Benthic foraminifera in Tertiary sediments of DSDP Leg 90 holes

Eocene through Pliocene benthic foraminifers were examined from seven sites located at middle and lower bathyal depths on the Lord Howe Rise in the Tasman Sea, from another site at lower bathyal depths in the Coral Sea, and from a site in the intermediate-depth, hemipelagic province of the Chatham Rise, east of southern New Zealand. Age-related, depth-related, and bioprovincial faunal variations are documented in this chapter. One new species, Rectuvigerina tasmana, is named. The paleoecologic indications of several key groups, including the miliolids, uvigerinids, nuttallitids, and cibicidids, are combined with sedimentologic and stable isotopic tracers to interpret paleoceanographic changes in the Tasman Sea. Because the total stratigraphic ranges of many bathyal benthic foraminifers are not yet known, most endpoints in the Tasman Sea are considered ecologically controlled events. The disappearances of Uvigerina rippensis and Cibicidoidesparki and the first appearances of U. pigmaea, Sphaeroidina bulloides, and Rotaliatina sulcigera at the Eocene/Oligocene boundary can be considered evolutionary events, as also can the first appearance of Cibicides wuellerstorfi in Zone NN5. Species which are restricted to the lower bathyal zone except during discrete pulses, most of which are related to the development of glacial conditions, include Melonis pompilioides, M. sphaeroides, Pullenia quinqueloba, Nuttallides umbonifera, and U. hispido-costata. Middle bathyal indigenes include U. spinulosa, U. gemmaeformis, Ehrenbergina marwicki, R. sulcigera, and all rectuvigerinids except Rectuvigerina spinea. Although the miliolids first occurred at lower bathyal depths, they were more common in the middle bathyal zone. Although the Neogene hispido-costate uvigerinids first developed at lower bathyal depths and at higher middle latitude sites, in the later Neogene this group migrated to shallower depths and became predominant also in the middle bathyal zone. Despite the relatively similar sedimentologic settings at the six middle bathyal Tasman sites, there was extensive intrageneric and intraspecific geographic variation. Mililiolids, strongly ornamented brizalinids, bolivinitids, Bulimina aculeata, Osangularia culter, and strongly porous morphotypes were more common at higher latitudes. Osangularia bengalensis, striate brizalinids such as Brizalina subaenariensis, Gaudryina solida, osangularids in general, and finely porous morphotypes were more common in the subtropics. There was strong covariance between faunas at lower middle latitude, lower bathyal Site 591, and higher middle latitude, middle bathyal Site 593. The following oceanographic history of the Tasman Sea is proposed; using the stable isotopic record as evidence for glacials and examining the ecologic correlations between (1) miliolids and carbonate saturation, (2) nuttallitids and undersaturated, cooled, or "new" water masses, (3) uvigerinids with high organic carbon in the sediment and high rates of sediment accumulation, and (4) cibicidids and terrestrial organic carbon. The glacial located near the Eocene/Oligocene boundary is characterized by the penetration of cooler, more corrosive waters at intermediate depths in high southern latitudes. This may have caused overturn, upwelling pulses, in other Tasman areas. The development of Neogenelike conditions began in the late Oligocene (Zone NP24/NP25) with the evolution of several common Neogene species. A large number of Paleogene benthics disappeared gradually through the course of the early Miocene, which was not well preserved at any Tasman site. Corrosive conditions shallowed into the middle bathyal zone in several pulses during the early Miocene. The development of glacial conditions in the middle Miocene was accompanied by major changes throughout the Tasman Sea. Sediment accumulation rates increased and high-productivity faunas and corrosive conditions developed at all but the lowest-latitude Site 588. This increase in productivity and accumulation rate is attributed to the eutrophication of Antarctic water masses feeding Tasman current systems, as well as to invigorated circulation in general. It overlaps with the beginning of the Pacific High-productivity Episode (10-5 Ma). During the latest Miocene glacial episode, corrosive conditions developed at lower bathyal depths, while cooler water and lower nutrient levels shallowed to middle bathyal depths. Lower input of terrestrial organic carbon may be related to the lower nutrient levels of this time and to the termination of the Pacific High-productivity Episode. The moderate glacial episode during the mid-Pliocene (Zone NN15/NN16, ~3.2 Ma) corresponds to a decline in sediment accumulation rates and a reorganization of faunas unlike that of all other times. New genera proliferate and indices for cool, noncorrosive conditions and high organic carbon expand throughout the middle bathyal zone coeval with the sedimentation rate decreases. By the latest Pliocene (about 2.5 Ma), however, during another glacial episode, faunal patterns typical of this and later glacials develop throughout the Tasman Sea. Benthic foraminiferal patterns suggest increased input of terrestrial organic matter to Tasman Sea sediments during this episode and during later glacials.

