The stratigraphy of the Fishguard volcanic complex and associated sediments, SW Dyfed, Wales

This thesis describes the geology of a Lower Palaeozoic terrain, situated west of the town of Fishguard, SW Dyfed, Wales. The area is dominated by the Fishguard Volcanic Complex (Upper Llanvirn), and sediments that range in age from the Middle Cambrian to the Lower Llandeilo. The successions represent an insight into sedimentation and volcanism for c. 100 Ma. along the south-western margin of the Lower Palaeozoic Welsh Basin. The stratigraphy of the sedimentary sequence has been completely revised and the existing volcanostratigraphy modified. The observed complexity of the stratigraphy is primarily the consequence of Caldedonide deformation which resulted in large scale repetition. Fold-thrust tectonics dominates the structural style of the area. Caledonide trending (NE-SW) cross-faults complicate preexisting structures. Middle Cambrian (?) sedimentation is documented by shallow marine clastics and red shales deposited within tidal - subtidal environments. Upper Cambrian sedimentation was dominated by shallow marine `storm' and `fair weather' sedimentation within a muddy shelf environment. Shallow marine conglomerates and heterolithic intertidal siliciclastics mark the onset of Ordovician sedimentation during the lower Arenig transgression. Mid-Arenig sediments reflect deposits influenced by storm, fair-weather and wave related processes in various shallow marine environments, including; shoreface, inner shelf, shoaling bar, and deltaic. Graptolitic marine shales were deposited from the upper mid-Arenig through to the lower Llandeilo; during which time sediments accumulated by pelagic processes and fine grained turbidites. The varied nature of sedimentation reflects both localised change within the depositional system and the influence of larger regional eustatic events. Ordovician subaqueous volcanic activity produced thick accumulations of lavas, pyroclastics, hydroclastics, and hyaloclastics. The majority of volcanism was effusive in nature, erupted below the Pressure Compensation Level. Basaltic volcanism was characterised by pillowed lavas and tube networks, whilst sheet-flow lavas, pillow breccias and minor hyaloclastites developed locally. Silicic volcanism was dominated by rhyolitic clastics of various affinities, although coherent silicic obsidian lavas, sheet-flow lavas and pyroclastics developed. Hypabyssal intrusives of variable composition and habit occur throughout the volcanic successions. Low-grade regional metamorphism has variably affected the area, conditions of the prehnite-pumpellyite and greenschist facies having been attained. Numerous secondary phases developed in response to the conditions imposed, which collectively indicate that P-T conditions were of low-pressure facies series in the range P= 1.2-2.0 kbars and T= 230-350oC, under an elevated geothermal gradient of 40-45oC km-1. Polymineralic cataclastites associated with Caledonide deformation indicate that tectonism and metamorphism were in part contemporaneous.

Water temperature reconstructions based on paired Uk'37 and Tex86 records, and paired surface and subsurface Mg/Ca records

This dataset contains the collection of available published paired Uk'37 and Tex86 records spanning multi-millennial to multi-million year time scales, as well as a collection of Mg/Ca-derived temperatures measured in parallel on surface and subsurface dwelling foraminifera, both used in the analyses of Ho and Laepple, Nature Geoscience 2016. As the signal-to-noise ratios of proxy-derived Holocene temperatures are relatively low, we selected records that contain at least the last deglaciation (oldest sample >18kyr BP).

Neodymium isotopic characterization of Ross Sea Bottom Water and its advection through the southern South Pacific, POLARSTERN cruise ANT-XXVI/2

