Morphometric measurements throughout ontogeny of selected planktic foraminiferal test species

Development plays an important part in shaping adult morphology and morphological disparity, yet its influence on evolutionary processes is seldom explored because of a lack of preservation of ontogenetic stages in the fossil record. By preserving their entire ontogenetic history within their test, and with the advent of high-resolution imaging techniques, planktic foraminifera allow us to investigate the influence of developmental constraints on disparity. Using Synchrotron radiation X-ray tomographic microscopy (SRXTM), we reconstruct the ontogenetic progression of seven species across several of the major morphotypic groups of planktic foraminifera, including morphotypes of a species exhibiting high phenotypic plasticity and closely related pseudo-cryptic sister-taxa. We show differences in growth patterns between the globigerinid species, which appear more tightly regulated within the framework of isometry from the neanic stage, and the globorotaliid species, whose adult stages present allometric trends. Morphological change through ontogeny results in a change in surface area to volume ratios. Different metabolic processes therefore dominate at different stages of ontogeny, changing the vulnerability of the organism to environmental influences over growth, from factors affecting diffusion rates in the juvenile to those affecting energy supply in the adult. These findings identify some of the parameters within which evolutionary mechanisms have to act.

Floral and faunal characteristics and content of photosynthetic pigment in soils of northern Victoria Land, Antarctica

Although soil algae are among the main primary producers in most terrestrial ecosystems of continental Antarctica, there are very few quantitative studies on their relative proportion in the main algal groups and on how their distribution is affected by biotic and abiotic factors. Such knowledge is essential for understanding the functioning of Antarctic terrestrial ecosystems. We therefore analyzed biological soil crusts from northern Victoria Land to determine their pH, electrical conductivity (EC), water content (W), total and organic C (TC and TOC) and total N (TN) contents, and the presence and abundance of photosynthetic pigments. In particular, the latter were tested as proxies for biomass and coarse-resolution community structure. Soil samples were collected from five sites with known soil algal communities and the distribution of pigments was shown to reflect differences in the relative proportions of Chlorophyta, Cyanophyta and Bacillariophyta in these sites. Multivariate and univariate models strongly indicated that almost all soil variables (EC, W, TOC and TN) were important environmental correlates of pigment distribution. However, a significant amount of variation is independent of these soil variables and may be ascribed to local variability such as changes in microclimate at varying spatial and temporal scales. There are at least five possible sources of local variation: pigment preservation, temporal variations in water availability, temporal and spatial interactions among environmental and biological components, the local-scale patchiness of organism distribution, and biotic interactions.

Neogene calcareous nannofossils in the Caribbean and the eastern equatorial Pacific

Principal components analysis of nannofossil assemblages in five ODP sites in the Caribbean and the eastern equatorial Pacific was used to monitor Neogene surface water circulation changes. In 15.83-10.71 Ma, floral distributions clearly show the existence of the Circum-tropical Current between the Caribbean and the eastern equatorial Pacific and no surface water communication between the northern and southern Caribbean. In 10.71-9.36 Ma, the Circum-tropical Current weakened and the northward intra-Caribbean current had been initiated. Northern and southern Caribbean surface waters again became separated in 8.35-3.65 Ma and the Circum-tropical Current was regenerated. After 2.76 Ma, the northward intra-Caribbean current has completely been established and the Circum-tropical Current disappeared. This suggests that a barrier to the Circum-tropical Current had formed, indicating the final closure of the Isthmus of Panama.

Sporomorphs and quantitative analysis of the Eocene record from IODP Hole 318-U1356A

The early Eocene epoch was characterized by extreme global warmth, which in terrestrial settings was characterized by an expansion of near-tropical vegetation belts into the high latitudes. During the middle to late Eocene, global cooling caused the retreat of tropical vegetation to lower latitudes. In high-latitude settings, near-tropical vegetation was replaced by temperate floras. This floral change has recently been traced as far south as Antarctica, where along the Wilkes Land margin paratropical forests thrived during the early Eocene and temperate Nothofagus forests developed during the middle Eocene. Here we provide both qualitative and quantitative palynological data for this floral turnover based on a sporomorph record recovered at Integrated Ocean Drilling Program (IODP) Site U1356 off the Wilkes Land margin. Following the nearest living relative concept and based on a comparison with modern vegetation types, we examine the structure and diversity patterns of the Eocene vegetation along the Wilkes Land margin. Our results indicate that the early Eocene forests along the Wilkes Land margin were characterized by a diverse canopy composed of plants that today occur in tropical settings; their richness pattern was similar to that of present-day forests from New Caledonia. The middle Eocene forests were characterized by a canopy dominated by Nothofagus and exhibited richness patterns similar to modern Nothofagus forests from New Zealand.

