Morphological variability of Globorotalia menardii in DSDP Sites 68-502 and 68-503

Variability in the test of Globorotalia menardii during the past 8 million years has been investigated at DSDP Site 502A (Caribbean Sea) and DSDP Site 503A (Eastern Equatorial Pacific). Measurements were made of spire height (delta x), maximum diameter (delta y), the tangent angles of the upper and lower peripheral keels (phi 1, phi 2, respectively), the number of chambers in the final whorl, and the area of the silhouette in keel view. Four morphotypes alpha, beta, gamma, and delta were distinguished. Morphotype alpha was found in strata ranging in age from the Late Miocene through the Holocene. It shows a continuous increase in delta x and delta y until the Late Pleistocene. During and after the final closure of the ancient Central American Seaway (between 2.4 Ma and 1.8 Ma) there was a rapid increase in the area of the test in keel view. At the Caribbean Sea site, morphotype beta evolved during the past 0.22 Ma. It is less inflated than alpha and has a more delicate test. In the morphospace of delta x vs. delta y, morphotypes alpha and beta can be distinguished by a separation line delta y = 3.2 * delta x - 160 (delta x and delta y in µm). Plots of morphotype alpha are below that line, those of beta are above it. Morphotype alpha is taken to be Globorotalia menardii menardii Parker, Jones & Brady (1865) and includes G. menardii 'A' Bolli (1970). Morphotype beta is identified as G. menardii cultrata (d'Orbigny). Morphotypes gamma and delta are extinct Upper Miocene to Pliocene forms which evolved from morphotype alpha. They have a narrower phi 1 angle and more chambers (>=7) than morphotype alpha commonly with 5 to 6 chambers (7 in transitional forms). In contemporaneous samples morphotype delta can be distinguished from gamma by a smaller value of phi 1 and 8 or more chambers in the final whorl. Morphotype gamma is taken to be G. limbata (Fornasini, 1902) and includes the junior synonym G. menardii 'B' Bolli (1970). Morphotype delta is G. multicamerata Cushman & Jarvis (1930). With the exception of the Late Pleistocene development of G. menardii cultrataonly in the Caribbean the morphological changes of G. menardii at DSDP Sites 502A and 503A are similar. The development from the ancestral G. menardii menardii of the G. limbata - G. multicamerata lineage during the Pliocene and of G. menardii cultrata during the Late Pleistocene suggests responses at the two sites to a changing palaeoceanography during and after the formation of the Isthmus of Panama.

Grain size distribution and sea level changes in Gåsfjärden fjord

We analysed long-term variations in grain-size distribution in sediments from Gåsfjärden, a fjord-like inlet on the south-west Baltic Sea, and explored potential drivers of the recorded changes in sediment grain-size data. Over the last 5.4 thousand years (ka), the relative sea level decreased 17 m in the study region, caused by isostatic land uplift. As a consequence, Gåsfjärden has been transformed from an open coastal setting into a semi-closed inlet surrounded on the east by numerous small islands. To quantitatively estimate the morphological changes in Gåsfjärden over the last 5.4 ka and to further link the changes to our grain-size data, a digital elevation model (DEM)-based openness index was calculated. In the period between 5.4 and 4.4 ka BP, the inlet was characterised by the largest openness index. During this interval, the highest sand contents (~0.4 %) and silt/clay ratios (~0. 3) in the sediment sequence were recorded, indicating relatively high bottom water energy. After 4.4 ka BP, the average sand content was halved to ~0.2 % and the silt/clay ratios showed a significant decreasing trend over the last 4 ka. These changes are found to be associated with the gradual embayment of Gåsfjärden as represented in the openness index. The silt/clay ratios exhibited a delayed and slower change compared with the sand contents, which further suggest that finer particles are less sensitive to changes in hydrodynamic energy. Our DEM-based coastal openness index has proved to be a useful tool for interpreting the sedimentary grain-size record.

Shoreline change on nine reef islands over a seven-decade period

Atoll islands are subject to a variety of processes that influence their geomorphological development. Analysis of historical shoreline changes using remotely sensed images has become an efficient approach to both quantify past changes and estimate future island response. However, the detection of long-term changes in beach width is challenging mainly for two reasons: first, data availability is limited for many remote Pacific islands. Second, beach environments are highly dynamic and strongly influenced by seasonal or episodic shoreline oscillations. Consequently, remote-sensing studies on beach morphodynamics of atoll islands deal with dynamic features covered by a low sampling frequency. Here we present a study of beach dynamics for nine islands on Takú Atoll, Papua New Guinea, over a seven-decade period. A considerable chronological gap between aerial photographs and satellite images was addressed by applying a new method that reweighted positions of the beach limit by identifying "outlier" shoreline positions. On top of natural beach variability observed along the reweighted beach sections, we found that one third of the analyzed islands show a statistically significant decrease in reweighted beach width since 1943. The total loss of beach area for all islands corresponds to 44% of the initial beach area. Variable shoreline trajectories suggest that changes in beach width on Takú Atoll are dependent on local control (that is, human activity and longshore sediment transport). Our results show that remote imagery with a low sampling frequency may be sufficient to characterize prominent morphological changes in planform beach configuration of reef islands.

