Coral-algae metabolism and diurnal changes in the CO2-carbonate system of bulk sea water

Precise measurements were conducted in continuous flow seawater mesocosms located in full sunlight that compared metabolic response of coral, coral-macroalgae and macroalgae systems over a diurnal cycle. Irradiance controlled net photosynthesis (Pnet), which in turn drove net calcification (Gnet), and altered pH. Pnet exerted the dominant control on [CO3]2- and aragonite saturation state (Omega arag) over the diel cycle. Dark calcification rate decreased after sunset, reaching zero near midnight followed by an increasing rate that peaked at 03:00 h. Changes in Omega arag and pH lagged behind Gnet throughout the daily cycle by two or more hours. The flux rate Pnet was the primary driver of calcification. Daytime coral metabolism rapidly removes dissolved inorganic carbon (DIC) from the bulk seawater and photosynthesis provides the energy that drives Gnet while increasing the bulk water pH. These relationships result in a correlation between Gnet and Omega arag, with Omega arag as the dependent variable. High rates of H+ efflux continued for several hours following mid-day peak Gnet suggesting that corals have difficulty in shedding waste protons as described by the Proton Flux Hypothesis. DIC flux (uptake) followed Pnet and Gnet and dropped off rapidly following peak Pnet and peak Gnet indicating that corals can cope more effectively with the problem of limited DIC supply compared to the problem of eliminating H+. Over a 24 h period the plot of total alkalinity (AT) versus DIC as well as the plot of Gnet versus Omega arag revealed a circular hysteresis pattern over the diel cycle in the coral and coral-algae mesocosms, but not the macroalgae mesocosm. Presence of macroalgae did not change Gnet of the corals, but altered the relationship between Omega arag and Gnet. Predictive models of how future global changes will effect coral growth that are based on oceanic Omega arag must include the influence of future localized Pnet on Gnet and changes in rate of reef carbonate dissolution. The correlation between Omega arag and Gnet over the diel cycle is simply the response of the CO2-carbonate system to increased pH as photosynthesis shifts the equilibria and increases the [CO3]2- relative to the other DIC components of [HCO3]- and [CO2]. Therefore Omega arag closely tracked pH as an effect of changes in Pnet, which also drove changes in Gnet. Measurements of DIC flux and H+ flux are far more useful than concentrations in describing coral metabolism dynamics. Coral reefs are systems that exist in constant disequilibrium with the water column.

Calcareous nannofossils of ODP Leg 101 holes

Leg 101 of the Ocean Drilling Program drilled 19 holes at 11 sites to investigate the geology of the Straits of Florida and the northern Bahamas. Drilling at Site 626 indicated that the Gulf Stream has had significant flow through the Straits of Florida for at least the last 24 million years. Winnowed, foraminiferal grainstones and packstones with sparse nannofossil assemblages and the reworking of older nannofossils suggest strong bottom-current activity throughout this interval. Drilling north of Little Bahama Bank and in Exuma Sound documents the growth of platform slopes during the late Cenozoic. Nannofossil biostratigraphy of the upper Cenozoic sediments from the Little Bahama Bank and Exuma Sound slope transects indicates relatively continuous deposition, with only short breaks in the periplatform ooze and/or calciturbidite accumulation during the late Pliocene. These unconformities may be linked to sea-level lowstands. Nannofossil assemblages are generally poorly preserved owing to accelerated diagenesis caused by high aragonite and high magnesium calcite contents of bank-derived material. High rates of influx of bank-derived materials appear to coincide with highstands of sea level. Periplatform sediments are largely limited to the upper Cenozoic at Little Bahama Bank. Pelagic and/or hemipelagic conditions existed during the Late Cretaceous and Paleogene. A relatively complete, continuous section of Oligocene is present in the Little Bahama Bank area, although the rest of the Paleogene is thin. Paleogene material is also present in Northeast Providence Channel, although its thickness is uncertain. A thick upper Campanian chalk sequence with abundant, moderately to well-preserved nannofossils occurs in the Little Bahama Bank area. Hemipelagic nannofossil marls and marly chalks at Little Bahama Bank contain an excellent nannofossil record, which indicates a continuous lowermost to middle Cenomanian sequence overlying the upper Albian drowned platform. These hemipelagic sediments are significantly younger than the organic-rich, middle Albian limestones in Northeast Providence Channel. The latter indicate that a deep-water channel was already well established by the middle Albian.