Since the inception of the international GEOTRACES program, studies investigating the distribution of trace elements and their isotopes in the global ocean have significantly increased. In spite of this large-scale effort, the distribution of neodymium isotopes (143Nd/144Nd) and concentrations ([Nd]) in the high latitude south Pacific is still understudied. Here we report dissolved Nd isotopes and concentrations from 11 vertical water column profiles from the south Pacific between South America and New Zealand. Results suggest that Ross Sea Bottom Water (RSBW) is represented by an epsilon-Nd value of ~ -7, and is thus more radiogenic than Circumpolar Deep Water (epsilon-Nd ~ -8). RSBW and its characteristic epsilon-Nd signature can be traced far into the SE Pacific until progressive mixing with ambient Lower Circumpolar Deep water (LCDW) dilutes this signal north of the Antarctic Polar Front (APF). The SW-NE trending Pacific-Antarctic Ridge restricts the advection of RSBW into the SW Pacific, where bottom water density, salinity, and epsilon-Nd values of -9 indicate the presence of bottom waters of an origin different from the Ross Sea. Neodymium concentrations show low surface concentrations and a linear increase with depth north of the Polar Front. South of the APF, surface [Nd] is high and increases with depth but remains almost constant below ~1000 m. This vertical and spatial [Nd] pattern follows the southward shoaling density surfaces of the Southern Ocean frontal system and hence suggests supply of Nd to the upper ocean through upwelling of Nd-rich deep water. Low particle abundance dominated by reduced opal production and seasonal sea ice cover likely contributes to the maintenance of the high upper ocean [Nd] south of the APF. The reported data highlights the use of Nd isotopes as a water mass tracer in the Southern Ocean, with the potential for paleocenaographic reconstructions, and contributes to an improved understanding of Nd biogeochemistry.

Rising CO2 and increased light exposure synergistically reduce marine primary productivity

Carbon dioxide and light are two major prerequisites of photosynthesis. Rising CO2 levels in oceanic surface waters in combination with ample light supply are therefore often considered stimulatory to marine primary production. Here we show that the combination of an increase in both CO2 and light exposure negatively impacts photosynthesis and growth of marine primary producers. When exposed to CO2 concentrations projected for the end of this century, natural phytoplankton assemblages of the South China Sea responded with decreased primary production and increased light stress at light intensities representative of the upper surface layer. The phytoplankton community shifted away from diatoms, the dominant phytoplankton group during our field campaigns. To examine the underlying mechanisms of the observed responses, we grew diatoms at different CO2 concentrations and under varying levels (5-100%) of solar radiation experienced by the phytoplankton at different depths of the euphotic zone. Above 22-36% of incident surface irradiance, growth rates in the high-CO2-grown cells were inversely related to light levels and exhibited reduced thresholds at which light becomes inhibitory. Future shoaling of upper-mixed-layer depths will expose phytoplankton to increased mean light intensities. In combination with rising CO2 levels, this may cause a widespread decline in marine primary production and a community shift away from diatoms, the main algal group that supports higher trophic levels and carbon export in the ocean.

(Table 1) Comparison between bulk ICP-OES measurements and LA-ICP-MS analyses for Mg/Ca and Sr/Ca ratios of Globigerinoides sacculifer from ODP Hole 165-1000A

We constructed a high-resolution Mg/Ca record on the planktonic foraminifer Globigerinoides sacculifer in order to explore the change in sea surface temperature (SST) due to the shoaling of the Isthmus of Panama as well as the impact of secondary factors like diagenesis and large salinity fluctuations. The study covers the latest Miocene and the early Pliocene (5.6-3.9 Ma) and was combined with d18O to isolate changes in sea surface salinity (SSS). Before 4.5 Ma, SSTMg/Ca and SSS show moderate fluctuations, indicating a free exchange of surface ocean water masses between the Pacific and the Atlantic. The increase in d18O after 4.5 Ma represents increasing salinities in the Caribbean due to the progressive closure of the Panamanian Gateway. The increase in Mg/Ca toward values of maximum 7 mmol/mol suggests that secondary influences have played a significant role. Evidence of crystalline overgrowths on the foraminiferal tests in correlation with aragonite, Sr/Ca, and productivity cyclicities indicates a diagenetic overprint on the foraminiferal tests. Laser ablation inductively coupled plasma-mass spectrometry analyses, however, do not show significantly increased Mg/Ca ratios in the crystalline overgrowths, and neither do calculations based on pore water data conclusively result in significantly elevated Mg/Ca ratios in the crystalline overgrowths. Alternatively, the elevated Mg/Ca ratios might have been caused by salinity as the d18O record of Site 1000 has been interpreted to represent large fluctuations in SSS, and cultivating experiments have shown an increase in Mg/Ca with increasing salinity. We conclude that the Mg/Ca record <4.5 Ma can only reliably be considered for paleoceanographical purposes when the minimum values, not showing any evidence of secondary influences, are used, resulting in a warming of central Caribbean surface water masses after 4.5 Ma of ~2°C.

