Thermal reaction norms of growth in Atlantic and Pacific silverside fishes

How organisms may adapt to rising global temperatures is uncertain, but concepts can emerge from studying adaptive physiological trait variations across existing spatial climate gradients. Many ectotherms, particularly fish, have evolved increasing genetic growth capacities with latitude (i.e. countergradient variation (CnGV) in growth), which are thought to be an adaptation primarily to strong gradients in seasonality. In contrast, evolutionary responses to gradients in mean temperature are often assumed to involve an alternative mode, 'thermal adaptation'. We measured thermal growth reaction norms in Pacific silverside populations (Atherinops affinis) occurring across a weak latitudinal temperature gradient with invariant seasonality along the North American Pacific coast. Instead of thermal adaptation, we found novel evidence for CnGV in growth, suggesting that CnGV is a ubiquitous mode of reaction-norm evolution in ectotherms even in response to weak spatial and, by inference, temporal climate gradients. A novel, large-scale comparison between ecologically equivalent Pacific versus Atlantic silversides (Menidia menidia) revealed how closely growth CnGV patterns reflect their respective climate gradients. While steep growth reaction norms and increasing growth plasticity with latitude in M. menidia mimicked the strong, highly seasonal Atlantic coastal gradient, shallow reaction norms and much smaller, latitude-independent growth plasticity in A. affinis resembled the weak Pacific latitudinal temperature gradient.

Strontium and oxygen isotope data for DSDP Hole 44-390A and ODP Leg 171 holes

Differences in regional responses to climate fluctuations are well documented on short time scales (e.g., El Niño-Southern Oscillation), but with the exception of latitudinal temperature gradients, regional patterns are seldom considered in discussions of ancient greenhouse climates. Contrary to the expectation of global warming or global cooling implicit in most treatments of climate evolution over millions of years, this paper shows that the North Atlantic warmed by as much as 6°C (1.5% decrease in d18O values of planktic foraminifera) during the Maastrichtian global cooling interval. We suggest that warming was the result of the importation of heat from the South Atlantic. Decreasing North Atlantic d18O values are also associated with increasing gradients in planktic d13C values, suggesting increasing surface-water stratification and a correlated strengthening of the North Atlantic Polar Front. If correct, this conclusion predicts arctic cooling during the late Maastrichtian. Beyond implications for the Maastrichtian, these data demonstrate that climate does not behave as if there is a simple global thermostat, even on geologic time scales.

ATPase activity, proton gradients and proton secretion inhibition during experiments with Sepia officinalis and Sepioteuthis lessoniana

The constraints of an active life in a pelagic habitat led to numerous convergent morphological and physiological adaptations that enable cephalopod molluscs and teleost fishes to compete for similar resources. Here, we show for the first time that such convergent developments are also found in the ontogenetic progression of ion regulatory tissues; as in teleost fish, epidermal ionocytes scattered on skin and yolk sac of cephalopod embryos appear to be responsible for ionic and acid-base regulation before gill epithelia become functional. Ion and acid-base regulation is crucial in cephalopod embryos, as they are surrounded by a hypercapnic egg fluid with a Pco2 between 0.2 and 0.4 kPa. Epidermal ionocytes were characterized via immunohistochemistry, in situ hybridization, and vital dye-staining techniques. We found one group of cells that is recognized by concavalin A and MitoTracker, which also expresses Na+/H+ exchangers (NHE3) and Na+-K+-ATPase. Similar to findings obtained in teleosts, these NHE3-rich cells take up sodium in exchange for protons, illustrating the energetic superiority of NHE-based proton excretion in marine systems. In vivo electrophysiological techniques demonstrated that acid equivalents are secreted by the yolk and skin integument. Intriguingly, epidermal ionocytes of cephalopod embryos are ciliated as demonstrated by scanning electron microscopy, suggesting a dual function of epithelial cells in water convection and ion regulation. These findings add significant knowledge to our mechanistic understanding of hypercapnia tolerance in marine organisms, as it demonstrates that marine taxa, which were identified as powerful acid-base regulators during hypercapnic challenges, already exhibit strong acid-base regulatory abilities during embryogenesis.