Stable carbon and oxygen isotope ratios of Miocene to Recent foraminifera from the Rio Grande Rise, South Atlantic

Miocene to Recent species of planktic foraminifera in the Globorotalia (Globoconella) lineage evolved entirely within the thermocline. All species are most abundant within subtropical-temperate watermasses throughout their history. The near stasis in distribution within the thermocline and the subtropical convergence suggests the major morphological changes in Globorotalia (Globoconella) may have occurred through habitat subdivision rather than by vicariant shifts into new watermasses. At the Rio Grande Rise, in the South Atlantic, modern G. inflata is 0.66-0.84? more positive for delta18O than the most enriched coexisting Globigerinoides sacculifer and probably grows in the mid thermocline deeper than 325 m. All extinct globoconellid species have mean delta18O ratios 0.5-0.8? more positive than Globigerinoides trilobus and G. sacculifer and probably lived within the thermocline as well. Major events in skeletal evolution are poorly correlated with changes in delta18O in this group. These include evolutionary transitions to compressed, smooth-walled tests and acquisition of keels. In addition, morphological reversals from the umbilically-inflated G. conomiozea to biconvex G. pliozea and to unkeeled G. puncticulata occur in the absence of changes in delta18O signature. Instead, the ranges of delta18O between different species almost completely overlap once corrected for temporal changes in delta18O of sea water. Foraminifera morphologies have been widely considered to evolve in response to changes in watermasses or depth habitats. However, the variety of skeletal shapes in the globoconellid lineage apparently are not adaptations to a progressive radiation from the surface mixed layer into deeper waters.

Skeletal trade-offs in coralline algae in response to ocean acidification

Ocean acidification is changing the marine environment, with potentially serious consequences for many organisms. Much of our understanding of ocean acidification effects comes from laboratory experiments, which demonstrate physiological responses over relatively short timescales. Observational studies and, more recently, experimental studies in natural systems suggest that ocean acidification will alter the structure of seaweed communities. Here, we provide a mechanistic understanding of altered competitive dynamics among a group of seaweeds, the crustose coralline algae (CCA). We compare CCA from historical experiments (1981-1997) with specimens from recent, identical experiments (2012) to describe morphological changes over this time period, which coincides with acidification of seawater in the Northeastern Pacific. Traditionally thick species decreased in thickness by a factor of 2.0-2.3, but did not experience a change in internal skeletal metrics. In contrast, traditionally thin species remained approximately the same thickness but reduced their total carbonate tissue by making thinner inter-filament cell walls. These changes represent alternative mechanisms for the reduction of calcium carbonate production in CCA and suggest energetic trade-offs related to the cost of building and maintaining a calcium carbonate skeleton as pH declines. Our classification of stress response by morphological type may be generalizable to CCA at other sites, as well as to other calcifying organisms with species-specific differences in morphological types.

Seawater carbonate chemistry, nutrients, chlorophyll and diatom Proboscia alata during experiments, 2012

The combined effects of different light and aqueous CO2 conditions were assessed for the Southern Ocean diatom Proboscia alata(Brightwell) Sundström in laboratory experiments. Selected culture conditions (light and CO2(aq)) were representative for the natural ranges in the modern Southern Ocean. Light conditions were 40 (low) and 240 (high) µmol photons/m**2/s. The three CO2(aq) conditions ranged from 8 to 34 µmol/kg CO2(aq) (equivalent to a pCO2 from 137 to 598 µatm, respectively). Clear morphological changes were induced by these different CO2(aq) conditions. Cells in low [CO2(aq)] formed spirals, while many cells in high [CO2(aq)] disintegrated. Cell size and volume were significantly affected by the different CO2(aq) concentrations. Increasing CO2(aq) concentrations led to an increase in particulate organic carbon concentrations per cell in the high light cultures, with exactly the opposite happening in the low light cultures. However, other parameters measured were not influenced by the range of CO2(aq) treatments. This included growth rates, chlorophyll aconcentration and photosynthetic yield (FV/FM). Different light treatments had a large effect on nutrient uptake. High light conditions caused an increased nutrient uptake rate compared to cells grown in low light conditions. Light and CO2 conditions co-determined in various ways the response of P. alata to changing environmental conditions. Overall P. alata appeared to be well adapted to the natural variability in light availability and CO2(aq) concentration of the modern Southern Ocean. Nevertheless, our results showed that P. alata is susceptible to future changes in inorganic carbon concentrations in the Southern Ocean.