Quantitative assessment of intrinsic noise for visually guided behaviour in zebrafish

Supported by Royal Society of London (University Research Fellowship), Medical Research Council (New Investigator Research Grant) and CNRS.

Surface wind stress modeled for the Panama Seaway with the Community Climate System Model 3

The early Pliocene warm phase was characterized by high sea surface temperatures and a deep thermocline in the eastern equatorial Pacific. A new hypothesis suggests that the progressive closure of the Panamanian seaway contributed substantially to the termination of this zonally symmetric state in the equatorial Pacific. According to this hypothesis, intensification of the Atlantic meridional overturning circulation (AMOC) - induced by the closure of the gateway - was the principal cause of equatorial Pacific thermocline shoaling during the Pliocene. In this study, twelve Panama seaway sensitivity experiments from eight ocean/climate models of different complexity are analyzed to examine the effect of an open gateway on AMOC strength and thermocline depth. All models show an eastward Panamanian net throughflow, leading to a reduction in AMOC strength compared to the corresponding closed-Panama case. In those models that do not include a dynamic atmosphere, deepening of the equatorial Pacific thermocline appears to scale almost linearly with the throughflow-induced reduction in AMOC strength. Models with dynamic atmosphere do not follow this simple relation. There are indications that in four out of five models equatorial wind-stress anomalies amplify the tropical Pacific thermocline deepening. In summary, the models provide strong support for the hypothesized relationship between Panama closure and equatorial Pacific thermocline shoaling.

(Table 2) Terrigenous, authigenic and biogenic composition of sediments at DSDP Hole 93-603B

Pelagic sedimentation during the Early Cretaceous at Site 603 produced alternations of laminated marly limestone and bioturbated limestone-a facies typical of the "Blake-Bahama Formation" of the western Atlantic. This limestone is a nannofossil micrite, rich in calcified radiolarians, with variable amounts of organic matter, pyritized radiolarian tests, fish debris, and micaceous silt. The laminated marly limestone layers are enriched in organic matter when compared with the intervals of bioturbated limestone. The organic carbon is predominantly terrestrial plant debris; where the organic-carbon content is in excess of 1%, there is also a significant marine-derived component. Laminations can result either from bands of alternately enriched and depleted opaque material and clay, or from bands of elongate lenses (microflasers) of micrite, which could be plastically deformed pellets or diagenetic features. The alternating intervals of laminated and bioturbated structures may have resulted from combined changes in surface productivity, in the influx of terrigenous organic matter, and in the intensity of bottom circulation, which led to episodic oxygen depletion in the bottom water and sediments. Variations in the relative proportions of laminated clay-rich and bioturbated lime-rich limestone and in the development of cycles between these structures make it possible to subdivide the Lower Cretaceous pelagic facies into several subunits which appear to be regional in extent. Bioturbated limestone is dominant in the Berriasian, laminated marly limestone in the Valanginian and Barremian-lower Aptian, and well-developed alternations between these end members in the Hauterivian. The Hauterivian to lower Aptian sediments contain abundant terrigenous clastic turbidites associated with a submarine fan complex. These changes in the general characteristics of the pelagic sediment component of the Blake-Bahama Formation at Site 603 are synchronous with those in the Blake-Bahama Basin (Sites 534 and 391) to the south. Carbonate sedimentation ended in the early Aptian, probably because of a regional shoaling of the carbonate compensation depth.

Stable carbon and oxygen isotopes and Mg/Ca ratios of foraminifers from ODP Site 202-1241 sediments

A combination of stable isotope records and Mg/Ca temperature estimates of four different planktonic foraminiferal species from Ocean Drilling Program Site 1241 allows differentiation between temperature and salinity changes in the tropical east Pacific (TEP) upper water column during the Pliocene (~5.7-2.1 Ma). The deviation of d18O records and Mg/Ca temperature estimates from thermocline-dwelling planktonic foraminifers suggests that local changes in salinity exerted a much stronger control on Pliocene TEP upper ocean water mass signatures than previously assumed. The most pronounced Pliocene change in TEP upper ocean stratification was the shoaling of the thermocline from ~4.8 to 4.0 Ma that was possibly triggered by changes in the configuration of low-latitude ocean gateways. During this time interval, mixed-layer temperatures and salinities remained relatively constant in contrast to a pronounced temperature (~6°C) and salinity decrease at the bottom of the photic zone. This change led to a new state in the thermal structure of the TEP, as the thermocline remained relatively shallow until ~2.1 Ma.