(Table 1) n-alkane distribution, and stable carbon and deuterium isotope ratios of DSDP Hole 10-94

Hydrology, source region, and timing of precipitation are important controls on the climate of the Great Plains of North America and the composition of terrestrial ecosystems. Moisture delivered to the Great Plains varies seasonally and predominately derives from the Gulf of Mexico/Atlantic Ocean with minor contributions from the Pacific Ocean and Arctic region. For this work, we evaluate long-term relationships for the past ~ 35 million years between North American hydrology, climate, and floral change, using isotopic records and average carbon chain lengths of higher plant n-alkanes from Gulf of Mexico sediments (DSDP Site 94). We find that carbon isotope values (d13C) of n-alkanes, corrected for variations in the d13C value of atmospheric CO2, provide minor evidence for contributions of C4 plants prior to the Middle Miocene. A sharp spike in C4 input is identified during the Middle Miocene Climatic Optimum, and the influence of C4 plants steadily increased during the Late Miocene into the Pleistocene - consistent with other North American records. Chain-length distributions of n-alkanes, indicative of the composition of higher plant communities, remained remarkably constant from 33 to 4 Ma. However, a trend toward longer chain lengths occurred during the past 4 million years, concurrent with an increase in d13C values, indicating increased C4 plant influence and potentially aridity. The hydrogen isotope values (dD) of n-alkanes are relatively invariant between 33 and 9 Ma, and then become substantially more negative (75 per mil) from 9 to 2 Ma. Changes in the plant community and temperature of precipitation can solely account for the observed variations in dD from 33 to 5 Ma, but cannot account for Plio-Pleistocene dD variations and imply substantial changes in the source region of precipitation and seasonality of moisture delivery. We posit that hydrological changes were linked to tectonic and oceanographic processes including the shoaling and closure of the Panamanian Seaway, amplification of North Atlantic Deep Water Production and an associated increase of meridional winds. The southerly movement of the Intertropical Convergence Zone near 4 Ma allowed for the development of a near-modern pressure/storm track system, driving increased aridity and changes in seasonality within the North American interior.

Calcareous nannofossil stratigraphy, nannofossil events and carbon isotopic ratios of ODP Leg 183 sites

Cores from Sites 1135, 1136, and 1138 of Ocean Drilling Program Leg 183 to the Kerguelen Plateau (KP) provide the most complete Paleocene and Eocene sections yet recovered from the southern Indian Ocean. These nannofossil-foraminifer oozes and chalks provide an opportunity to study southern high-latitude biostratigraphic and paleoceanographic events, which is the primary subject of this paper. In addition, a stable isotope profile was established across the Cretaceous/Tertiary (K/T) boundary at Site 1138. An apparently complete K/T boundary was recovered at Site 1138 in terms of assemblage succession, isotopic signature, and reworking of older (Cretaceous) nannofossil taxa. There is a significant color change, a negative carbon isotope shift, and nannofossil turnover. The placement of the boundary based on these criteria, however, is not in agreement with the available shipboard paleomagnetic stratigraphy. We await shore-based paleomagnetic study to confirm or deny those preliminary results. The Paleocene nannofossil assemblage is, in general, characteristic of the high latitudes with abundant Chiasmolithus, Prinsius, and Toweius. Placed in context with other Southern Ocean sites, the biogeography of Hornibrookina indicates the presence of some type of water mass boundary over the KP during the earliest Paleocene. This boundary disappeared by the late Paleocene, however, when there was an influx of warm-water discoasters, sphenoliths, and fasciculiths. This not only indicates that during much of the late Paleocene water temperatures were relatively equable, but preliminary floral and stable isotope analyses also indicate that a relatively complete record of the late Paleocene Thermal Maximum event was recovered at Site 1135. It was only at the beginning of the middle Eocene that water temperatures began to decline and the nannofossil assemblage became dominated by cool-water species while discoaster and sphenolith abundances and diversity were dramatically reduced. One new taxonomic combination is proposed, Heliolithus robustus Arney, Ladner, and Wise.