Morphometric redefinition of the component chronospecies of the Globorotalia conoidea - G. inflata lineage in DSDP Hole 29-284

Phylo-zonations (or lineage-zonations) are based upon morphological changes within individual evolutionary lineages. These zonations, although potentially of use for stratigraphic subdivision and correlation, often suffer from a lack of quantitative exactness in the definitions of chronospecies. Thus exact reproducibility is hindered for stratigraphic determinations. The potential of morphometrically defined phylo-zonations is demonstrated on a temperate South Pacific Late Cenozoic lineage of planktonic foraminifera (Globorotalia conoidea through intermediate forms to Globorotalia inflata in DSDP Site 284) exhibiting phyletic gradualism. Our sampling interval is about 0.1 m.y. during the last 8 m.y. Changes in the number of chambers in the final whorl, test conicalness, percentage of keeled forms, and test roundness or inflatedness, are used to quantitatively define the following five chronospecies: G. conoidea (Late Miocene; 6.1->8.3 m.y.), G. conomiozea (latest Miocene ; 5.3-6.1 m.y.), G. puncticulata sphericomiozea (earliest Pliocene; 4.5-5.3 m.y.), G. puncticulata puncticulata (Early-Middle Pliocene; 2.9-4.5 m.y.), and G. inflata (Late Pliocene-Quaternary; 0-2.9 m.y.). This phylo-zonation is directly applicable to temperate cool subtropical Southern Hemisphere areas where the evolution took place (Kennett, 1967, 1973; Scott, 1979). It is still not known if the lineage occurs elsewhere; thus the applicability of the phylo-zonation over broader areas is still uncertain. Trends in general size and aperture shape seem to be climatically controlled, and thus may be only of local stratigraphic utility. The practical applications of morphometric phylo-zonation for stratigraphy is to a large extent dependent upon the amount of time and effort required to statistically define the trends. Experiments with large numbers of subsamples from this lineage demonstrate that accurate stratigraphic determinations are possible from measurements on only 15 specimens per sample, except for those very close to chronospecies boundaries.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Seawater carbonate chemistry and biological processes of sand dollars Dendraster excentricus during experiments, 2011

Reduction in global ocean pH due to the uptake of increased atmospheric CO2 is expected to negatively affect calcifying organisms, including the planktonic larval stages of many marine invertebrates. Planktonic larvae play crucial roles in the benthic-pelagic life cycle of marine organisms by connecting and sustaining existing populations and colonizing new habitats. Calcified larvae are typically denser than seawater and rely on swimming to navigate vertically structured water columns. Larval sand dollars Dendraster excentricus have calcified skeletal rods supporting their bodies, and propel themselves with ciliated bands looped around projections called arms. Ciliated bands are also used in food capture, and filtration rate is correlated with band length. As a result, swimming and feeding performance are highly sensitive to morphological changes. When reared at an elevated PCO2 level (1000 ppm), larval sand dollars developed significantly narrower bodies at four and six-arm stages. Morphological changes also varied between four observed maternal lineages, suggesting within-population variation in sensitivity to changes in PCO2 level. Despite these morphological changes, PCO2 concentration alone had no significant effect on swimming speeds. However, acidified larvae had significantly smaller larval stomachs and bodies, suggesting reduced feeding performance. Adjustments to larval morphologies in response to ocean acidification may prioritize swimming over feeding, implying that negative consequences of ocean acidification are carried over to later developmental stages.

The morphological response of Emiliania huxleyi to seawater carbonate chemistry changes: an inter-strain comparison

Four strains of the coccolithophore Emiliania huxleyi (RCC1212, RCC1216, RCC1238, RCC1256) were grown in dilute batch culture at four CO2 levels ranging from ~200 µatm to ~1200 µatm. Coccolith morphology was analyzed based on scanning electron micrographs. Three of the four strains did not exhibit a change in morphology over the CO2 range tested. One strain (RCC1256) displayed an increase in the percentage of malformed coccoliths with increasing CO2 concentration. We conclude that the sensitivity of the coccolith-shaping machinery to carbonate chemistry changes is strain-specific. Although it has been shown before that carbonate chemistry related changes in growth- and calcification rate are strain-specific, there seems to be no consistent correlation between coccolith morphology and growth or calcification rate. We did not observe an increase in the percentage of incomplete coccoliths in RCC1256, indicating that the coccolith-shaping machinery per se is affected by acidification and not the signalling pathway that produces the stop-signal for coccolith growth.

Morphological plasticity of the coral skeleton under CO2-driven seawater acidification

Ocean acidification causes corals to calcify at reduced rates, but current understanding of the underlying processes is limited. Here, we conduct a mechanistic study into how seawater acidification alters skeletal growth of the coral Stylophora pistillata. Reductions in colony calcification rates are manifested as increases in skeletal porosity at lower pH, while linear extension of skeletons remains unchanged. Inspection of the microstructure of skeletons and measurements of pH at the site of calcification indicate that dissolution is not responsible for changes in skeletal porosity. Instead, changes occur by enlargement of corallite-calyxes and thinning of associated skeletal elements, constituting a modification in skeleton architecture. We also detect increases in the organic matrix protein content of skeletons formed under lower pH. Overall, our study reveals that seawater acidification not only causes decreases in calcification, but can also cause morphological change of the coral skeleton to a more porous and potentially fragile phenotype.