Respiration rates and abundances of dominant copepods of the northern Benguela Current System

Respiration rates of 16 calanoid copepod species from the northern Benguela upwelling system were measured on board RRS Discovery in September/October 2010 to determine their energy requirements and assess their significance in the carbon cycle. Individual respiration rates were standardised to a mean copepod body mass and a temperature regime typical of the northern Benguela Current. These adjusted respiration rates revealed two different activity levels (active and resting) in copepodids C5 of Calanoides carinatus and females of Rhincalanus nasutus, which reduced their metabolism during dormancy by 82% and 62%, respectively. An allometric function (Imax) and an energy budget approach were performed to calculate ingestion rates. Imax generally overestimated the ingestion rates derived from the energy budget approach by >75%. We suggest that the energy budget approach is the more reliable approximation with a total calanoid copepod (mainly females) consumption of 78 mg C m-2 d-1 in neritic regions and 21 mg C m-2 d-1 in oceanic regions. The two primarily herbivorous copepods C. carinatus (neritic) and Nannocalanus minor (oceanic) contributed 83% and 5%, respectively, to total consumption by calanoid copepods. Locally, C. carinatus can remove up to 90% of the diatom biomass daily. In contrast, the maximum daily removal of dinoflagellate biomass by N. minor was 9%. These estimates imply that C. carinatus is an important primary consumers in the neritic province of the northern Benguela system, while N. minor has little grazing impact on phytoplankton populations further offshore. Data on energy requirements and total consumption rates of dominant calanoid copepods of this study are essential for the development of realistic carbon budgets and food-web models for the northern Benguela upwelling system.

Organic carbon, reduced sulfur, and iron in Miocene to Holocene sediments from the Oman and Beguela upwelling systems

We examined sediments from Neogene and Quaternary sections of the Benguela and Oman upwelling systems (DSDP Site 532, ODP Sites 723 and 722) to determine environmental and geochemical factors which control and limit pyrite formation in organic-carbon-rich marine sediments. Those samples from the upwelling sites, which contained low to moderate concentrations of total organic carbon (0.7%-3%), had C/S ratios typical of normal marine sediments, i.e., around 2.8. In these sediments, TOC availability probably limited pyrite formation. Results that do not conform with accepted models were found for the sediments high in TOC (3^0-12.4%). The organic matter was of marine origin and contained considerable pyrolytic hydrocarbons, a fact that we take as a sign of low degradation, yet significant concentrations of dissolved sulfate coexisted with it (> 5 mmol/L in the case of Sites 532 and 723). Detrital iron was probably not limiting in either case, because the degree of pyritization was always less than 0.65. Therefore, controls on sulfate reduction and pyrite formation in the organic matter-rich sediments do not appear to conform simply to generally accepted diagenetic models. The data from these thermally immature, old, and organic-rich marine sediments imply that (1) the total reduced sulfur content of organic-rich marine upwelling sediments rarely exceeds an approximate boundary of 1.5% by weight, (2) the C/S ratio of these sediments is not constant and usually much higher than the empirical values proposed for marine sediments. We conclude that sedimentary pyrite formation in upwelling sediments is limited by an as yet unknown factor, and that caution is advised in using C/S ratios and C vs. S diagrams in paleoenvironmental reconstructions for organic-rich sediments.

The Impacts of Inter-El Nino Variability on the Tropical Atlantic and Northeast Brazil Climate

In this study, observations and numerical simulations are used to investigate how different El Nino events affect the development of SST anomalies in the Atlantic and how this relates to the Brazilian northeast (NE) precipitation. The results show that different types of El Nino have different impacts on the SST anomalies of the equatorial and tropical South Atlantic but a similar SST response in the tropical North Atlantic. Strong and long (weak and short) El Ninos with the main heating source located in the eastern (central) Pacific generate cold (warm) anomalies in the cold tongue and Benguela upwelling regions during boreal winter and spring. When the SST anomalies in the eastern equatorial and tropical South Atlantic are cold (warm), the meridional SST gradient across the equator is positive (negative) and the ITCZ is not allowed (allowed) to move southward during the boreal spring; as a consequence, the precipitation is below (above) the average over the NE. Thus, strong and long (weak and short) El Ninos are followed by dry (wet) conditions in the NE. During strong and long El Ninos, changes in the Walker circulation over the Atlantic and in the Pacific-South Atlantic (PSA) wave train cause easterly wind anomalies in the western equatorial Atlantic, which in turn activate the Bjerknes mechanism, establishing the cold tongue in boreal spring and summer. These easterly anomalies are also responsible for the Benguela upwelling. During short and weak El Ninos, westerly wind anomalies are present in the western equatorial Atlantic accompanied by warm anomalies in the eastern equatorial and tropical South Atlantic; a positive phase of the South Atlantic dipole develops during boreal winter. The simulations highlight the importance of ocean dynamics in establishing the correct slope of the equatorial thermocline and SST anomalies, which in turn determine the correct rainfall response over the NE.