Have we been underestimating the effects of ocean acidification in zooplankton?

Understanding how copepods may respond to ocean acidification (OA) is critical for risk assessments of ocean ecology and biogeochemistry. The perception that copepods are insensitive to OA is largely based on experiments with adult females. Their apparent resilience to increased carbon dioxide (pCO2) concentrations has supported the view that copepods are 'winners' under OA. Here, we show that this conclusion is not robust, that sensitivity across different life stages is significantly misrepresented by studies solely using adult females. Stage-specific responses to pCO2 (385-6000 µatm) were studied across different life stages of a calanoid copepod, monitoring for lethal and sublethal responses. Mortality rates varied significantly across the different life stages, with nauplii showing the highest lethal effects; nauplii mortality rates increased threefold when pCO2 concentrations reached 1000 µatm (year 2100 scenario) with LC50 at 1084 µatm pCO2. In comparison, eggs, early copepodite stages, and adult males and females were not affected lethally until pCO2 concentrations >= 3000 µatm. Adverse effects on reproduction were found, with >35% decline in nauplii recruitment at 1000 µatm pCO2. This suppression of reproductive scope, coupled with the decreased survival of early stage progeny at this pCO2 concentration, has clear potential to damage population growth dynamics in this species. The disparity in responses seen across the different developmental stages emphasizes the need for a holistic life-cycle approach to make species-level projections to climate change. Significant misrepresentation and error propagation can develop from studies which attempt to project outcomes to future OA conditions solely based on single life history stage exposures.

Seawater carbonate chemistry, clearance rates, respiration rates, condition index and cellular turnover (RNA: DNA) of Juvenile King Scallop, Pecten maximus in a laboratory experiment

The decline in ocean water pH and changes in carbonate saturation states through anthropogenically mediated increases in atmospheric CO2 levels may pose a hazard to marine organisms. This may be particularly acute for those species reliant on calcareous structures like shells and exoskeletons. This is of particular concern in the case of valuable commercially exploited species such as the king scallop, Pecten maximus. In this study we investigated the effects on oxygen consumption, clearance rates and cellular turnover in juvenile P. maximus following 3 months laboratory exposure to four pCO2 treatments (290, 380, 750 and 1140 µatm). None of the exposure levels were found to have significant effect on the clearance rates, respiration rates, condition index or cellular turnover (RNA: DNA) of individuals. While it is clear that some life stages of marine bivalves appear susceptible to future levels of ocean acidification, particularly under food limiting conditions, the results from this study suggest that where food is in abundance, bivalves like juvenile P. maximus may display a tolerance to limited changes in seawater chemistry.

Offspring sensitivity to ocean acidification changes seasonally in a coastal marine fish

