20 movies of x-ray tomograms of different Alpinia fossils

The fossil floras described by Dieter MAI and Harald WALTHER are invaluable for understanding the past plant diversity in Europe, and provide important information on the occurrence of taxa in the fossil record that is critical for evolutionary studies. Among the taxa they recognized were seeds assigned to the extant genus Alpinia ROXB. (Zingiberaceae, Zingiberales). We reinvestigated 28 specimens that were assigned to Alpinia arnensis (CHANDLER) MAI, Alpinia cf. arnensis, and Alpinia bivascularis MAI from the Ypresian (lower Eocene) of the UK, upper Eocene of Germany, and lower Miocene of Germany using non-destructive synchrotron-based X-ray tomography to reveal internal anatomy. None of the samples studied show an anatomy consistent with extant Alpinia or even Zingiberales. The fossils lack the globose shape, often striate external surface, seed coat structure, operculum, and micropylar collar seen in all Alpinia, and lack the chalazal chamber seen in many Alpinia species. Two specimens from the lower Miocene of Germany showed the structure of fruits of Caricoidea CHANDLER (Cyperaceae) with a single-layered exocarp, thick mesocarp, and sclerified endocarp. The other specimens are recognized as Carpolithes albolutum nom. nov. (incertae sedis) from the Ypresian of the UK, C. phoenixnordensis sp. nov. (incertae sedis) from the upper Eocene of Germany, C. bivascularis comb. nov. (incertae sedis) from the lower Miocene of Germany as well as indeterminate tegmens from the lower Miocene of Germany. This reinvestigation demonstrates that there is, as yet, no confirmed fossil record for the extant genus Alpinia. Furthermore, at least four different taxa are recognized from what had been two extinct species, enhancing our understanding of these important European Cenozoic carpofloras.

Morphology of Orbulina universa and Globorotalia scitula across local extinctions during Sapropel S5 (Eastern Mediterranean Sea, c.126-121 ka)

Extinction is a remarkably difficult phenomenon to study under natural conditions. This is because the outcome of stress exposure and associated fitness reduction is not known until the extinction occurs and it remains unclear whether there is any phenotypic reaction of the exposed population that can be used to predict its fate. Here we take advantage of the fossil record, where the ecological outcome of stress exposure is known. Specifically, we analyze shell morphology of planktonic Foraminifera in sediment samples from the Mediterranean, during an interval preceding local extinctions. In two species representing different plankton habitats, we observe shifts in trait state and decrease in variance in association with non-terminal stress, indicating stabilizing selection. At terminal stress levels, immediately before extinction, we observe increased growth asymmetry and trait variance, indicating disruptive selection and bet-hedging. The pre-extinction populations of both species show a combination of trait states and trait variance distinct from all populations exposed to non-terminal levels of stress. This finding indicates that the phenotypic history of a population may allow the detection of threshold levels of stress, likely to lead to extinction. It is thus an alternative to population dynamics in studying and monitoring natural population ecology.

Planktonic Foraminifera shell calcification intensity from sediment trap L1/K276 from the Azores Front (2002/2003)

Using shells collected from a sediment trap series in the Madeira Basin, we investigate the effects of seasonal variation of temperature, productivity, and optimum growth conditions on calcification in three species of planktonic Foraminifera. The series covers an entire seasonal cycle and reflects conditions at the edge of the distribution of the studied species, manifesting more suitable growth conditions during different parts of the year. The seasonal variation in seawater carbonate saturation at the studied site is negligible compared to other oceanic regions, allowing us to assess the effect of parameters other than carbonate saturation. Shell calcification is quantified using weight and size of individual shells. The size-weight scaling within each species is robust against changes in environmental parameters, but differs among species. An analysis of the variation in calcification intensity (size-normalized weight) reveals species-specific response patterns. In Globigerinoides ruber (white) and Globigerinoides elongatus, calcification intensity is correlated with temperature (positive) and productivity (negative), whilst in Globigerina bulloides no environmental forcing is observed. The size-weight scaling, calcification intensity, and response of calcification intensity to environmental change differed between G. ruber (white) and G. elongatus, implying that patterns extracted from pooled analyses of these species may reflect their changing proportions in the samples. Using shell flux as a measure of optimum growth conditions, we observe significant positive correlation with calcification intensity in G. elongatus, but negative correlation in G. bulloides. The lack of a consistent response of calcification intensity to optimum growth conditions is mirrored by the results of shell size analyses. We conclude that calcification intensity in planktonic Foraminifera is affected by factors other than carbonate saturation. These factors include temperature, productivity, and optimum growth conditions, but the strength and sign of the relationships differ among species, potentially complicating interpretations of calcification data from the fossil record.