Pliocene-Pleistocene calcareous nannofossils from the Iberian Abyssal Plain

During Ocean Drilling Program Leg 149, five sites were drilled on the Iberia Abyssal Plain, west of the Iberian Peninsula. Five holes (Holes 897A, 897C, 898A, 899A, and 900A) yielded Pliocene-Pleistocene sediments, which consist mainly of turbidites. Among these, Holes 897C and 898A yielded significant Pliocene-Pleistocene sediments that provided a high-resolution nannofossil biostratigraphy essential for locating paleomagnetic polarity events and for interpreting the age and frequency of turbidite sedimentation in the Iberia Abyssal Plain. Pliocene-Pleistocene nannofossils recovered during Leg 149 are generally abundant and well to moderately preserved. Although reworking is evident in most samples, the Pliocene-Pleistocene nannofossils proved quite reliable for dating the sediments. Most Pleistocene zonal boundaries proposed by S. Gartner in 1977 and the Pliocene standard zonal boundaries proposed by E. Martini in 1971 were easily recognized. In addition, several other nannofossil events proposed by D. Rio et al. in 1990 and by T. Sato and T. Takayama in 1992 were recognized and proved valuable for improving the resolution of Pliocene-Pleistocene nannofossil biostratigraphy. The Pliocene-Pleistocene nannofossil biostratigraphic results of Holes 897C and 900A coincide rather well with the discerned paleomagnetic polarity events. As a result, the combination of nannofossil biostratigraphic and paleomagnetic studies provides important information for fulfilling the second objective of this leg: to determine the history of turbidite sedimentation in the Iberia Abyssal Plain. The general trend of sedimentation rates inferred by nannofossil biostratigraphy indicates that sedimentation rates increase from the continental margin to the deep sea along with increasing water depth.

Calcareous nannofossils in ODP Hole 131-808C

ODP Leg 131 recovered nannofossil-bearing sediments from Site 808 in the Nankai Trough, western Pacific Ocean. Three holes were examined for nannofossils, 808A, 808B, and 808C. A total of 22 nannofossil events were recognized, of which 10 are used as zonal markers. The sediments recovered from Hole 808A (0-111.4 mbsf) contain Pleistocene nannofossil assemblages that are mostly well preserved. All samples from this hole were assigned to nannofossil Zone NN21. The nannofossil assemblages observed in Hole 808B (111.0-358.8 mbsf) are poorly to well preserved and were all assigned to the Pleistocene. The NN21/NN20 Boundary is placed at 230.7 ± 4.4 mbsf. Hole 808C was cored from 298.5 to 1327 mbsf and basalt was reached at 1289.9 mbsf. The sediments recovered range in age from the upper part of Zone NN20 of the Pleistocene to Zone NN5 of the middle Miocene and contain poorly to well-preserved nannofossil assemblages. The Pliocene/Pleistocene Boundary, marked by the FO Gephyrocapsa caribbeanica, was placed at 776.3 ±1.6 mbsf, and the Miocene/Pliocene Boundary is tentatively placed at 955.9 ±1.5 mbsf. The lowermost sediments above basement as well as a sediment sample intercalated between basalt flows are assigned to Zone NN5, with an age of approximately 15 Ma. Age estimates provided by nannofossils show that the sedimentation rate in the trench-fill deposits of the Nankai Trough was very high, 800-1350 m/m.y (0-0.46 Ma), whereas in the Shikoku Basin deposits (> 0.46 Ma), the sedimentation rate was much lower (24-200 m/m.y). These age estimates also provide an extrapolated age of approximately 15 Ma for the basaltic basement at Site 808.

Dolomite-rich coralline algae in reefs resist dissolution in acidified conditions

Coral reef ecosystems develop best in high-flow environments but their fragile frameworks are also vulnerable to high wave energy. Wave-resistant algal rims, predominantly made up of the crustose coralline algae (CCA) Porolithon onkodes and P. pachydermum, are therefore critical structural elements for the survival of many shallow coral reefs. Concerns are growing about the susceptibility of CCA to ocean acidification because CCA Mg-calcite skeletons are more susceptible to dissolution under low pH conditions than coral aragonite skeletons. However, the recent discovery of dolomite (Mg0.5Ca0.5(CO3)), a stable carbonate, in P. onkodes cells necessitates a reappraisal of the impacts of ocean acidification on these CCA. Here we show, using a dissolution experiment, that dried dolomite-rich CCA have 6-10 times lower rates of dissolution than predominantly Mg-calcite CCA in both high-CO2 (~ 700 ppm) and control (~ 380 ppm) environments, respectively. We reveal this stabilizing mechanism to be a combination of reduced porosity due to dolomite infilling and selective dissolution of other carbonate minerals. Physical break-up proceeds by dissolution of Mg-calcite walls until the dolomitized cell eventually drops out intact. Dolomite-rich CCA frameworks are common in shallow coral reefs globally and our results suggest that it is likely that they will continue to provide protection and stability for coral reef frameworks as CO2 rises.