Experimental assessments of species vulnerabilities to ocean acidification are rapidly increasing in number, yet the potential for short- and long-term adaptation to high CO2 by contemporary marine organisms remains poorly understood. We used a novel experimental approach that combined bi-weekly sampling of a wild, spawning fish population (Atlantic silverside Menidia menidia) with standardized offspring CO2 exposure experiments and parallel pH monitoring of a coastal ecosystem. We assessed whether offspring produced at different times of the spawning season (April to July) would be similarly susceptible to elevated (1100 µatm, pHNIST = 7.77) and high CO2 levels (2300 µatm, pHNIST = 7.47). Early in the season (April), high CO2 levels significantly (p < 0.05) reduced fish survival by 54% (2012) and 33% (2013) and reduced 1 to 10 d post-hatch growth by 17% relative to ambient conditions. However, offspring from parents collected later in the season became increasingly CO2-tolerant until, by mid-May, offspring survival was equally high at all CO2 levels. This interannually consistent plasticity coincided with the rapid annual pH decline in the species' spawning habitat (mean pH: 1 April/31 May = 8.05/7.67). It suggests that parents can condition their offspring to seasonally acidifying environments, either via changes in maternal provisioning and/or epigenetic transgenerational plasticity (TGP). TGP to increasing CO2 has been shown in the laboratory but never before in a wild population. Our novel findings of direct CO2-related survival reductions in wild fish offspring and seasonally plastic responses imply that realistic assessments of species CO2-sensitivities must control for parental environments that are seasonally variable in coastal habitats.

Temperature is the evil twin: Effects of increased temperature and ocean acidification on reproduction in a reef fish

Reproduction in many organisms can be disrupted by changes to the physical environment, such as those predicted to occur during climate change. Marine organisms face the dual climate change threats of increasing temperature and ocean acidification, yet no studies have examined the potential interactive effects of these stressors on reproduction in marine fishes. We used a long-term experiment to test the interactive effects of increased temperature and CO2 on the reproductive performance of the anemonefish, Amphiprion melanopus. Adult breeding pairs were kept for 10 months at three temperatures, 28.5°C (+0.0°C), 30.0°C (+1.5°C) and 31.5°C (+3.0°C), cross-factored with 3 CO2 levels, a current day control (417 µatm) and moderate (644 µatm) and high (1134 µatm) treatments consistent with the range of CO2 projections for the year 2100 under RCP8.5. We recorded each egg clutch produced during the breeding season, the number of eggs laid per clutch, average egg size, fertilization success, survival to hatching, hatchling length and yolk provisioning. Adult body condition, hepatosomatic index, gonadosomatic index, and plasma 17beta-estradiol concentrations were measured at the end of the breeding season to determine the effect of prolonged exposure to increased temperature and elevated CO2 on adults, and to examine potential physiological mechanisms for changes in reproduction. Temperature had by far the stronger influence on reproduction, with clear declines in reproduction occurring in the +1.5°C treatment and ceasing altogether in the +3.0°C treatment. In contrast, CO2 had a minimal effect on the majority of reproductive traits measured, but caused a decline in offspring quality in combination with elevated temperature. We detected no significant effect of temperature or CO2 on adult body condition or hepatosomatic index. Elevated temperature had a significant negative effect on plasma 17beta-estradiol concentrations, suggesting that declines in reproduction with increasing temperature were due to the thermal sensitivity of reproductive hormones rather than a reduction in energy available for reproduction. Our results show that elevated temperature exerts a stronger influence than high CO2 on reproduction in A. melanopus. Understanding how these two environmental variables interact to affect the reproductive performance of marine organisms will be important for predicting the future impacts of climate change.

Microenvironmental changes support evidence of photosynthesis and calcification inhibition in Halimeda under ocean acidification and warming

The effects of elevated CO2 and temperature on photosynthesis and calcification of two important calcifying reef algae (Halimeda macroloba and Halimeda cylindracea) were investigated with O2 microsensors and chlorophyll a fluorometry through a combination of two pCO2 (400 and 1,200 µatm) and two temperature treatments (28 and 32 °C) equivalent to the present and predicted conditions during the 2100 austral summer. Combined exposure to pCO2 and elevated temperature impaired calcification and photosynthesis in the two Halimeda species due to changes in the microenvironment around the algal segments and a reduction in physiological performance. There were no significant changes in controls over the 5-week experiment, but there was a 50-70 % decrease in photochemical efficiency (maximum quantum yield), a 70-80 % decrease in O2 production and a threefold reduction in calcification rate in the elevated CO2 and high temperature treatment. Calcification in these species is closely coupled with photosynthesis, such that a decrease in photosynthetic efficiency leads to a decrease in calcification. Although pH seems to be the main factor affecting Halimeda species, heat stress also has an impact on their photosystem II photochemical efficiency. There was a strong combined effect of elevated CO2 and temperature in both species, where exposure to elevated CO2 or temperature alone decreased photosynthesis and calcification, but exposure to both elevated CO2 and temperature caused a greater decline in photosynthesis and calcification than in each stress individually. Our study shows that ocean acidification and ocean warming are drivers of calcification and photosynthesis inhibition in Halimeda. Predicted climate change scenarios for 2100 would therefore severely affect the fitness of Halimeda, which can result in a strongly reduced production of carbonate sediments on coral reefs under such changed climate conditions.

