Benthic foraminifera across the Cretaceous/Paleogene boundary in the Southern Ocean

The impact of an asteroid at the Cretaceous/Paleogene (K/Pg) boundary triggered dramatic biotic, biogeochemical and sedimentological changes in the oceans that have been intensively studied. Paleo-biogeographical differences in the biotic response to the impact and its environmental consequences, however, have been less well documented. We present a high-resolution analysis of benthic foraminiferal assemblages at Southern Ocean ODP Site 690 (Maud Rise, Weddell Sea, Antarctica). At this high latitude site, late Maastrichtian environmental variability was high, but benthic foraminiferal assemblages were not less diverse than at lower latitudes, in contrast to those of planktic calcifiers. Also in contrast to planktic calcifiers, benthic foraminifera did not suffer significant extinction at the K/Pg boundary, but show transient assemblage changes and decreased diversity. At Site 690, the extinction rate was even lower (~3%) than at other sites. The benthic foraminiferal accumulation rate varied little across the K/Pg boundary, indicating that food supply to the sea floor was affected to a lesser extent than at lower latitude sites. Compared to Maastrichtian assemblages, Danian assemblages have a lower diversity and greater relative abundance of heavily calcified taxa such as Stensioeina beccariiformis and Paralabamina lunata. This change in benthic foraminiferal assemblages could reflect post-extinction proliferation of different photosynthesizers (thus food for the benthos) than those dominant during the Late Cretaceous, therefore changes in the nature rather than in the amount of the organic matter supplied to the seafloor. However, severe extinction of pelagic calcifiers caused carbonate supersaturation in the oceans, thus might have given competitive advantage to species with large, heavily calcified tests. This indirect effect of the K/Pg impact thus may have influenced the deep-sea dwellers, documenting the complexity of the effects of major environmental disturbance.

Response of nannoplankton to major changes in sea-surface temperature at the South Chatham Rise, southeastern New Zealand

A high-resolution history of paleoceanographic changes in the subpolar waters of the southern margin of the Subtropical Convergence Zone during the last 130 kyr, is present in foraminiferal assemblages of DSDP Site 594. The foraminifera indicate that sea-surface temperatures during the Last Interglacial Climax were warmer than today, and that between substage 5d through to the end of isotope stage 2, temperatures were mostly cooler than Holocene temperatures. The paleotemperatures suggest that (1) the Subtropical Convergence was located over the site during substage 5e, later moving further north, then moving southwards to near the site during the Holocene, and (2) the Polar Front was positioned over the Site during glacial stages 6, 4, 2 and possibly parts of stage 3. Several major events are indicated by the nannofloral assemblages during these large changes in sea-surface temperature and associated reorganization of ocean circulation. First, the time-progressive trends between E. huxleyi and medium to large Gephyrocupsa are unique to this site, with E. huxleyi dominating over medium Gephyrocupsa during stages 5c-a, middle part of stage 4 and after the middle point of stage 3. This unusual trend may (at least partly) be caused by the shift of the Polar Front across the site. Second, upwelling flora (E. huxleyi and small placoliths) increase in abundance during stages 1, 3 and 5, suggesting that upwelling or disturbance of water stratification took place during the interglacials. Thirdly, there are no significant differences between the distribution patterns of the various morphotypes of medium to large Gephyrocupsu, and the combined value of all medium Gephyrocupsu increases in abundance during glacials (stages 2 and 4 and the end of stage 6), similar to the abundance trends in benthic foraminifera. Finally, subordinate nannofossil taxa also show distinctive climatic trends during the last glacial cycle: (1) Syrucosphaera spp. are present in increased abundance during warmer extremes in climate (substages 5e, 5a, and stage 1); (2) Coccolithus pelagicus and Culcidiscus leptoporus dominate the subordinate nannofossil taxa, and their relative proportions seem to provide a useful paleoceanographic index, with C. pelagicus dominating when the Polar Front Zone is over the site (stages 6, 4 and 2), whilst C. leptoporus is relatively more abundant when the STC is positioned over the site (stages 1 and 5e). Increased abundance of C. pelagicus also can indicate intensified coastal upwelling.

Southern Ocean CO2 uptake and buffer factor from model run 2012-2100, with links to model results

The Southern Ocean is a key region for global carbon uptake and is characterised by a strong seasonality with the annual CO2 uptake being mediated by biological carbon draw-down in summer. Here, we show that the contribution of biology to CO2 uptake will become even more important until 2100. This is the case even if biological production remains unaltered and can be explained by the decreasing buffer capacity of the ocean as its carbon content increases. The same amount of biological carbon draw-down leads to a more than twice as large reduction in CO2 (aq) concentration and hence to a larger CO2 gradient between ocean and atmosphere that drives the gas-exchange. While the winter uptake south of 44°S changes little, the summer uptake increases largely and is responsible for the annual mean response. The combination of decreasing buffer capacity and strong seasonality of biological carbon draw-down introduces a strong and increasing seasonality in the anthropogenic carbon uptake.