Chemistry of Globigerinoides ruber shells from ODP Site 160-963

Here we present a high-resolution faunal, floral and geochemical (stable isotopes and trace elements) record from the sediments of Ocean Drilling Program Site 963 (central Mediterranean basin), which shows centennial/millennial-scale resemblance to the high-northern latitude rapid temperature fluctuations documented in the Greenland ice cores between 20 and 70 kyr BP. Oxygen and carbon isotopes, planktic foraminifera and calcareous nannofossil distributions suggest that Dansgaard-Oeschger (D/O) and Heinrich events (HE) are distinctly expressed in the Mediterranean climate record. Moreover, recurrent though subdued oscillations not previously identified in the Lateglacial Mediterranean sediments document a significant centennial-scale climate variability in the basin that is greater than previously thought. Alternations between climate regimes dominated by polar outbreaks during D/O stadials and warm D/O interstadials, with associated intensification of continental runoff, are well expressed in the ODP Site 963. These place the Mediterranean basin as an often overlooked recorder of the interplay between large- and regional- scale climate controls at intermediate latitudes, and of the possible interactions between different components of the climate system. Significant changes in Ba/Ca values measured in Globigerinoides ruber shells from a number of D/O stadials and interstadials suggest enhanced freshwater input from the north-eastern Mediterranean borderland during the D/O interstadials. However, the short duration of 3D stratification events never led to complete oxygen consumption along the water column, but clear effects of sluggish 3D circulation in the basin are testified to by negative excursions in d13C measured in selected species of planktic and benthic foraminifera. HEs are constantly associated with lightening in the d18O record of planktic foraminifera, possibly because of the impact of iceberg melting in the Iberian Margin on Mediterranean thermohaline circulation. Interestingly, in two cases in particular, HE2 and HE5, fresher water inputs also affected deeper horizons of intermediate waters, suggesting a basin-wide impact.

Haemolymph inorganic ion composition, physiology and behaviour dependent on temperature and ambient magnesium concentration in early life history stages of Paralomis granulosa (Decapoda, Anomura, Lithodidae)

Marine brachyuran and anomuran crustaceans are completely absent from the extremely cold (-1.8 °C) Antarctic continental shelf, but caridean shrimps are abundant. This has at least partly been attributed to low capacities for magnesium excretion in brachyuran and anomuran lithodid crabs ([Mg2+]HL = 20-50 mmol/L) compared to caridean shrimp species ([Mg2+]HL = 5-12 mmol/L). Magnesium has an anaesthetizing effect and reduces cold tolerance and activity of adult brachyuran crabs. We investigated whether the capacity for magnesium regulation is a factor that influences temperature-dependent activity of early ontogenetic stages of the Sub-Antarctic lithodid crab Paralomis granulosa. Ion composition (Na+, Mg2+, Ca2+, Cl-, [SO4]2-) was measured in haemolymph withdrawn from larval stages, the first and second juvenile instars (crabs I and II) and adult males and females. Magnesium excretion improved during ontogeny, but haemolymph sulphate concentration was lowest in the zoeal stages. Neither haemolymph magnesium concentrations nor Ca2+:Mg2+ ratios paralleled activity levels of the life stages. Long-term (3 week) cold exposure of crab I to 1 °C caused a significant rise of haemolymph sulphate concentration and a decrease in magnesium and calcium concentrations compared to control temperature (9 °C). Spontaneous swimming activity of the zoeal stages was determined at 1, 4 and 9 °C in natural sea water (NSW, [Mg2+] = 51 mmol/L) and in sea water enriched with magnesium (NSW + Mg2+, [Mg2+] = 97 mmol/L). It declined significantly with temperature but only insignificantly with increased magnesium concentration. Spontaneous velocities were low, reflecting the demersal life style of the zoeae. Heart rate, scaphognathite beat rate and forced swimming activity (maxilliped beat rate, zoea I) or antennule beat rate (crab I) were investigated in response to acute temperature change (9, 6, 3, 1, -1 °C) in NSW or NSW + Mg2+. High magnesium concentration reduced heart rates in both stages. The temperature-frequency curve of the maxilliped beat (maximum: 9.6 beats/s at 6.6 °C in NSW) of zoea I was depressed and shifted towards warmer temperatures by 2 °C in NSW + Mg2+, but antennule beat rate of crab I was not affected. Magnesium may therefore influence cold tolerance of highly active larvae, but it remains questionable whether the slow-moving lithodid crabs with demersal larvae would benefit from an enhanced magnesium excretion in nature.