(Table S1) Age determination of sediment cores from the Labrador Sea

High resolution flow speed reconstructions of two core sites located on Gardar Drift in the northeast Atlantic Basin and Orphan Knoll in the northwest Atlantic Basin reveal a long-term decrease in flow speed of Northeast Atlantic Deep Water (NEADW) after 6,500 years. Benthic foraminiferal oxygen isotopes of sites currently bathed in NEADW show a 0.2per mil depletion after 6,500 years, shortly after the start of the development of a carbon isotope gradient between NEADW and Norwegian Sea Deep Water. We consider these changes in near-bottom flow vigor and benthic foraminiferal isotope records to mark a significant reorganization of the Holocene deep ocean circulation, and attribute the changes to a weakening of NEADW flow during the mid to late Holocene that allowed the shoaling of Lower Deep Water and deeper eastward advection of Labrador Sea Water into the northeast Atlantic Basin.

Delta 44/40 Ca, Mg/Ca and Sr/Ca record of Globigerinoides sacculifer and calculated temperatures and salinity changes of ODP Hole 165-999A

The delta18O values of planktonic foraminifera increased in the Caribbean by about 0.5? relative to the equatorial East Pacific values between 4.6 and 4.2 Ma as a consequence of the closure of the Central American Gateway (CAG). This increase in delta18O can be interpreted either as an increase in Caribbean sea surface (mixed layer) salinity (SSS) or as a decrease in sea surface temperatures (SST). This problem represents an ideal situation to apply the recently developed paleotemperature proxy delta44/40Ca together with Mg/Ca and d18O on the planktic foraminifer Globigerinoides sacculifer from ODP Site 999. Although differences in absolute temperature calibration of delta44/40Ca and Mg/Ca exist, the general pattern is similar indicating a SST decrease of about 2-3 8C between 4.4 and 4.3 Ma followed by an increase in the same order of magnitude between 4.3 and 4.0 Ma. Correcting the delta18O record for this temperature change and assuming that changes in global ice volume are negligible, the salinity-induced planktonic delta18O signal decreased by about 0.4? between 4.4 and 4.3 Ma and increased by about 0.9? between 4.3 and 4.0 Ma in the Caribbean. The observed temperature and salinity trends are interpreted to reflect the restricted exchange of surface water between the Caribbean and the Pacific in response to the shoaling of the Panamanian Seaway, possibly accompanied by a southward shift of the Intertropical Convergence Zone (ITCZ) between 4.4 and 4.3 Ma. Differences in Mg/Ca- and delta44/40Ca-derived temperatures can be reconciled by corrections for secular variations of the marine Mg/Ca[sw] and delta44/40Ca, a salinity effect on the Mg/Ca ratio and a constant temperature offset of ~2.5 °C between both SST proxy calibrations.

Sea surface temperatures reconstructed from sediment core U1338

This study presents a new alkenone-derived Sea Surface Temperature (SST) record and d18ONoelaerhabdaceae data of the 2-5 µm carbonate fractions from the IODP site U1338 located in the Eastern Equatorial Pacific (EEP), over the Miocene-Pliocene. Our data and those available from other sites of the same area show the establishment of a cold tongue during the early Pliocene (4.4 - 3.6 Ma). SST and d18ONoelaerhabdaceae time-series indicate periods of significant salinity variations. Comparison with the d18Obenthic curve from sediment cores of the Equatorial Pacific Ocean allow us to distinguish between global changes and local surface salinity variations in the EEP. Ice sheet growth and evaporation-precipitation are then discussed as possible drivers of such changes, as well as the role of Central American and the Indonesian seaway restriction. Our data suggest a shallowing of the thermocline in the EEP, between 6.8 and 6 Ma, and its shoaling between 4.8 and 4.0 Ma, suggesting the appearance of the cold tongue (Steph et al., 2010). The Pliocene climate transition would therefore not be primarily driven by Northern hemisphere glaciation.