The evolutionary history of size variation of planktic foraminiferal assemblages in the Cenozoic

The iterative evolutionary radiation of planktic foraminifers is a well-documented macroevolutionary process. Here we document the accompanying size changes in entire planktic foraminiferal assemblages for the past 70 My and their relationship to paleoenvironmental changes. After the size decrease at the Cretaceous/Paleogene (K/P) boundary, high latitude assemblages remained consistently small. Size evolution in low latitudes can be divided into three major phases: the first is characterized by dwarfs (65-42 Ma), the second shows moderate size fluctuations (42-14 Ma), and in the third phase, planktic foraminifers have grown to the unprecedented sizes observed today. Our analyses of size variability with paleoproxy records indicate that periods of size increase coincided with phases of global cooling (Eocene and Neogene). These periods were characterized by enhanced latitudinal and vertical temperature gradients in the oceans and high diversity (polytaxy). In the Paleocene and during the Oligocene, the observed (minor) size changes of the largely low-diversity (oligotaxic) assemblages seem to correlate with productivity changes. However, polytaxy per se was not responsible for larger test sizes.

(Table A1) Isotopic ratios of Sr, Nd, and Pb for sediments in ODP Leg 138 sites

The provenance of eolian dust supplied to deep-sea sediments has the potential to offer insights into changes in past atmospheric circulation. Specifically, measuring temporal changes in dust provenance can shed light on changes in the mean position of the Intertropical Convergence Zone (ITCZ), a region acting as a barrier separating wind-blown material derived from northern versus southern hemisphere sources. Here we have analyzed Nd, Sr, and Pb isotope ratios in the operationally-defined detrital component extracted from deep-sea sediments in the eastern equatorial Pacific (EEP) along a meridional transect at 110°W from 3°S to 7°N (ODP Leg 138, sites 848-853). Sr isotope results show that barite Sr has a significant influence on 87Sr/86Sr isotope ratios of samples in the upwelling zone of the EEP. However, sites located >3° or more away from the equator (sites 852 and 853) are believed to not be affected by barite Sr and provide useful detrital Sr signals. 208Pb/206Pb and 207Pb/206Pb ratios in all cores fall into the Pb-isotope space of five potential dust sources (Asia, North and Central/South America, Sahara, and Australia), with no distinct isotopic fingerprinting of the dominant source(s). epsilon-Nd values were most valuable for discerning detrital source provenance, and their values at all sites, ranging from ~5.46 to ~3.25, were more unradiogenic for sediments deposited during the last glacial than for those deposited during the Holocene. There are distinct latitudinal trends in the epsilon-Nd values, with more radiogenic values further south and less radiogenic values further north, excluding site 848. This distinction holds true for both Holocene and last glacial periods. For the most southerly site, 848, we invoke, for the first time, a distinct southern hemisphere Australian source as being responsible for the unradiogenic Nd isotope ratios. Both average last glacial and Holocene epsilon-Nd values show similar sharp gradients along the transect between 5.29°N and 2.77°N, suggesting little movement of the glacial ITCZ in the EEP. However, during the deglacial, this gradient is stronger and shifted further north between 5.29°N and 7.21°N, suggesting a more northerly, possibly stronger, deglacial ITCZ.

Sea surface temperture reconstruction for ODP Site 189-1172

Relative to the present day, meridional temperature gradients in the Early Eocene age (~56-53 Myr ago) were unusually low, with slightly warmer equatorial regions (Pearson et al., 2007, doi:10.1130/G23175A.1 ) but with much warmer subtropical Arctic (Sluijs et al., 2008, doi:10.1029/2007PA001495) and mid-latitude (Sluijs et al., 2007, doi:10.1038/nature06400) climates. By the end of the Eocene epoch (~34 Myr ago), the first major Antarctic ice sheets had appeared (Zachos et al., 1992, doi:10.1130/0091-7613(1992)020<0569:EOISEO>2.3.CO;2; Barker et al., 2007, doi:10.1016/j.dsr2.2007.07.027), suggesting that major cooling had taken place. Yet the global transition into this icehouse climate remains poorly constrained, as only a few temperature records are available portraying the Cenozoic climatic evolution of the high southern latitudes. Here we present a uniquely continuous and chronostratigraphically well-calibrated TEX86 record of sea surface temperature (SST) from an ocean sediment core in the East Tasman Plateau (palaeolatitude ~65° S). We show that southwest Pacific SSTs rose above present-day tropical values (to ~34° C) during the Early Eocene age (~53 Myr ago) and had gradually decreased to about 21° C by the early Late Eocene age (~36 Myr ago). Our results imply that there was almost no latitudinal SST gradient between subequatorial and subpolar regions during the Early Eocene age (55-50 Myr ago). Thereafter, the latitudinal gradient markedly increased. In theory, if Eocene cooling was largely driven by a decrease in atmospheric greenhouse gas concentration Zachos et al. (2008, doi:10.1038/nature06588), additional processes are required to explain the relative stability of tropical SSTs given that there was more significant cooling at higher latitudes.

