On the basis of the cluster analysis 16 elements have been combined into 5 geochemical groups. Two of them (1 - Ag-Mo group and 2 - Mn-Ba-Ni-Sr group) were considered in detail. Analysis of Ag and Mo distributions in the sediments and findings of molybdenite permitted to conclude that enrichment of the sediments by these elements resulted from edaphic washing of magmatic rocks containing Ag-Mo mineralization and probably located on the northwestern slope of the Deryugin trough. The second geochemical group is most likely connected with hydrothermal barite mineralization found in the northeastern part of the Deryugin trough. Considering Mn distribution in bottom sediment cores supply of Mn was pulsating. This allows concluding that during Holocene - Late Pleistocene three cycles of hydrothermal activity occurred in the Deryugin Trough, and the most intensive one was in Holocene.

Manganese contents and accumulation rates in reduced bottom sediments of the Sea of Okhotsk

Manganese contents in reduced sediments and accumulation rates were investigated. Their values in sediments of most of cores are background (0.03-0.07 %).Anomalous concentrations (up to 2.5 %) and accumulation rates (up to 60 mg/cm**2/ka) occur near the known region of hydrothermal barite mineralization in the Derugin Basin. High accumulation rates of Mn (>10 mg/cm**2/ka) also occur in Holocene sediments to south-east from the Derugin Basin. It can be assumed that high Mn contents and accumulation rates occur there due to transportation of Mn-rich water from the Derugin Basin in the near-bottom layer under the lower border of the Sea of Okhotsk Intermediate Water. Intensive Mn accumulation is also typical for the South Okhotsk Basin near the Bussol Strait. Mn accumulation rates of glacial sediments of the second oxygen isotope stage are less significant, which is presumed to be caused by paleoceanological reasons.