(Table 2) Age determination of sediment cores from the Portuguese Margin

We demonstrate that changes in the behavior of the Mediterranean Outflow Water (MOW) prior to and through the last deglaciation played an important role in promoting Meridional Overturning Circulation (MOC). Estimation of past MOW salt and heat fluxes indicates that they gradually increased through the last deglaciation. Between 17.5 and 14.6 thousand years ago (ka B.P., where B.P. references year 1950), net evaporation from the Mediterranean exported sufficient fresh water from the North Atlantic catchment to cause an average salinity increase of 0.5 psu throughout the upper 2000 m of the entire North Atlantic to the north of 25°N. Combined with rapid intensification and shoaling of the MOW plume, which we identify around 15-14.5 ka B.P., this deglacial MOW-related salt accumulation preconditioned the North Atlantic for abrupt resumption of the MOC at 14.6 ka B.P.

Middle Miocene evolution and stable isotope ratios of Globorotalia (Fohsella) in the western equatorial Pacific

Evolution of the planktic foraminiferal lineage Globorotalia (Fohsella) occurred during the Miocene between 23.7 and 11.8 Ma and forms the basis for stratigraphic subdivision of the early middle Miocene (Zones N 10 through N 12). Important morphologic changes within the G. (Fohsella) lineage included a marked increase in test size, a transition from a rounded to an acute periphery, and the development of a keel in later forms. We found that the most rapid changes in morphology of G. (Fohsella) occurred between 13 and 12.7 Ma and coincided with an abrupt increase in the delta18O ratios of shell calcite. Comparison of isotopic results of G. (Fohsella) with other planktic foraminifers indicate that delta18O values of the lineage diverge from surface-dwelling species and approach deep-dwelling species after 13.0 Ma, indicating a change in depth habitat from the surface mixed layer to intermediate depth near the thermocline. Isotopic and faunal evidence suggests that this change in depth stratification was associated with an expansion of the thermocline in the western equatorial Pacific. After adapting to a deeper water habitat at 13.0 Ma, the G. (Fohsella) lineage became extinct abruptly at 11.8 Ma during a period when isotopic and faunal evidence suggest a shoaling of the thermocline. Following the extinction of G. (Fohsella), the ecologic niche of the lineage was filled by the Globorotalia (Menardella) group, which began as a deep-water form and later evolved to an intermediate-water habitat. We suggest that the evolution of G. (Fohsella) and G. (Menardella) were tightly linked to changes in the structure of the thermocline in the western equatorial Pacific.

SOCIAL AND ECOLOGICAL FACTORS INFLUENCING COLLECTIVE BEHAVIOR IN GIANT DANIO

A fundamental problem in biology is understanding how and why things group together. Collective behavior is observed on all organismic levels - from cells and slime molds, to swarms of insects, flocks of birds, and schooling fish, and in mammals, including humans. The long-term goal of this research is to understand the functions and mechanisms underlying collective behavior in groups. This dissertation focuses on shoaling (aggregating) fish. Shoaling behaviors in fish confer foraging and anti-predator benefits through social cues from other individuals in the group. However, it is not fully understood what information individuals receive from one another or how this information is propagated throughout a group. It is also not fully understood how the environmental conditions and perturbations affect group behaviors. The specific research objective of this dissertation is to gain a better understanding of how certain social and environmental factors affect group behaviors in fish. I focus on two ecologically relevant decision-making behaviors: (i) rheotaxis, or orientation with respect to a flow, and (ii) startle response, a rapid response to a perceived threat. By integrating behavioral and engineering paradigms, I detail specifics of behavior in giant danio Devario aequipinnatus (McClelland 1893), and numerically analyze mathematical models that may be extended to group behavior for fish in general, and potentially other groups of animals as well. These models that predict behavior data, as well as generate additional, testable hypotheses. One of the primary goals of neuroethology is to study an organism's behavior in the context of evolution and ecology. Here, I focus on studying ecologically relevant behaviors in giant danio in order to better understand collective behavior in fish. The experiments in this dissertation provide contributions to fish ecology, collective behavior, and biologically-inspired robotics.