Benthic–pelagic links and rocky intertidal communities: Bottom-up effects on top-down control?

Insight into the dependence of benthic communities on biological and physical processes in nearshore pelagic environments, long considered a “black box,” has eluded ecologists. In rocky intertidal communities at Oregon coastal sites 80 km apart, differences in abundance of sessile invertebrates, herbivores, carnivores, and macrophytes in the low zone were not readily explained by local scale differences in hydrodynamic or physical conditions (wave forces, surge flow, or air temperature during low tide). Field experiments employing predator and herbivore manipulations and prey transplants suggested top-down (predation, grazing) processes varied positively with bottom-up processes (growth of filter-feeders, prey recruitment), but the basis for these differences was unknown. Shore-based sampling revealed that between-site differences were associated with nearshore oceanographic conditions, including phytoplankton concentration and productivity, particulates, and water temperature during upwelling. Further, samples taken at 19 sites along 380 km of coastline suggested that the differences documented between two sites reflect broader scale gradients of phytoplankton concentration. Among several alternative explanations, a coastal hydrodynamics hypothesis, reflecting mesoscale (tens to hundreds of kilometers) variation in the interaction between offshore currents and winds and continental shelf bathymetry, was inferred to be the primary underlying cause. Satellite imagery and offshore chlorophyll-a samples are consistent with the postulated mechanism. Our results suggest that benthic community dynamics can be coupled to pelagic ecosystems by both trophic and transport linkages.

Stable isotope record of planktonic foraminifera of the Atlantic Ocean

The thermal structure of the upper ocean (0-1000 m) is set by surface heat fluxes, shallow wind-driven circulation, and the deeper thermohaline circulation. Its long-term variability can be reconstructed using deep-dwelling planktonic foraminifera that record subsurface conditions. Here we used six species (Neogloboquadrina dutertrei, Globorotalia tumida, Globorotalia inflata, Globorotalia truncatulinoides, Globorotalia hirsuta, and Globorotalia crassaformis) from 66 core tops along a meridional transect spanning the mid-Atlantic (42°N to 25°S) to develop a method for reconstructing past thermocline conditions. We estimated the calcification depths from d18O measurements and the Mg/Ca-temperature relationships for each species. This systematic strategy over this large latitudinal section reveals distinct populations with different Mg/Ca-temperature relationships for G. inflata, G. truncatulinoides, and G. hirsuta in different areas. The calcification depths do not differ among the different populations, except for G. hirsuta, where the northern population calcifies much shallower than the southern population. N. dutertrei and G. tumida show a remarkably constant calcification depth independent of oceanographic conditions. The deepest dweller, G. crassaformis, apparently calcifies in the oxygen-depleted zone, where it may find refuge from predators and abundant aggregated matter to feed on. We found a good match between its calcification depth and the 3.2 ml/l oxygen level. The results of this multispecies, multiproxy study can now be applied down-core to facilitate the reconstruction of open-ocean thermocline changes in the past.

(Table 2) Age determination of sediment cores from the Panama Basin

Application of the 230Th normalization method to estimate sediment burial fluxes in six cores from the eastern equatorial Pacific (EEP) reveals that bulk sediment and organic carbon fluxes display a coherent regional pattern during the Holocene that is consistent with modern oceanographic conditions, in contrast with estimates of bulk mass accumulation rates (MARs) derived from core chronologies. Two nearby sites (less than 10 km apart), which have different MARs, show nearly identical 230Th-normalized bulk fluxes. Focusing factors derived from the 230Th data at the foot of the Carnegie Ridge in the Panama Basin are >2 in the Holocene, implying that lateral sediment addition is significant in this part of the basin. New geochemical data and existing literature provide evidence for a hydrothermal source of sediment in the southern part of the Panama Basin and for downslope transport from the top of the Carnegie Ridge. The compilation of core records suggests that sediment focusing is spatially and temporally variable in the EEP. During oxygen isotope stage 2 (OIS 2, from 13-27 ka BP), focusing appears even higher compared to the Holocene at most sites, similar to earlier findings in the eastern and central equatorial Pacific. The magnitude of the glacial increase in focusing factors, however, is strongly dependent on the accuracy of age models. We offer two possible explanations for the increase in glacial focusing compared to the Holocene. The first one is that the apparent increase in lateral sediment redistribution is partly or even largely an artifact of insufficient age control in the EEP, while the second explanation, which assumes that the observed increase is real, involves enhanced deep sea tidal current flow during periods of low sea level stand.