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.

Planktonic foraminiferal faunas and stable isotope ratios of sediments from the Benguela Upwelling System

Two sediment cores retrieved from the continental slope in the Benguela Upwelling System, GeoB 1706 (19°33.7'S 11°10.5'E) and GeoB 1711 (23°18.9'S, 12°22.6'E), reveal striking variations in planktonic foraminiferal abundances during the last 160,000 years. These fluctuations are investigated to assess changes in the intensity and position of the upwelling centres off Namibia. Four species make up over 95% of the variation within the core, and enable the record to be divided into episodes characterized by particular planktonic foraminiferal assemblages. The fossil assemblages have meaningful ecological significance when compared to those of the modern day and the relationship to their environment. The cold-water planktonic foraminifer, Neogloboquadrina pachyderma sinistral [N. pachyderma (s)], dominates the modern-day, coastal upwelling centres, and Neogloboquadrina pachyderma dextral and Globigerina bulloides characterize the fringes of the upwelling cells. Globorotalia inflata is representative of the offshore boundary between newly upwelled waters and the transitional, reduced nutrient levels of the subtropical waters. In the fossil record, episodes of high N. pachyderma (s) abundances are interpreted as evidence of increased upwelling intensity, and the associated increase in nutrients. The N. pachyderma (s) record suggests temporal shifts in the intensity of upwelling, and corresponding trophic domains, that do not follow the typical glacial-interglacial pattern. Periods of high N. pachyderma (s) abundance describe rapid, discrete events dominating isotope stages 3 and 2. The timing of these events correlates to the temporal shifts of the Angola-Benguela Front (Jansen et al., 1997) situated to the north of the Walvis Ridge. Absence of high abundances of N. pachyderma (s) from the continental slope of the southern Cape Basin indicates that Southern Ocean surface water advection has not exerted a major influence on the Benguela Current System. The coincidence of increased upwelling intensity with the movement of the Angola-Benguela Front can be interpreted mainly by changes in strength and zonality of the trade wind system.

Foraminifera species in the CO2 vents near Ischia, Italy, 2010

The seas around the island of Ischia (Italy) have a lowered pH as a result of volcanic gas vents that emit carbon dioxide from the sea floor at ambient seawater temperatures. These areas of acidified seawater provide natural laboratories in which to study the long-term biological response to rising CO2 levels. Benthic foraminifera (single-celled protists) are particularly interesting as they have short life histories, are environmentally sensitive and have an excellent fossil record. Here, we examine changes in foraminiferal assemblages along pH gradients at CO2 vents on the coast of Ischia and show that the foraminiferal distribution, diversity and nature of the fauna change markedly in the living assemblages as pH decreases.