Foraminiferal oxygen and carbon isotope data for DSDP Sites 32-305 and 62-463 and ODP Sites 171-1049 and 171-1050

Oxygen isotope analyses of well-preserved foraminifera from Blake Nose (30°N paleolatitude, North Atlantic) and globally distributed deep-sea sites provide a long-term paleotemperature record for the late Albian-Maastrichtian interval that is difficult to reconcile with the existence of significant Cretaceous ice sheets. Given reasonable assumptions about the isotopic composition of Cretaceous seawater, our results suggest that middle bathyal water temperatures at Blake Nose increased from ~12°C in the late Albian through middle Cenomanian to a maximum of 20°C during the latest Cenomanian and earliest Turonian. Bottom waters were again ~12°C during the middle Campanian and cooled to a minimum of 9°C during the Maastrichtian. Correlative middle bathyal foraminifera from other ocean basins yield paleotemperature estimates that are very similar to those from Blake Nose. Comparison of global bottom-water temperatures and latitudinal thermal gradients suggests that global climate changed from a warm greenhouse state during the late Albian through late Cenomanian to a hot greenhouse phase during the latest Cenomanian through early Campanian, then to cool greenhouse conditions during the mid-Campanian through Maastrichtian.

Age profiles and isotopes d18O and d13C in Globigerina angiporoides and Oridorsalis umbonatus at DSDP Site 90-593 and Holes 12-111, 29-277 and 90-592

The late Eocene through earliest Miocene stable-isotope composition of southwest Pacific microfossils has been examined in a traverse of high-quality sedimentary sequences ranging from subantarctic (DSDP Site 277) through temperate regions (DSDP Sites 592 and 593). Changes in oxygen-isotope values, measured in benthic and planktonic foraminifers, document the Oligocene development and strengthening of latitudinal thermal zonation from water masses with broad temperature gradients during the Eocene to the steeper gradients and more distinct latitudinally distributed surface water-mass belts of the Neogene. The oxygen-isotope records can be divided into three intervals: late Eocene, early Oligocene, and middle to late Oligocene. Each interval represents a successive stage in the evolution of latitudinal thermal gradients between subantarctic and temperate regions in the Southern Hemisphere. During the late Eocene, oxygen-isotope values at subantarctic Site 277 were similar to those at temperate Sites 592 and 593. The isotope values suggest that, although the inferred paleotemperatures at Site 277 are slightly cooler on average than those at the temperate sites, there is no evidence for a major thermal boundary between the regions at this time. All three sites record the well-known oxygen-isotope enrichment of about 1 per mil in both planktonic and benthic foraminifers in close association with the Eocene/Oligocene boundary. In contrast to the earliest Oligocene enrichments in the planktonic and benthic oxygen-isotope composition at Site 277, more northern Sites 592 and 593 exhibit a depletion through the early-middle Oligocene. This documents the beginning of thermal segregation as subantarctic waters cooled relative to those at temperate latitudes. During the Oligocene, this surface-water differentiation continued, as measured by planktonic d18O values. The oxygen-isotope records of the benthic foraminifers also began to diverge in the earliest Oligocene. The most enriched oxygen-isotope values in all records cluster in the middle Oligocene, marked by oscillating episodes of enrichments >0.5 per mil occurring most prominently in the subantarctic record of Site 277. These values can be interpreted as recording either the coldest oceanic temperatures of the Paleogene and/or accumulations of Antarctic ice. After this interval, latitudinal thermal differentiation developed rapidly during the middle Oligocene, especially in the surface waters which actually warmed in temperate areas. If the enriched Oligocene oxygen-isotope values indicate that ice had accumulated, this ice must have disappeared by the early Miocene, when depleted oxygen-isotope values suggest very warm conditions. The data presented in this chapter document the progressive increase of latitudinal temperature gradients from the late Eocene through the late Oligocene. This pattern of increasing isotopic offset between latitudinally distributed southwest Pacific sites is linked to the establishment and strengthening of the Circum-Antarctic Current, previously considered to have developed during the middle to late Oligocene. The intensification of this current system progressively decoupled the warm subtropical gyres from cool polar circulation, in turn leading to increased Antarctic glaciation.