Impact of CO2-driven acidification on the development of the sea cucumber Apostichopus japonicus (Selenka) (Echinodermata: Holothuroidea)

We evaluated the impact of ocean acidification on the early development of sea cucumber Apostichopus japonicus. The effect of pH-levels (pH 8.04, 7.85, 7.70 and 7.42) were tested on post-fertilization success, developmental (stage duration) and growth rates. Post-fertilization success decreased linearly with pH leading to a 6% decrease at pH 7.42 as compared to pH 8.1. The impact of pH on developmental time was stage-dependent: (1) stage duration increased linearly with decreasing pH in early-auricularia stage; (2) decreased linearly with decreasing pH in the mid-auricularia stage; but (3) pH decline had no effect on the late-auricularia stage. At the end of the experiment, the size of doliolaria larvae linearly increased with decreasing pH. In conclusion, a 0.62 unit decrease in pH had relatively small effects on A. japonicus early life-history compared to other echinoderms, leading to a maximum of 6% decrease in post-fertilization success and subtle effects on growth and development.

(Appendix) Late Cretaceous calcareous nannofossils of DSDP Holes 80-549 and 80-551

he early late Cretaceous (Cenomanian-early Turonian) is thought to have been one of the warmest periods of the Phanerozoic. This period was characterised by tropical sea surface temperatures of up to 36 °C and a pole-to-equator-gradient of less than 10 °C. The subsequent Turonian-Maastrichtian was characterised by a continuous climatic cooling, peaking in the Maastrichtian. This climatic cooling and the resulting palaeoceanographic changes had an impact on planktic primary producer communities including calcareous nannofossils. In order to gain a better understanding of these Cenomanian-Maastrichtian palaeoceanographic changes, calcareous nannofossils have been studied from the proto North Atlantic (Goban Spur, DSDP Sites 549, 551). In order to see potential differences between open oceanic and shelf dwelling nannofossils, the data from Goban Spur have been compared to findings from the European shelf (northern Germany). A total of 77 samples from Goban Spur were studied for calcareous nannofossils revealing abundant (mean 6.2 billion specimens/g sediment) and highly diverse (mean 63 species/sample) nannofossil assemblages. The dominant taxa are Watznaueria spp. (mean 30.7%), Prediscosphaera spp. (mean 18.3%), Zeugrhabdotus spp. (mean 8.3%), Retecapsa spp. (mean 7.2%) and Biscutum spp. (mean 6.6%). The Cenomanian assemblages of both Goban Spur (open ocean) and Wunstorf (shelf) are characterised by elevated abundances of high fertility taxa like Biscutum spp., Zeugrhabdotus spp. and Tranolithus orionatus. Early Turonian to Maastrichtian calcareous nannofossil assemblages of Goban Spur are, however, quite different to those described from European sections. Oceanic taxa like Watznaueria spp., Retecapsa spp. and Cribrosphearella ehrenbergii dominate in Goban Spur whereas the fertility indicators Biscutum spp. and T. orionatus are more abundant in the European shelf assemblages. This shift from a homogeneous distribution of calcareous nannofossils in the Cenomanian towards a heterogeneous one in the Turonian-Maastrichtian implies a change of the ocean circulation. The "eddy ocean" system of the Cenomanian was replaced by an oceanic circulation similar to the modern one in the Turonian-Maastrichtian, caused by the cooling. The increased pole-to-equator-gradients resulted in an oceanic circulation similar to the modern one.

Impact of atmospheric pCO2, seawater temperature, and calcification rate on the delta 18O and delta 13C composition of echinoid calcite (Echinometra viridis)