Relative abundances of planktonic foraminifera from surface sediments of the Drake Passage, Southern Ocean (Table 1)

Based on a qualitative and quantitative evaluation of Recent sediments samples (top 3 cm of cores as well as Petersen grab samples) from the Drake Passage, between South America and Antarctica, the distribution of planktonic foraminifera and their relation to oceanographic conditions was investigated. The Antarctic Convergence - the northern limit of the cold Antarctic Surface Water - is shown to be of major importance in controlling the distributional pattern of planktonic species as well as their total numbers. South of the convergence, Globigerina pachyderma is usually the only species found in the sediment. It occurs with abundances not greater than 6000 per gram dry sediment, and at most stations less than 100 specimens per gram of dry sediment were recovered. At a number of deep-sea stations below 3700 m depth approx. no planktonic foraminifera were found at all. It is most probable, that at least some of these stations are located below the limit of CaCO3 dissolution. North of the Antarctic Convergence planktonic foraminiferal numbers are much higher and range from 1800 to 120000 per gram of dry sediment. Eight species are the major constituents of the population: Globigerina pachyderma, Globigerina bulloides, Globogerina quinqueloba, Globigerina inflata, Globorotalia truncatolinoides, Globorotalia scitula, Globigerinita glutinata and Globigerinita uvula. The widespread occurrence of Globorotalia truncatulinoides, which in the northern hemisphere is usually a subtropical form, is especially noteworthy. Another Globigerina, morphologically similar to G. pachyderma, has been recognized frequently north of the Antarctic Convergence. Globigerina megastoma which has its type area in the Drake Passage, has been found only rarely. Orbulina universa occurs in samples from the areas of higher water temperature around the South American Continent. Globigerina pachyderma is predominantly sinistrally coiled throughout the area investigated, but a slight increase in the percentage of dextrally coiled specimens may be noticed with increasing water temperature, i.e. from south to north.

Short-term variations in radiolarian faunal dsitribution in the eastern equatorial Pacific

Two short time intervals centered at 2.3 and 4.7 Ma were studied to investigate short-term variations in surface-ocean processes as indicated by changes in the radiolarian microfossil population. These time intervals represent two different settings of late Neogene climate. The older interval represents a time when tropical circulation between the Pacific and Atlantic oceans was not blocked by the Isthmus of Panama, whereas the younger interval represents a time when Northern Hemisphere glaciation was present but did not display the dominance of the 100,000-yr cycle that characterizes the late Pleistocene. The younger time slice at 2.3 Ma was sampled at all Leg 138 sites except Site 844, where significant reworking was evident. All sites except 844, 853, and 854 were sampled for the older time slice. Samples were taken at 10- to 20-cm intervals at each site and spanned a GRAPE density maximum and minimum. Thus, it was possible to investigate whether the changes in carbonate content (as indicated by GRAPE density) were associated with changes in surface-ocean conditions (indicated by radiolarian assemblage variations). For both time slices, the radiolarian data indicate that intervals of decreased carbonate content are periods of cooler water conditions and possibly enhanced biogenic production. Times of increased carbonate content are associated with inferred warmer oceanographic conditions, as indicated by the dominance of tropical assemblages at 2.3 Ma and tropical and western Pacific assemblages during the time slice centered at 4.8 Ma. However, the spatial patterns of change during each time slice show a distinct difference in the mapped patterns of radiolarian assemblage dominance. The older time slice, representing a period before the closing of the Isthmus of Panama, shows more zonal patterns presumably associated with a more zonal character of equatorial circulation. After the closing of the isthmus, the shifts in faunal patterns between times of high and low carbonates are characterized by shifts in the dominance of the tropical and transitional assemblages, respectively, throughout the region.