Distribution of calcareous nannofossils in upper Cretaceous and Cenozoic sediments of the Kerguelen Plateau

ODP Leg 119 drilled 11 sites on the Kerguelen Plateau (southern Indian Ocean) and Prydz Bay (East Antarctica). Upper Pliocene through Quaternary sediments were recovered at Site 736 on the northern Kerguelen Plateau; calcareous nannofossils occurred in only a few samples. Over 700 m of middle Eocene through Quaternary sediments was cored at Site 737 on the northern Kerguelen Plateau, and calcareous nannofossils are abundant in the middle Eocene through the middle Miocene sediments. Nearly 500 m of sediments ranging from the lower Turanian to the Quaternary was recovered at Site 738 on the southern Kerguelen Plateau; calcareous nannofossils are abundant from the Miocene downward. Calcareous nannofossils are also abundant in the upper Eocene through Miocene section from Site 744 on the southern Kerguelen Plateau. Except for Core 119-746A-13H, the Neogene sequences drilled at deep-water Sites 745 and 746 off the southern Kerguelen Plateau are devoid of calcareous nannofossils. Occurrences of calcareous nannofossils were generally rare and sporadic at Sites 739 and 742 in Prydz Bay and suggest that the diamictite sequences recovered is as old as middle Eocene-early Oligocene age. Other sites drilled in Prydz Bay (Sites 740, 741, and 743) did not yield calcareous nannofossils. Species diversity of calcareous nannofossils was low (about a dozen) in the southern Indian Ocean in the Late Cretaceous. High-latitude nanno floral characteristics are apparent after the Cretaceous/Tertiary boundary extinctions. Cold climatic conditions limited Oligocene calcareous nannofossil assemblages to fewer than a dozen species, and extinctions of species generally were not compensated by originations of new species. Only a few species of calcareous nannofossils were found in the Miocene sequences, in which Coccolithuspelagicus and one or two species of Reticulofenestra exhibit extreme (0%-100%) fluctuations in assemblage dominance, and these fluctuations may reflect rapid fluctuations in the surface-water temperatures. Further deterioration of climate in the late Neogene essentially excluded calcareous nannoplankton from the Southern Ocean. Significantly warmer water conditions during part of the early-middle Pleistocene were inferred by a few lower-middle Pleistocene calcareous nannofossil species found on the Kerguelen Plateau. The calcareous nannofossil zonation of Roth (1978 doi:10.2973/dsdp.proc.44.134.1978) can be applied to the Upper Cretaceous section recovered at Site 738, and the zonation of Okada and Bukry (1980 doi:10.1016/0377-8398(80)90016-X) can be applied without much difficulty to the Paleocene to middle Eocene sequences from the Kerguelen Plateau. However, some conventional upper Paleogene markers are not useful for southern high latitudes, whereas a few nonconventional species events are useful for subdividing the upper Paleogene sequences. The latter species events include the first occurrence (FO) of Reticulofenestra reticulata, the FO and last occurrence (LO) of Reticulofenestra oamaruensis, the LO of Isthmolithus recurvus, and the LO of Chiasmolithus altus. As the Neogene sequences from the southern Indian Ocean contain only a few long-ranging, cold-water species, or are devoid of coccoliths, calcareous nannofossil zonations remain virtually unworkable for the Neogene in the high-latitude southern Indian Ocean as in other sectors of the Southern Ocean.

Oxygen isotope values from biogenic apatie (conodont elements and fish teeth) from the Lower Triassic Mianwali Formation (Salt Range, Pakistan)

Recovery from the end-Permian mass extinction is frequently described as delayed, with complex ecological communities typically not found in the fossil record until the Middle Triassic epoch. However, the taxonomic diversity of a number of marine groups, ranging from ammonoids to benthic foraminifera, peaked rapidly in the Early Triassic. These variations in biodiversity occur amidst pronounced excursions in the carbon isotope record, which are compatible with episodes of massive CO2 outgassing from the Siberian Large Igneous Province. Here we present a high-resolution Early Triassic temperature record based on the oxygen isotope composition of pristine apatite from fossil conodonts. Our reconstruction shows that the beginning of the Smithian substage of the Early Triassic was marked by a cooler climate, followed by an interval of warmth lasting until the Spathian substage boundary. Cooler conditions resumed in the Spathian. We find the greatest increases in taxonomic diversity during the cooler phases of the early Smithian and early Spathian. In contrast, a period of extreme warmth in the middle and late Smithian was associated with floral ecological change and high faunal taxonomic turnover in the ocean. We suggest that climate upheaval and carbon-cycle perturbations due to volcanic outgassing were important drivers of Early Triassic biotic recovery.