Paleoclimate reconstruction from Miocene macroflora in Kazakhstan compiled from various publications

25 datasets (13 fossil leaf and pollen assemblages, 12 quantitative palaeoclimatic datasets) are provided in order to analyse Early Miocene palaeoclimate in Kazakhstan. The rich fossil record in Kazakhstan documents that during the Oligocene and Early Miocene this area in Central Eurasia was densely forested with warm-temperate deciduous trees and shrubs of the so-called "Turgayan flora". 29 fossil floras from 13 localities have been selected for a quantitative analysis of the Aquitanian (early Early Miocene) climate situation in Kazakhstan. The assessed mean annual temperatures generally place around 15 °C, while values of mean annual precipitation are of about 1000 mm. In combination with several other climate parameters estimated (temperatures of warmest and coldest months, precipitation rates of wettest, driest and warmest months), these data reflect uniform climatic conditions over several thousands of square kilometres. Data of temperature parameters show slight spatial differentiations, with generally cooler mean annual temperatures and higher seasonality (i.e. warmer summers and colder winters) in the north-eastern part of the study area compared with the south-western area around Lake Aral. As compared with palaeoclimate estimates for the European and East Asian Aquitanian, the central part of the Eurasian continent reveals evident signals of higher seasonality and slightly increased continentality.

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.

(Supplementary Material 2) Landmark coordinates of the analyzed specimens with associated metadata

The use of planktonic foraminifera in paleoceanographic studies relies on the assumption that morphospecies represent biological species with ecological preferences that are stable through time and space. However, genetic surveys unveiled a considerable level of diversity in most morphospecies of planktonic foraminifera. This diversity is significant for paleoceanographic applications because cryptic species were shown to display distinct ecological preferences that could potentially help refine paleoceanographic proxies. Subtle morphological differences between cryptic species of planktonic foraminifera have been reported, but so far their applicability within paleoceanographic studies remains largely unexplored. Here we show how information on genetic diversity can be transferred to paleoceanography using Globorotalia inflata as a case study. The two cryptic species of G. inflata are separated by the Brazil-Malvinas Confluence (BMC), a major oceanographic feature in the South Atlantic. Based on this observation, we developed a morphological model of cryptic species detection in core top material. The application of the cryptic species detection model to Holocene samples implies latitudinal oscillations in the position of the confluence that are largely consistent with reconstructions obtained from stable isotope data. We show that the occurrence of cryptic species in G. inflata, can be detected in the fossil record and used to trace the migration of the BMC. Since a similar degree of morphological separation as in G. inflata has been reported from other species of planktonic foraminifera, the approach presented in this study can potentially yield a wealth of new paleoceanographical proxies.

Benthic foraminifera of the Red Sea

The distribution of living (Rose Bengal-stained), dead and fossil benthic foraminifera was investigated in six short cores (multicores, 30-32 cm total length) recovered from the central Red Sea. The ecological preferences as well as the relationship between the live and dead/fossil assemblages (preserved down-core) were examined. The sites, located along a W-E profile and between the depth of 366 and 1782 m, extend from the center of the oxygen minimum zone (OMZ, ~200-650 m), through its margin at ~600 m, and down to the well-aerated deep-water environment. Live (Rose-Bengal stained) and coexisting dead foraminifera were studied in the upper 5 cm of each of the sites, and the fossil record was studied down to ~32 cm. Q-mode Principal Component Analysis was used and four distinct foraminiferal fossil assemblages were determined. These assemblages follow different water mass properties. In the center of the OMZ, where the organic carbon content is highest and the oxygen concentration is lowest (<=0.5 ml O2/l), the Bolivina persiensis-Bulimina marginata-Discorbinella rhodiensis assemblage dominates. The slightly more aerated and lower organic-carbon-content seafloor, at the margin of the OMZ, is characterized by the Neouvigerina porrecta-Gyroidinoides cf. G. soldanii assemblage. The transitional environment, between 900-1200 m, with its well-aerated and oligotrophic seafloor, is dominated by the Neouvigerina ampullacea-Cibicides mabahethi assemblage. The deeper water (>1500 m), characterized by the most oxygenated and oligotrophic seafloor conditions, is associated with the Astrononion sp. A-Hanzawaia sp. A assemblage. Throughout the Red Sea extremely high values of temperature and salinity are constant below ~200 m depth, but the flux of organic matter to the sea floor varies considerably with bathymetry and appears to be the main controlling factor governing the distribution pattern of the benthic foraminifera. Comparison between live and the dead/fossil assemblages reveals a large difference between the two. Processes that may control this difference include species-specific high turnover rates, and preferential predation and loss of fragile taxa (either by chemical or microbial processes). Significant variations in the degree of loss of the organic-cemented agglutinants were observed down core. This group is preserved down to 5-10 cm at the shallow OMZ sites and down to greater depths at well-aerated and oligotrophic sites. The lower rate of disintegration of these forms, in the deeper locations of the Red Sea, may be related to low microbial activity. This results in the preservation of increasing numbers of organic-cemented shells down-core.