The tropical echinoid Echinometra viridis was reared in controlled laboratory experiments at temperatures of approximately 20°C and 30°C to mimic winter and summer temperatures and at carbon dioxide (CO2) partial pressures of approximately 487 ppm-v and 805 ppm-v to simulate current and predicted-end-of-century levels. Spine material produced during the experimental period and dissolved inorganic carbon (DIC) of the corresponding culture solutions were then analyzed for stable oxygen (delta 18Oe, delta 18ODIC) and carbon (The tropical echinoid Echinometra viridis was reared in controlled laboratory experiments at temperatures of approximately 20°C and 30°C to mimic winter and summer temperatures and at carbon dioxide (CO2) partial pressures of approximately 487 ppm-v and 805 ppm-v to simulate current and predicted-end-of-century levels. Spine material produced during the experimental period and dissolved inorganic carbon (DIC) of the corresponding culture solutions were then analyzed for stable oxygen (delta18Oe, delta18ODIC) and carbon (delta13Ce, delta13CDIC) isotopic composition. Fractionation of oxygen stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (delta18O = delta18Oe - delta18ODIC) was significantly inversely correlated with seawater temperature but not significantly correlated with atmospheric pCO2. Fractionation of carbon stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (Delta delta13C = delta13Ce - delta13CDIC) was significantly positively correlated with pCO2 and significantly inversely correlated with temperature, with pCO2 functioning as the primary factor and temperature moderating the pCO2-delta13C relationship. Echinoid calcification rate was significantly inversely correlated with both delta18O and delta13C, both within treatments (i.e., pCO2 and temperature fixed) and across treatments (i.e., with effects of pCO2 and temperature controlled for through ANOVA). Therefore, calcification rate and potentially the rate of co-occurring dissolution appear to be important drivers of the kinetic isotope effects observed in the echinoid spines. Study results suggest that echinoid delta18O monitors seawater temperature, but not atmospheric pCO2, and that echinoid delta13C monitors atmospheric pCO2, with temperature moderating this relationship. These findings, coupled with echinoids' long and generally high-quality fossil record, supports prior assertions that fossil echinoid delta18O is a viable archive of paleo-seawater temperature throughout Phanerozoic time, and that delta13C merits further investigation as a potential proxy of paleo-atmospheric pCO2. However, the apparent impact of calcification rate on echinoid delta18O and delta13C suggests that paleoceanographic reconstructions derived from these proxies in fossil echinoids could be improved by incorporating the effects of growth rate.13Ce, delta13CDIC) isotopic composition. Fractionation of oxygen stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (delta18O = delta18Oe - delta18ODIC) was significantly inversely correlated with seawater temperature but not significantly correlated with atmospheric pCO2. Fractionation of carbon stable isotopes between the echinoid spines and DIC of their corresponding culture solutions (delta13C = delta13Ce - delta13CDIC) was significantly positively correlated with pCO2 and significantly inversely correlated with temperature, with pCO2 functioning as the primary factor and temperature moderating the pCO2-delta13C relationship. Echinoid calcification rate was significantly inversely correlated with both delta18O and delta13C, both within treatments (i.e., pCO2 and temperature fixed) and across treatments (i.e., with effects of pCO2 and temperature controlled for through ANOVA). Therefore, calcification rate and potentially the rate of co-occurring dissolution appear to be important drivers of the kinetic isotope effects observed in the echinoid spines. Study results suggest that echinoid delta18O monitors seawater temperature, but not atmospheric pCO2, and that echinoid delta13C monitors atmospheric pCO2, with temperature moderating this relationship. These findings, coupled with echinoids' long and generally high-quality fossil record, supports prior assertions that fossil echinoid delta18O is a viable archive of paleo-seawater temperature throughout Phanerozoic time, and that delta13C merits further investigation as a potential proxy of paleo-atmospheric pCO2. However, the apparent impact of calcification rate on echinoid delta18O and delta13C suggests that paleoceanographic reconstructions derived from these proxies in fossil echinoids could be improved by incorporating the effects of growth rate.

Phenotypical responses of Fucus vesiculosus germlings to warming, acidification, nutrient enrichment and local upwelling under natural fluctuations (July - September 2014)

During summer 2014 (mid-July - mid-September 2014), early life-stage Fucus vesiculosus were exposed to combined ocean acidification and warming (OAW) in the presence and absence of enhanced nutrient levels (OAW x N experiment). Subsequently, F. vesiculosus germlings were exposed to a final upwelling disturbance during 3 days (mid-September 2014). Experiments were performed in the near-natural scenario "Kiel Outdoor Benthocosms" including natural fluctuations in the southwestern Baltic Sea, Kiel Fjord, Germany (54°27 'N, 10°11 'W). Genetically different sibling groups and different levels of genetic diversity were employed to test to which extent genetic variation would result in response variation. The data presented here show the phenotypical response (growth and survival) of the different experimental populations of F. vesiculosus under OAW, nutrient enrichment and the upwelling event. Log effect ratios demonstrate the responses to enhanced OAW and nutrient concentrations relative to the ambient conditons. Carbon, nitrogen content (% DW) and C:N ratios were measured after the exposure of ambient and high nutrient levels. Abiotic conditions the OAW x nutrient experiment and the upwelling event, are shown.