Effects of ocean acidification on growth and physiology of Ulva lactuca (Chlorophyta) in a rockpool-scenario, link to supplementary data

Rising atmospheric CO2-concentrations will have severe consequences for a variety of biological processes. We investigated the responses of the green alga Ulva lactuca (Linnaeus) to rising CO2-concentrations in a rockpool scenario. U. lactuca was cultured under aeraton with air containing either preindustrial pCO2 (280µatm) or for the end of the 21st century predicted (700µatm) pCO2 for 31 days. We addressed the following question: Will elevated CO2-concentrations affect photosynthesis (net photosynthesis, rETR(max), Fv/Fm, pigment composition) and growth of U. lactuca in rockpools with limited water exchange? Two phases of the experiment were distinguished: In the initial phase (day 1-4) the Seawater Carbonate System (SWCS) of the culture medium could be adjusted to the selected atmospheric pCO2 condition by continuous aeration with target pCO2 values. In the second phase (day 4-31) the SWCS was largely determined by the metabolism of the growing U. lactuca biomass. In the initial phase, Fv/Fm and rETR(max) were only slightly elevated at high CO2-concentrations whereas growth was significantly enhanced. After 31 days the Chl a content of the thalli was significantly lower under future conditions and the photosynthesis of thalli grown under preindustrial conditions was not dependent on external carbonic anhydrase. Biomass increased significantly at high CO2-concentrations. At low CO2-concentrations most adult thalli disintegrated between day 14 and 21, whereas at high CO2-concentrations most thalli remained integer until day 31. Thallus disintegration at low CO2-concentrations was mirrored in a drastic decline in seawater DIC and HCO3-. Accordingly, the SWCS differed significantly between the treatments. Our results indicated a slight enhancement of photosynthetic performance and significantly elevated growth of U. lactuca at future CO2-concentrations. The accelerated thallus disintegration at high CO2-concentrations under conditions of limited water exchange indicates additional CO2 effects on the life cycle of U. lactuca when living in rockpools.

Age determination and paleotemperatures of sediment cores TTR17-1_384B and TTR17-1_436B in the Alboran Sea

Climate conditions in the westernmost Mediterranean (Alboran Sea basin) over the last two millennia have been reconstructed through integration of molecular proxies applied for the first time in this region at such high resolution. Two temperature proxies, one based on isoprenoid membrane lipids of marine Thaumarchaeota (TEXH86-tetraether index of compounds consisting of 86 carbons) and the other on alkenones produced by haptophytes (UK'37 ratio) were applied to reconstruct sea surface temperature (SST). Both records reveal a progressive long term decline in SST over the last two millennia and an increased rate of warming during the second half of the twentieth century. This is in accord with previous temperature reconstructions for the Northern Hemisphere. TEXH86 temperature values are higher than those inferred from UK'37, probably due to differences in the bloom season of haptophytes and Thaumarchaeota, and reflect summer SST. The branched vs. isoprenoid tetraether index (BIT index) suggests a low contribution of soil organic matter (OM) to the sedimentary OM. The stable carbon isotopic composition of long chain n-alkanes indicates a predominant C3 plant contribution, with no major change in vegetation over the last 2000 yr. The distribution of long chain 1,14-diols (most likely sourced by Proboscia species in this setting) provided insight into variation in upwelling conditions during the last 2000 yr and depicts a correlation with the North Atlantic Oscillation (NAO) index, providing evidence of enhanced wind induced upwelling during periods of a persistent positive mode of the NAO.