Aprubt climate change events in the Northeastern Atlantic Ocean

The western Iberian margin has been one of the key locations to study abrupt glacial climate change and associated interhemispheric linkages. The regional variability in the response to those events is being studied by combining a multitude of published and new records. Looking at the trend from Marine Isotope Stage (MIS) 10 to 2, the planktic foraminifer data, conform with the alkenone record of Martrat et al. [2007], shows that abrupt climate change events, especially the Heinrich events, became more frequent and their impacts in general stronger during the last glacial cycle. However, there were two older periods with strong impacts on the Atlantic meridional overturning circulation (AMOC): the Heinrich-type event associated with Termination (T) IV and the one occurring during MIS 8 (269 to 265 ka). During the Heinrich stadials of the last glacial cycle, the polar front reached the northern Iberian margin (ca. 41°N), while the arctic front was located in the vicinity of 39°N. During all the glacial periods studied, there existed a boundary at the latter latitude, either the arctic front during extreme cold events or the subarctic front during less strong coolings or warmer glacials. Along with these fronts sea surface temperatures (SST) increased southward by about 1°C per one degree of latitude leading to steep temperature gradients in the eastern North Atlantic and pointing to a close vicinity between subpolar and subtropical waters. The southern Iberian margin was always bathed by subtropical water masses - surface and/ or subsurface ones -, but there were periods when these waters also penetrated northward to 40.6°N. Glacial hydrographic conditions were similar during MIS 2 and 4, but much different during MIS 6. MIS 6 was a warmer glacial with the polar front being located further to the north allowing the subtropical surface and subsurface waters to reach at minimum as far north as 40.6°N and resulting in relative stable conditions on the southern margin. In the vertical structure, the Greenland-type climate oscillations during the last glacial cycle were recorded down to 2465 m during the Heinrich stadials, i.e. slightly deeper than in the western basin. This deeper boundary is related to the admixing of Mediterranean Outflow Water, which also explains the better ventilation of the intermediate-depth water column on the Iberian margin. This compilation revealed that latitudinal, longitudinal and vertical gradients existed in the waters along the Iberian margin, i.e. in a relative restricted area, but sufficient paleo-data exists now to validate regional climate models for abrupt climate change events in the northeastern North Atlantic Ocean.

Nitrogen isotope gradients off Peru and Ecuador related to upwelling, productivity, nutrient uptake and oxygen deficiency

We present new nitrogen isotope data from the water column and surface sediments for paleo-proxy validation collected along the Peruvian and Ecuadorian margins between 1°N and 18°S. Productivity proxies in the bulk sediment (organic carbon, total nitrogen, biogenic opal, C37 alkenone concentrations) and 15N/14N ratios were measured at more than 80 locations within and outside the present-day Peruvian oxygen minimum zone (OMZ). Microbial N-loss to N2 in subsurface waters under O2 deficient conditions leaves a characteristic 15N-enriched signal in underlying sediments. We find that phytoplankton nutrient uptake in surface waters within the high nutrient, low chlorophyll (HNLC) regions of the Peruvian upwelling system influences the sedimentary signal as well. How the d15Nsed signal is linked to these processes is studied by comparing core-top values to the 15N/14N of nitrate and nitrite (d15N[NOx]) in the upper 200 m of the water column. Between 1°N and 10°S, subsurface O2 is still high enough to suppress N-loss keeping d15NNOx values relatively low in the subsurface waters. However d15N[NOx] values increase toward the surface due to partial nitrate utilization in the photic zone in this HNLC portion of the system. d15N[sed] is consistently lower than the isotopic signature of upwelled [NO3]-, likely due to the corresponding production of 15N depleted organic matter. Between 10°S and 15°S, the current position of perennial upwelling cells, HNLC conditions are relaxed and biological production and near-surface phytoplankton uptake of upwelled [NO3]- are most intense. In addition, subsurface O2 concentration decreases to levels sufficient for N-loss by denitrification and/or anammox, resulting in elevated subsurface d15N[NOx] values in the source waters for coastal upwelling. Increasingly higher production southward is reflected by various productivity proxies in the sediments, while the north-south gradient towards stronger surface [NO3]- utilization and subsurface N-loss is reflected in the surface sediment 15N/14N ratios. South of 10°S, d15N[sed] is lower than maximum water column d15N[NOx] values most likely because only a portion of the upwelled water originates from the depths where highest d15N[NOx] values prevail. Though the enrichment of d15N[NOx] in the subsurface waters is unambiguously reflected in d15N[sed] values, the magnitude of d15N[sed] enrichment depends on both the depth of upwelled waters and high subsurface d15N[NOx] values produce by N-loss. Overall, the degree of N-loss influencing subsurface d15N[NOx] values, the depth origin of upwelled waters, and the degree of near-surface nitrate utilization under HNLC conditions should be considered for the interpretation of paleo d15N[sed] records from the Peruvian oxygen minimum zone.