Diffusion boundary layers ameliorate the negative effects of ocean acidification on the temperate coralline macroalga Arthrocardia corymbosa

Anthropogenically-modulated reductions in pH, termed ocean acidification, could pose a major threat to the physiological performance, stocks, and biodiversity of calcifiers and may devalue their ecosystem services. Recent debate has focussed on the need to develop approaches to arrest the potential negative impacts of ocean acidification on ecosystems dominated by calcareous organisms. In this study, we demonstrate the role of a discrete (i.e. diffusion) boundary layer (DBL), formed at the surface of some calcifying species under slow flows, in buffering them from the corrosive effects of low pH seawater. The coralline macroalga Arthrocardia corymbosa was grown in a multifactorial experiment with two mean pH levels (8.05 'ambient' and 7.65 a worst case 'ocean acidification' scenario projected for 2100), each with two levels of seawater flow (fast and slow, i.e. DBL thin or thick). Coralline algae grown under slow flows with thick DBLs (i.e., unstirred with regular replenishment of seawater to their surface) maintained net growth and calcification at pH 7.65 whereas those in higher flows with thin DBLs had net dissolution. Growth under ambient seawater pH (8.05) was not significantly different in thin and thick DBL treatments. No other measured diagnostic (recruit sizes and numbers, photosynthetic metrics, %C, %N, %MgCO3) responded to the effects of reduced seawater pH. Thus, flow conditions that promote the formation of thick DBLs, may enhance the subsistence of calcifiers by creating localised hydrodynamic conditions where metabolic activity ameliorates the negative impacts of ocean acidification.

Maastrichtian calcareous nannofossils of Kronsmoor-Section

The latest Campanian-earliest Maastrichtian interval is well known as a period of intense climate cooling. This cooling caused a distinctive bipolar biogeographic distribution of calcareous nannofossil assemblages: High latitude settings were dominated by newly evolving endemic taxa, former cosmopolitan species disappeared at the same time and equatorial communities experienced an invasion of cool water taxa. The impact of this cooling on northern mid-latitude assemblages is, however, less well known. In order to overcome this gap we studied the Kronsmoor section (northwest Germany). This section provides a continuous upper Campanian - lower Maastrichtian succession with moderately to well preserved nannofossils. Uppermost Campanian assemblages are dominated by Prediscosphaera cretacea; other common taxa include Prediscosphaera stoveri, Watznaueria barnesiae and Micula staurophora. The lower Maastrichtian is characterized by lower numbers of P. cretacea and frequent Kamptnerius magnificus, Arkhangelskiella cymbiformis and Cribrosphaerella ehrenbergii. These changes reflect, in part, the Campanian-Maastrichtian boundary cooling since some successful taxa (e.g. K. magnificus) are related to cool surface waters. Other shifts in the nannofossil communities were perhaps the result of a changing nutrient regime. Stronger latitudinal gradients may have increased wind velocities and thus the eolian input of ferruginous dust required by N-fixing bacteria. The enhanced high latitude deep-water formation probably changed the bottom-water environment in disfavor of denitrificating organisms. A decline of chemical weathering and fluviatile transport may have reduced the amount of bioavailable phosphate. These processes led to an increased nitrate and a decreased phosphate content shifting the nutrient regime from nitrate towards phosphate limitation.

Calcareous nannofossils across the Danian/Selandian boundary

A high-resolution calcareous nannofossil analysis of the Danian/Selandian boundary was conducted at Site 1262 (Walvis Ridge, South Atlantic) to pinpoint the lowest occurrence of fasciculiths and to unravel the evolutionary trends throughout nannofossil Zone NP4. Using quantitative analyses, numerous primary and secondary bioevents were identified, improving the biostratigraphic resolution of this interval. The main events recorded at Site 1262 were also identified at the Zumaia section Global Stratotype Section and Point (GSSP) of the base of the Selandian and at the Qreiya section (Egypt). The lowest occurrence of fasciculiths (represented by the LO of Gomphiolithus magnicordis and Gomphiolithus magnus) was observed in the middle part of Chron C27r, above the LO of Toweius pertusus and prior to the LO of the genus Sphenolithus. The synchroneity of the LO of fasciculiths was also verified at various latitudes, such as DSDP Site 384 (NW Atlantic), ODP Site 761B (Indian Ocean) and DSDP Site 577A (Pacific Ocean). The first and second diversification events (Steurbaut and Sztrákos, 2008, doi:10.1016/j.marmicro.2007.08.004), or radiation events (Bernaola et al., 2009, doi:10.1344/105.000000272), of fasciculiths have been thoroughly discussed and well characterized by a succession of events. The occurrence of the Latest Danian Event (LDE) and several paleoenvironmental changes recognized during this time interval, coupled with an ecological competition with Sphenolithus, appear to be the probable causes of the First and Second Radiations and the fasciculith barren interval between them. The occurrence of new morphostructures and taxa suggests evolutionary trends and a strict link between morphological evolution and paleoclimate.