Die Gletscher der bolivianischen Anden, Verbreitung von Gletschern und neuzeitlichen Moränen in der Cordillera Apolobamba Cololo Region (Ostkordillere)

The uniform triangulation of the whole glacierized area of the Eastern Cordillera of Bolivia provided the first comparable geometrie basis for an accurate photogrammetric height evaluation of glaciers in the mountain range from the far south up to the Peruvian border. A total of 1775 glaciers, larger than 0.1 ha, were recorded in 16 mountain ranges ofthe East- ern Cordillera. Snow lines in the outer tropics can now be analysed with an accuracy of European standards. In spite of the widemeshed network of meteorological stations this allows a more pre- eise climatic differentiation to be made, particularly at high altitude sites. The methods and results of the analysis are described. Because of these results it becomes necessary to revise and refine previous concepts of the spatial distribution of climatic phenomena. Characteristics of glaciers in the Bolivian Cordillera obviously depend on the NE to SW decline in precipitation. The snow lines which are found at an altitude of 4500 m on the north- eastern slope directed towards the Yungas climb up to 5200 m on the lee slope towards the Alti- plano. The N to S increase in snowline elevation is not as significant as expected. With some res- ervation a 100 m fall in the snowline elevation in the Bolivian Cordillera can be explained by an increase in precipitation of about 50 mm.

Biological impacts of enhanced alkalinity in Carcinus maenas

Further steps are needed to establish feasible alleviation strategies that are able to reduce the impacts of ocean acidification, whilst ensuring minimal biological side-effects in the process. Whilst there is a growing body of literature on the biological impacts of many other carbon dioxide reduction techniques, seemingly little is known about enhanced alkalinity. For this reason, we investigated the potential physiological impacts of using chemical sequestration as an alleviation strategy. In a controlled experiment, Carcinus maenas were acutely exposed to concentrations of Ca(OH)2 that would be required to reverse the decline in ocean surface pH and return it to pre-industrial levels. Acute exposure significantly affected all individuals' acid-base balance resulting in slight respiratory alkalosis and hyperkalemia, which was strongest in mature females. Although the trigger for both of these responses is currently unclear, this study has shown that alkalinity addition does alter acid-base balance in this comparatively robust crustacean species.

Diatom-bound N isotope records over the last two glacial cycles in sediment core PS75/072-4

In a sediment core from the Pacific sector of the Antarctic Zone (AZ) of the Southern Ocean, we report diatom-bound N isotope (d15Ndb) records for total recoverable diatoms and two distinct diatom assemblages (pennate and centric rich). These data indicate tight coupling between the degree of nitrate consumption and Antarctic climate across the last two glacial cycles, with d15Ndb (and thus the degree of nitrate consumption) increasing at each major Antarctic cooling event. Coupled with evidence from opal- and barium-based proxies for reduced export production during ice ages, the d15Ndb increases point to ice age reductions in the supply of deep ocean-sourced nitrate to the AZ surface. The two diatom assemblages and species abundance data indicate that the d15Ndb changes are not the result of changing species composition. The pennate and centric assemblage d15Ndb records indicate similar changes but with a significant decline in their difference during peak ice ages. A tentative seasonality-based interpretation of the centric-to-pennate d15Ndb difference suggests that late summer surface waters became nitrate free during the peak glacials.