Geographic distribution of dinoflagellate cysts in surface sediments

Dinoflagellate cysts are useful for reconstructing upper water conditions. For adequate reconstructions detailed information is required about the relationship between modern day environmental conditions and the geographic distribution of cysts in sediments. This Atlas summarises the modern global distribution of 71 organicwalled dinoflagellate cyst species. The synthesis is based on the integration of literature sources together with data of 2405 globally distributed surface sediment samples that have been preparedwith a comparable methodology and taxonomy. The distribution patterns of individual cyst species are being comparedwith environmental factors that are knownto influence dinoflagellate growth, gamete production, encystment, excystment and preservation of their organic-walled cysts: surface water temperature, salinity, nitrate, phosphate, chlorophyll-a concentrations and bottom water oxygen concentrations. Graphs are provided for every species depicting the relationship between seasonal and annual variations of these parameters and the relative abundance of the species. Results have been compared with previously published records; an overview of the ecological significance as well as information about the seasonal production of each individual species is presented. The relationship between the cyst distribution and variation in the aforementioned environmental parameters was analysed by performing a canonical correspondence analysis. All tested variables showed a positive relationship on the 99% confidence level. Sea-surface temperature represents the parameter corresponding to the largest amount of variance within the dataset (40%) followed by nitrate, salinity, phosphate and bottom-water oxygen concentration, which correspond to 34%, 33%, 25% and 24% of the variance, respectively. Characterisations of selected environments as well as a discussion about how these factors could have influenced the final cyst yield in sediments are included.

Age model and ostracod countings from IODP Site 306-U1314

We present a detailed study of glacial/interglacial deep sea benthic ostracod assemblage variability at IODP Site U1314 (subpolar North Atlantic) in relation to the history of ice-rafting events and changes in deep ocean circulation over the past 170 ky. Our records of ostracod diversity, abundance and dissolution and sediment properties (IRD and CaCO3) show an excellent correspondence to high amplitude orbital and millennial variability observed in the climate records (d13C and d18O) from neighboring deep water sites, suggesting that the benthic meiofauna fluctuates synchronously with the prevailing oceanographic conditions (surface ocean conditions, deep ocean circulation and water temperature and food flux). Krithe (dominant), Argilloecia and Cytheropteron are the most abundant and diverse genera in association with Rockallia enigmatica. Three ostracod assemblages are recognized. The genera Pennyella, Argilloecia, Pelecocythere, Ambocythere, Pseudobosquetina, Bradleya and Nannocythere are associated with interglacials and interstadials, and possibly reflect increased flux of food to the sediments and more vigorous NADW formation. A transitional assemblage composed of species of Cytheropteron, Xestoleberis and Eucythere is restricted to climatic transitions and indicate moderate environmental conditions and seasonal productivity. A glacial/stadial assemblage is characterized by a temporal predominance of either intermediate-depth and shallow water Arctic/subarctic species (belonging to Cytheropteron, Polycope, Pedicythere, Swainocythere, Cluthia, Heterocyprideis, Elofsonella and Finmarchinella) or abyssal North Atlantic ostracods (Bythocythere, Dutoitella, Bathycythere and Bythocypris). The influx of high latitude taxa can be partially explained by ice-rafting, but may also represent a shift of the location of intermediate and deep water convection to the area south of Iceland. Therefore the combination of species characteristic of different watermasses during glacials may reflect shifts in the influence of high nutrient southern source water (e.g. AABW) vs. low nutrient GNAIW during glacials.