Calcareous dinoflagellate cysts and geochemistry of Mediterranean Sea sediments

Comparison of calcareous dinoflagellate cyst assemblages with Ba, Al, Mn, and Fe records from three sediment cores collected in the eastern Mediterranean Sea indicate that calcareous dinoflagellate cysts are generally resistant to postdepositional dissolution. Cyst association changes during and after sapropel S1 formation can therefore be closely related to variability in surface water productivity. Two groups of cysts are defined: those having highest abundances within the sapropelic and postsapropelic sediments. The temporal cyst distributions suggest increased freshwater input mainly from the Nile and a shallowing of the pycnocline as the most important processes increasing nutrient concentration in the photic zone, thus leading to increased productivity and organic carbon fluxes during sapropel formation. Furthermore, a general warming trend at the beginning of S1 formation and a slight salinity decrease are reconstructed.

Cenozoic calcareous nannofossils in DSDP Leg 77 holes

Three of the six DSDP Leg 77 sites drilled in the western approaches to the Straits of Florida yielded thick sequences of Cenozoic sediment rich in calcareous nannofossils. Hiatuses are prominent in each of these continuously cored intervals. A prominent upper Oligocene hiatus, observed at each of these three sites, can be correlated to a large-scale "global" regression event. Other disconformable horizons present in the study area cannot be positively related to sealevel fluctuations and may be caused by a number of factors including local tectonic activity. Paleogene sections are generally marked by thick accumulations within the upper Oligocene Sphenolithus ciperoensis Zone and by a pronounced braarudosphaerid-holococcolith bloom recorded in the lower Oligocene and upper Eocene. This bloom is particularly well developed at Site 540. All samples examined contain abundant nannofossils. Preservation fluctuates throughout the sections from good to poor.

Calcareous dinoflagellates in Maastrichtian to early Miocene sediments of DSDP Hole 39-357

The evolution of calcareous dinoflagellate communities has been investigated for the latest Cretaceous to earliest Neogene interval of the mid-latitude South Atlantic. In doing so, the response of calcareous dinoflagellates to Cenozoic climatic change has been addressed for the first time. Trends in species composition and distribution patterns of wall types indicate significant changes which correlate with major palaeoenvironmental modifications. A first major shift concerning the relative abundance of species and wall types occurred across the Cretaceous-Tertiary boundary. The associations remained stable during the entire Paleocene and Eocene. Only in the late Eocene did a dramatic decrease in temperature cause a slight diversification. A second major shift in the abundance patterns occurred across the Eocene-Oligocene boundary. The early Miocene warming is possibly reflected in the distinct increase in relative abundance of one species. The assemblages of calcareous dinoflagellates evidently react to major climatic changes during the Cenozoic. These poorly investigated organisms may thus provide an important contribution to the understanding of earth's palaeoclimatic evolution.

Seawater carbonate chemistry and physiological responses of three temperate coralline algae in a laboratory experiment

Coralline algae are major calcifiers of significant ecological importance in marine habitats but are among the most sensitive calcifying organisms to ocean acidification. The elevated pCO2 effects were examined in three coralline algal species living in contrasting habitats from intertidal to subtidal zones on the north-western coast of Brittany, France: (i) Corallina elongata, a branched alga found in tidal rock pools, (ii) Lithophyllum incrustans, a crustose coralline alga from the low intertidal zone, and (iii) Lithothamnion corallioides (maerl), a free-living form inhabiting the subtidal zone. Metabolic rates were assessed on specimens grown for one month at varying pCO2: 380 (current pCO2), 550, 750 and 1000 µatm (elevated pCO2). There was no pCO2 effect on gross production in C. elongata and L. incrustans but L. incrustans respiration strongly increased with elevated pCO2. L. corallioides gross production slightly increased at 1000 µatm, while respiration remained unaffected. Calcification rates decreased with pCO2 in L. incrustans (both in the light and dark) and L. corallioides (only in the light), while C. elongata calcification was unaffected. This was consistent with the lower skeletal mMg/Ca ratio of C. elongata (0.17) relative to the two other species (0.20). L. incrustans had a higher occurrence of bleaching that increased with increasing pCO2. pCO2 could indirectly impact this coralline species physiology making them more sensitive to other stresses such as diseases or pathogens. These results underlined that the physiological response of coralline algae to near-future ocean acidification is species-specific and that species experiencing naturally strong pH variations were not necessarily more resistant to elevated pCO2 than species from more stable environment.