Mortality and climatic/environmental correlates of Widdringtonia cedargensis trees in the Cederberg, South Africa

Background: Conifer populations appear disproportionately threatened by global change. Most examples are, however, drawn from the northern hemisphere and long-term rates of population decline are not well documented as historical data are often lacking. We use a large and long-term (1931-2013) repeat photography dataset together with environmental data and fire records to account for the decline of the critically endangered Widdringtonia cedarbergensis. Eighty-seven historical and repeat photo-pairs were analysed to establish 20th century changes in W. cedarbergensis demography. A generalized linear mixed-effects model was fitted to determine the relative importance of environmental factors and fire-return interval on mortality for the species. Results: From an initial total of 1313 live trees in historical photographs, 74% had died and only 44 (3.4%) had recruited in the repeat photographs, leaving 387 live individuals. Juveniles (mature adults) had decreased (increased) from 27% (73%) to 8% (92%) over the intervening period. Our model demonstrates that mortality is related to greater fire frequency, higher temperatures, lower elevations, less rocky habitats and aspect (i.e. east-facing slopes had the least mortality). Conclusions: Our results show that W. cedarbergensis populations have declined significantly over the recorded period, with a pronounced decline in the last 30 years. Individuals that established in open habitats at lower, hotter elevations and experienced a greater fire frequency appear to be more vulnerable to mortality than individuals growing within protected, rocky environments at higher, cooler locations with less frequent fires. Climate models predict increasing temperatures for our study area (and likely increases in wildfires). If these predictions are realised, further declines in the species can be expected. Urgent management interventions, including seedling out-planting in fire-protected high elevation sites, reducing fire frequency in higher elevation populations, and assisted migration, should be considered.

Biogenic barium in Cretaceous-Paleogene sediments

One of the best-studied aspects of the K-Pg mass extinction is the decline and subsequent recovery of open ocean export productivity (e.g., the flux of organic matter from the surface to deep ocean). Some export proxies, including surface-to-deep water d13C gradients and carbonate sedimentation rates, indicate a global decline in export productivity triggered by the extinction. In contrast, benthic foraminiferal and other geochemical productivity proxies suggest spatially and temporally heterogeneous K-Pg boundary effects. Here we address these conflicting export productivity patterns using new and compiled measurements of biogenic barium. Unlike a previous synthesis, we find that the boundary effect on export productivity and the timing of recovery varied considerably between different oceanic sites. The northeast and southwest Atlantic, Southern Ocean, and Indian Ocean records saw export production plummet and remain depressed for 350 thousand to 2 million years. Biogenic barium and other proxies in the central Pacific and some upwelling or neritic Atlantic sites indicate the opposite, with proxies recording either no change or increased export production in the early Paleocene. Our results suggest that widespread declines in surface-to-deep ocean d13C do not record a global decrease in export productivity. Rather, independent proxies, including barium and other geochemical proxies, and benthic community structure, indicate that some regions were characterized by maintained or rapidly recovered organic flux from the surface ocean to the deep seafloor, while other regions had profound reductions in export productivity that persisted long into the Paleocene.

Under-ice export flux measurements by short-term drifting sediment traps at 27 stations in the Arctic Ocean during summer 1995, 1997, and 2012

A critical question regarding the organic carbon cycle in the Arctic Ocean is whether the decline in ice extent and thickness and the associated increase in solar irradiance in the upper ocean will result in increased primary production and particulate organic carbon (POC) export. To assess spatial and temporal variability in POC export, under-ice export fluxes were measured with short-term sediment traps in the northern Laptev Sea in July-August-September 1995, north of the Fram Strait in July 1997, and in the Central Arctic in August-September 2012. Sediment traps were deployed at 2-5 m and 20-25 m under ice for periods ranging from 8.5 to 71 h. In addition to POC fluxes, total particulate matter, chlorophyll a, biogenic particulate silica, phytoplankton, and zooplankton fecal pellet fluxes were measured to evaluate the amount and composition of the material exported in the upper Arctic Ocean. Whereas elevated export fluxes observed on and near the Laptev Sea shelf were likely the combined result of high primary production, resuspension, and release of particulate matter from melting ice, low export fluxes above the central basins despite increased light availability during the record minimum ice extent of 2012 suggest that POC export was limited by nutrient supply during summer. These results suggest that the ongoing decline in ice cover affects export fluxes differently on Arctic shelves and over the deep Arctic Ocean and that POC export is likely to remain low above the central basins unless additional nutrients are supplied to surface waters.