Fidelity bias in mollusk assemblages from coastal lagoons of Southern Brazil

The South America southern coast exhibits many outcrops with abundant shell beds, from the Pleistocene through the Recent. How much biological information is preserved within these shell beds? Or, what is the actual probability a living community has to leave a fossil record corresponding to these shell deposits? Although ecological and biogeographical aspects might had been pointed, considering these temporal scales, up to the moment there is no taphonomically-oriented studies available. Quantitative comparisons between living (LAs), death (DAs) and fossil assemblages (FAs) are important not only in strictly taphonomic studies, but have grown a leading tool for conservation paleobiology analysis. Comparing LAs, DAs and FAs from estuaries and lagoons in the Rio Grande do Sul Coastal Plain makes possible to quantitatively understand the nature and quantity of biological information preserved in fossil associations in Holocene lagoon facies. As already noted by several authors, spatial scale parts the analysis, but we detected that the FAs refl ects live ones, rather than dead ones, as previously not realized. The results herein obtained illustrates that species present in DA are not as good preserved in recent (Holocene) fossil record as originally thought. Strictly lagoon species are most prone to leave fossil record. The authors consider that the fi delity pattern here observed for estuarine mollusks to be driven by (i) high temporal and spatial variability in the LAs, (ii) spatial mixing in the DA and (iii) differential preservation of shells, due to long residence times in the taphonomically active zone.

(Table 1) Sample descriptions, carbon contents, and lipid yields across the K/T boundary at DSDP Hole 93-605 and at Stevns Klint, Denmark

This reconnaissance study was undertaken to determine whether the mass extinctions and faunal successions that mark the Cretaceous/Tertiary (K/T) boundary left a discernible molecular fossil record in the sediments of this period. Lipid signatures of sediments taken from above and below the K/T boundary were compared in core and outcrop samples taken from two locations: the U.S. east coast continental margin (western Atlantic Ocean, DSDP Site 605) and Stevns Klint, Denmark. Four calcareous sediments taken from above and below the K/T boundary in DSDP Hole 605, Section 605-66-1, revealed changing lipid signatures between above and below that are characterized by a large component of unresolved naphthenic hydrocarbons and a homologous series of n-alkanes ranging from Ci6 to C33. These lipid signatures are attributed to an influx of a terrestrial higher plant component and to bacterial reworking of the sediments under partially anoxic depositional and/or diagenetic conditions. The outcrop samples from Stevns Klint had extremely low concentrations of indigenous lipids. The fish clay at the K/T boundary contained traces of microbial hydrocarbons and fatty acids, whereas the carbonates above and below had only microbial fatty acids and additional terrestrial resin acids. The data from both sites indicate a perturbation in the deposition of lipid compound classes across the K/T boundary.