Mg/Ca ratios in planktonic foraminifera

The primary Mg/Ca ratio of foraminiferal shells is a potentially valuable paleoproxy for sea surface temperature (SST) reconstructions. However, the reliable extraction of this ratio from sedimentary calcite assumes that we can overcome artifacts related to foraminiferal ecology and partial dissolution, as well as contamination by secondary calcite and clay. The standard batch method for Mg/Ca analysis involves cracking, sonicating, and rinsing the tests to remove clay, followed by chemical cleaning, and finally acid-digestion and single-point measurement. This laborious procedure often results in substantial loss of sample (typically 30-60%). We find that even the earliest steps of this procedure can fractionate Mg from Ca, thus biasing the result toward a more variable and often anomalously low Mg/Ca ratio. Moreover, the more rigorous the cleaning, the more calcite is lost, and the more likely it becomes that any residual clay that has not been removed by physical cleaning will increase the ratio. These potentially significant sources of error can be overcome with a flow-through (FT) sequential leaching method that makes time- and labor-intensive pretreatments unnecessary. When combined with time-resolved analysis (FT-TRA) flow-through, performed with a gradually increasing and highly regulated acid strength, produces continuous records of Mg, Sr, Al, and Ca concentrations in the leachate sorted by dissolution susceptibility of the reacting material. Flow-through separates secondary calcite from less susceptible biogenic calcite and clay, and further resolves the biogenic component into primary and more resistant fractions. FT-TRA reliably separates secondary calcite (which is not representative of original life habitats) from the more resistant biogenic calcite (the desired signal) and clay (a contaminant of high Mg/Ca, which also contains Al), and further resolves the biogenic component into primary and more resistant fractions that may reflect habitat or other changes during ontogeny. We find that the most susceptible fraction of biogenic calcite in surface dwelling foraminifera gives the most accurate value for SST and therefore best represents primary calcite. Sequential dissolution curves can be used to correct the primary Mg/Ca ratio for clay, if necessary. However, the temporal separation of calcite from clay in FT-TRA is so complete that this correction is typically <=2%, even in clay-rich sediments. Unlike hands-on batch methods, that are difficult to reproduce exactly, flow-through lends itself to automation, providing precise replication of treatment for every sample. Our automated flow-through system can process 22 samples, two system blanks, and 48 mixed standards in <12 hours of unattended operation. FT-TRA thus represents a faster, cheaper, and better way to determine Mg/Ca ratios in foraminiferal calcite.

Danian calcareous nannofossils in ODP Hole 119-738C

An apparently complete Danian section was recovered at ODP Site 738 on the southern Kerguelen Plateau. Calcareous nannofossils are abundant and moderately preserved in the section. A number of taxa common in middle or low latitudes, such as Braarudosphaera, Biscutum? romeinii, Biscutum? parvulum, Cyclagelosphaera, Octolithus multiplus, and Toweius petalosus are absent at Site 738. On the other hand, a bloom of Hornibrookina occurs at Site 738 only slightly (15 cm) above the Cretaceous/Tertiary boundary as defined by the iridium peak. Species of Chiasmolithus and Prinsius are very abundant. This gives the nannofossil assemblages distinct high-latitude characteristics and suggests significant latitudinal thermal gradients in the Danian oceans. A Danian nannofossil zonation for the Antarctic region is proposed, which utilizes traditional markers and several nontraditional markers, i.e., the first occurrences of Hornibrookina, Prinsius martinii, and Chiasmolithus bidens, and the last occurrence of Hornibrookina teuriensis. Quantitative analyses of the calcareous nannofossil assemblages from Site 738 reveal four steps of rapid floral changes in the early Danian before relatively stable nannofloral conditions were reached at about 63.8 Ma.