Carbonate and terrigenous content and chemical composition of sediments in the Central Equatorial Pacific

This study addresses changes in the absolute magnitude and spatial geometry of particle flux and export production in a meridional transect across the central equatorial Pacific Ocean's upwelling system during oxygen isotope Stage 11 and Stage 12 and compares these time periods to the current Holocene interglacial system. Temporal and spatial variability in several chemical proxies of export production, and in particular the distributions of Ba, scavenged Al, and P, are studied in a suite of sediment cores gathered along a cross-equator transect at 5°S, 2°S, 0°, 2°N, and 4°N. Because this latitudinal range preserves strong gradients in biogenic particle flux in the modern equatorial Pacific Ocean, we are able to assess variations in the relative magnitude of export production as well as the meridional width of the equatorial system through the late Quaternary glacial/interglacial cycles. During interglacial oxygen isotope Stage 11 the chemical proxies each indicate lower particle flux and export production than during Stage 12. These records are consistent throughout the transect during this time period, but geographic narrowing (during the interglacial) and widening (during the glacial) of the meridional gradient also occurs. Although carbonate concentration varies dramatically through glacial/interglacial cycles at all latitudes studied, the productivity proxies record only minimal glacial/interglacial change at 5°S and 4°N, indicating that the carbonate minima at these latitudes is controlled dominantly by dissolution rather than production. The chemical data indicate that although the spatial geometry of the system during Stages 11 and 12 indicates maximum productivity at the equator during both glacial and interglacial conditions, the absolute magnitude of export production integrated from 5°S to 4°N during Stage 11 was 25-50% less than during Stage 12, and also was 25-50% less than it is now.

Sensitivity to ocean acidification parallels natural pCO2 gradients experienced by Arctic copepods under winter sea ice

The Arctic Ocean is a bellwether for ocean acidification, yet few direct Arctic studies have been carried out and limited observations exist, especially in winter. We present unique under-ice physicochemical data showing the persistence of a mid water column area of high CO2 and low pH through late winter, Zooplankton data demonstrating that the dominant copepod species are distributed across these different physicochemical conditions, and empirical data demonstrating that these copepods show sensitivity to pCO2 that parallels the range of natural pCO2 they experience through their daily vertical migration behavior. Our data, collected as part of the Catlin Arctic Survey, provide unique insight into the link between environmental variability, behavior, and an organism's physiological tolerance to CO2 in key Arctic biota.

Seawater carbonate chemistry and calcium carbonate of Padina spp., photosynthesis of Padina pavonica in nature CO2 gradients experiment

Predicting the impacts of ocean acidification on coastal ecosystems requires an understanding of the effects on macroalgae and their grazers, as these underpin the ecology of rocky shores. Whilst calcified coralline algae (Rhodophyta) appear to be especially vulnerable to ocean acidification, there is a lack of information concerning calcified brown algae (Phaeophyta), which are not obligate calcifiers but are still important producers of calcium carbonate and organic matter in shallow coastal waters. Here, we compare ecological shifts in subtidal rocky shore systems along CO2 gradients created by volcanic seeps in the Mediterranean and Papua New Guinea, focussing on abundant macroalgae and grazing sea urchins. In both the temperate and tropical systems the abundances of grazing sea urchins declined dramatically along CO2 gradients. Temperate and tropical species of the calcifying macroalgal genus Padina (Dictyoaceae, Phaeophyta) showed reductions in CaCO3 content with CO2 enrichment. In contrast to other studies of calcified macroalgae, however, we observed an increase in the abundance of Padina spp. in acidified conditions. Reduced sea urchin grazing pressure and significant increases in photosynthetic rates may explain the unexpected success of decalcified Padina spp. at elevated levels of CO2. This is the first study to provide a comparison of ecological changes along CO2 gradients between temperate and tropical rocky shores. The similarities we found in the responses of Padina spp. and sea urchin abundance at several vent systems increases confidence in predictions of the ecological impacts of ocean acidification over a large geographical range.