Coccosphere geometry measurements from culture experiments on the coccolithophore species Calcidiscus leptoporus, Calcidiscus quadriperforatus and Helicosphaera carteri

Coccolithophores are a key phytoplankton group that exhibit remarkable diversity in their biology, ecology, and calcitic exoskeletons (coccospheres). An understanding of the physiological processes that underpin coccosphere architecture is essential for maximizing the information that can be retrieved from their extensive fossil record. Using culturing experiments on four modern species from three long-lived families, we investigate how coccosphere architecture responds to population shifts from rapid (exponential) to slowed (stationary) growth phases as nutrients become depleted. These experiments reveal statistical differences in cell size and the number of coccoliths per cell between these two growth phases, specifically that cells in exponential-phase growth are typically smaller with fewer coccoliths, whereas cells experiencing growth-limiting nutrient depletion have larger coccosphere sizes and greater numbers of coccoliths per cell. Although the exact numbers are species-specific, these growth-phase shifts in coccosphere geometry are common to four different coccolithophore families (Calcidiscaceae, Coccolithaceae, Isochrysidaceae, Helicosphaeraceae), demonstrating that this is a core physiological response to nutrient depletion across a representative diversity of this phytoplankton group. Polarised light microscopy was used for all coccosphere geometry measurements.

Carbon and oxygen isotope ratios of Selinde River, southeastern Siberian platform

Carbon isotopic data from the Selinde section in the southeastern part of the Siberian platform area are correlated with the reference isotopic profile from the Lower Cambrian stratotype sections of the Lena Aldan region, but also show additional d13C excursions unrecognized there. The chemostratigraphic correlation suggests that the geological and fossil record of the lower Pestrotsvet Formation in the Selinde section has a deeper history than the stratotype region. This conclusion is important for both constraining the age of the earliest Cambrian marine transgression on the Siberian platform and providing a clearer understanding of the pace and order of early Cambrian geochemical and biological events.

Fossil pollen record of composite sediment core TMD from Tso Moriri, analysis of modern surface pollen samples, mean annual precipitation reconstruction, and digitisation and recalibration of different discussed palaeoclimate proxy records

This paper presents a new fossil pollen record from Tso Moriri (32°54'N, 78°19'E, 4512 m a.s.l.) and seeks to reconstruct changes in mean annual precipitation (MAP) during the last 12,000 years. This high-alpine lake occupies an area of 140 km**2 in a glacial-tectonic valley in the northwestern Himalaya. The region has a cold climate, with a MAP <300 mm, and open vegetation. The hydrology is controlled by the Indian Summer Monsoon (ISM), but winter westerly-associated precipitation also affects the regional water balance. Results indicate that precipitation levels varied significantly during the Holocene. After a rapid increase in MAP, a phase of maximum humidity was reached between ca. 11 to 9.6 cal ka BP, followed by a gradual decline in MAP. This trend parallels the reduction in the Northern Hemisphere summer insolation. Comparison of different palaeoclimate proxy records reveal evidence for a stronger Holocene decrease in precipitation in the northern versus the southern parts of the ISM domain. The long-term trend of ISM weakening is overlaid with several short periods of greater dryness, which are broadly synchronous with the North Atlantic cold spells, suggesting reduced amounts of westerly-associated winter precipitation. Compared to the mid and late Holocene, it appears that westerlies had a greater influence on the western parts of the ISM domain during the early Holocene. During this period, the westerly-associated summer precipitation belt was positioned at Mediterranean latitudes and amplified the ISM-derived precipitation. The Tso Moriri pollen record and moisture reconstructions also suggest that changes in climatic conditions affected the ancient Harappan Civilisation, which flourished in the greater Indus Valley from approximately 5.2 to 3 cal ka BP. The prolonged Holocene trend towards aridity, punctuated by an interval of increased dryness (between ca. 4.5 to 4.3 cal ka BP), may have pushed the Mature Harappan urban settlements (between ca. 4.5 to 3.9 cal ka BP) to develop more efficient agricultural practices to deal with the increasingly acute water shortages. The amplified aridity associated with North Atlantic cooling between ca. 4 to 3.6 and around 3.2 cal ka BP further hindered local agriculture, possibly causing the deurbanisation that occurred from ca. 3.9 cal ka BP and eventual collapse of the Harappan Civilisation between ca. 3.5 to 3 cal ka BP.