Bulk geochemical and lipid biomarker data for sediment core W8402A-14

Eleven sediment samples taken downcore and representing the past 26 kyr of deposition at MANOP site C (0°57.2°N, 138°57.3°W) were analyzed for lipid biomarker composition. Biomarkers of both terrestrial and marine sources of organic carbon were identified. In general, concentration profiles for these biomarkers and for total organic carbon (TOC) displayed three common stratigraphic features in the time series: (1) a maximum within the surface sediment mixed layer (<=4 ka); (2) a broad minimum extending throughout the interglacial deposit; and (3) a deep, pronounced maximum within the glacial deposit. Using the biomarker records, a simple binary mixing model is described that assesses the proportion of terrestrial to marine TOC in these sediments. Best estimates from this model suggest that ~20% of the TOC is land-derived, introduced by long-range eolian transport, and the remainder is derived from marine productivity. The direct correlation between the records for terrestrial and marine TOC with depth in this core fits an interpretation that primary productivity at site C has been controlled by wind-driven upwelling at least over the last glacial/interglacial cycle. The biomarker records place the greatest wind strength and highest primary productivity within the time frame of 18 to 22 kyr B.P. Diagenetic effects limit our ability to ascertain directly from the biomarker records the absolute magnitude that different types of primary productivity have changed at this ocean location over the past 26 kyr.

First-year and second-year snow properties on sea ice in the Weddell Sea during POLARSTERN cruise ANT-XXII/2 (ISPOL) during the drift

Observations of snow properties, superimposed ice, and atmospheric heat fluxes have been performed on first-year and second-year sea ice in the western Weddell Sea, Antarctica. Snow in this region is particular as it does usually survive summer ablation. Measurements were performed during Ice Station Polarstern (ISPOL), a 5-week drift station of the German icebreaker RV Polarstern. Net heat flux to the snowpack was 8 W/m**2, causing only 0.1 to 0.2 m of thinning of both snow cover types, thinner first-year and thicker second-year snow. Snow thinning was dominated by compaction and evaporation, whereas melt was of minor importance and occurred only internally at or close to the surface. Characteristic differences between snow on first-year and second-year ice were found in snow thickness, temperature, and stratigraphy. Snow on second-year ice was thicker, colder, denser, and more layered than on first-year ice. Metamorphism and ablation, and thus mass balance, were similar between both regimes, because they depend more on surface heat fluxes and less on underground properties. Ice freeboard was mostly negative, but flooding occurred mainly on first-year ice. Snow and ice interface temperature did not reach the melting point during the observation period. Nevertheless, formation of discontinuous superimposed ice was observed. Color tracer experiments suggest considerable meltwater percolation within the snow, despite below-melting temperatures of lower layers. Strong meridional gradients of snow and sea-ice properties were found in this region. They suggest similar gradients in atmospheric and oceanographic conditions and implicate their importance for melt processes and the location of the summer ice edge.

Nematode absolute densities and genus counts from four stations near the Antarctic Peninsula, collected with a multicorer device during POLARSTERN cruise ANT-XXIX/3 in January-March 2013

Marine free-living nematode communities were studied at similar depths (~500m) at two sides of the Antarctic Peninsula, characterised by different environmental and oceanographic conditions. At the Weddell Sea side, benthic communities are influenced by cold deep-water formation and seasonal sea-ice conditions, whereas the Drake Passage side experiences milder oceanic conditions and strong dynamics of the Antarctic Circumpolar Current. Surface primary productivity contrasted with observed benthic pigment patterns and varied according to the area studied: chlorophyll a concentrations (as a proxy for primary production) were high in the Weddell Sea sediments, but low in the surface waters above; this pattern was reversed in the Drake Passage. Differences between areas were largely mirrored by the nematode communities: nematode densities peaked in Weddell stations and showed deeper vertical occurrence in the sediment, associated with deeper penetration of chlorophyll a. Generic composition did not differ markedly between both areas, but rather showed distinct community shifts with depth in the sediment.

(Table 1) Radiocarbon dates of core JM05-085-GC, South-Barents Sea

The Late Weichselian-Early Holocene variability of the North Atlantic Current has been studied with focus on the zonal component of this meridional transport during the transition from glacial to interglacial conditions. The investigated sediment core is from 409 m water depth in the SW Barents Sea. Eight Accelerator mass spectrometry (AMS) 14C dates show that the core covers the last 20,000 cal yr B.P. with a centennial scale resolution during Late Weichselian-Early Holocene. Planktic foraminiferal assemblages were analyzed using the >100 ?m size fraction and foraminiferal planktic and benthic d13C and d18O isotopes were measured. Furthermore, a range of physical and chemical analyses has been carried out on the bulk sediment samples. Four time periods have been identified which represent the varying oceanographic conditions in Ingøydjupet, a glacial trough located off the north coast of Norway in the SW Barents Sea. 1) The late glacial (before ca 15,000 cal yr B.P.) influenced by the nearby ice sheets with high amounts of sea ice- or iceberg-transported detritus. 2) The late Oldest Dryas stadial and the Bølling-Allerød interstadial (ca 15,000-12,700 cal yr B.P.) with cold surface water conditions influenced by the collapse of the nearby ice sheets, high amounts of sea ice- or iceberg-transported detritus and melt water and weak subsurface inflow of Atlantic Water. 3) The Younger Dryas cold stadial (12,700-11,650 cal yr B.P.) with low primary productivity and extensive sea ice cover and 4) The Preboreal and Early Holocene (11,650-6800 cal yr B.P. cal yr B.P.) with strong influx of Atlantic Water into the area, near absence of ice rafted debris and generally ameliorated conditions in both surface and bottom water masses as seen from a high flux of foraminifera and increased marine primary production.

Expanded GHOST database

In this study we review a global set of alkenone- and foraminiferal Mg/Ca-derived sea surface temperatures (SST) records from the Holocene and compare them with a suite of published Eemian SST records based on the same approach. For the Holocene, the alkenone SST records belong to the actualized GHOST database (Kim, J.-H., Schneider R.R., 2004). The actualized GHOST database not only confirms the SST changes previously described but also documents the Holocene temperature evolution in new oceanic regions such as the Northwestern Atlantic, the eastern equatorial Pacific, and the Southern Ocean. A comparison of Holocene SST records stemming from the two commonly applied paleothermometry methods reveals contrasting - sometimes divergent - SST evolution, particularly at low latitudes where SST records are abundant enough to infer systematic discrepancies at a regional scale. Opposite SST trends at particular locations could be explained by out-of-phase trends in seasonal insolation during the Holocene. This hypothesis assumes that a strong contrast in the ecological responses of coccolithophores and planktonic foraminifera to winter and summer oceanographic conditions is the ultimate reason for seasonal differences in the origin of the temperature signal provided by these organisms. As a simple test for this hypothesis, Eemian SST records are considered because the Holocene and Eemian time periods experienced comparable changes in orbital configurations, but had a higher magnitude in insolation variance during the Eemian. For several regions, SST changes during both interglacials were of a similar sign, but with higher magnitudes during the Eemian as compared to the Holocene. This observation suggests that the ecological mechanism shaping SST trends during the Holocene was comparable during the penultimate interglacial period. Although this "ecology hypothesis" fails to explain all of the available results, we argue that any other mechanism would fail to satisfactorily explain the observed SST discrepancies among proxies.

(Table 2) Abundance of Radiolaria species in sediment samples from Core AMK4-340

Over the past 13 kyr the most significant natural changes in the Reykjanes ridge region took place within 13-7.8 kyr B.P. They resulted from alternating intensifications of the influence of the Labrador (LWM) and Norwegian-Greenland (NGWM) water masses. During 13-11.7 kyr B.P. natural conditions were governed by influence of LWM with sea surface temperature (SST) 3-5°C lower present one. During 11.7-10.3 kyr B.P. NGWM with SST 6-7°C lower present one predominated. During 10.3-9.5 kyr B.P. oceanographic conditions were rapidly transforming and approaching present ones controlled by interaction between LWM and North Atlantic water masses; SST abruptly increased almost to the present value. During 9.5-8.3 kyr B.P. intensification of NGWM led to small decrease of SST (1.5-2.5°C below present value; between 8.3 and 7.8 kyr B.P. natural conditions had approximated present ones and later on remained relatively stable; SST fluctuated with an amplitude of about 1.5°C.