Análise genética do tubarão-raposa Alopias superciliosus no Oceano Atlântico, utilizando região controle do DNA mitocondrial

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Oxygen and Chlorophyll Distribution in the Eastern Atlantic Ocean from 0º to 35ºS.

Programa de doctorado en Oceanografía. Trabajo presentado para la obtención del Diploma de Estudios Avanzados.

Stable isotope record of planktonic foraminifera from sediment core MD96-2085

A high-resolution (~4-5cm/kyr) giant piston core record (MD962085) retrieved during an IMAGES II-NAUSICAA cruise from the continental slope of the southeast Atlantic Ocean reveals striking variations in planktonic foraminifer faunal abundances and sea-surface temperatures (SST) during the past 600 000 yr. The location and high-quality sedimentary record of the core provide a good opportunity to assess the variability of the Benguela Current system and associated important features of the ocean-climate system in the southeast Atlantic. The planktonic foraminifer faunal abundances of the core are dominated by three assemblages: (1) Neogloboquadrina pachyderma (right coiling) + Neogloboquadrina dutertrei, (2) Globigerina bulloides, and (3) Globorotalia inflata. The assemblage of N. pachyderma (right coiling) + N. dutertrei shows distinctive abundance changes which are nearly in-phase with glacial-interglacial variations. The high abundances of this assemblage are associated with major glacial conditions, possibly representing low SST/high nutrient level conditions in the southwestern Africa margin. In contrast, the G. bulloides and G. inflata assemblages show greater high-frequency abundance change patterns, which are not parallel to the glacial-interglacial changes. These patterns may indicate rapid oceanic frontal movements from the south, and a rapid change in the intensity of the Benguela upwelling system from the east. A single episode of maximum abundances of a polar water species N. pachyderma (left coiling) occurred in the beginning of stage 9 (~340-330 kyr). The event of the maximum occurrence of this species shown in this record may indicate instability in the Benguela coastal upwelling, or the Antarctic polar front zone position. A winter season SST estimate using transfer function techniques for this record shows primarily glacial-interglacial variations. The SST is maximal during the transitions from the major glacial to interglacial stages (Terminations I, II, IV, V), and is associated with the abundance maxima of a warm water species indicator Globigerinoides ruber. Cross-spectral analyses of the SST record and the SPECMAP stack reveal statistically significant concentrations of variance and coherencies in three major orbital frequency bands. The SST precedes changes in the global ice volume in all orbital frequency bands, indicating a dominant southern Hemispheric climate effect over the Benguela Current region in the southeast Atlantic.

Dinoflagellate cyst counts and age model from sediment samples of ODP Hole 175-1085A

Sediment samples from ODP Site 1085 were investigated in order to obtain more information on the initiation and development of the Benguela upwelling system during the middle and upper Miocene. In particular, our intent was to establish the causes of the upwelling as well as the response of the upwelling regime to the development of the Antarctic Circumpolar Current. Based on changes in the calcareous dinoflagellate cyst association, we found an initial increase of the dinoflagellate cyst productivity, probably related to the initiation of upwelling about 11.8 Ma ago. Two distinct increases in cyst productivity in conjunction with temperature decreases of the upper water masses reflect upwelling pulses off Namibia and occur at the end of the Miocene cooling events Mi5 (about 11.5 Ma) and Mi6 (about 10.5 Ma). Both cooling events are associated with an ice volume increase in Antarctica and are thought to have led to an increase in southeasterly winds, possibly causing these two upwelling pulses. We demonstrate a decrease in dinoflagellate cyst productivity and enhanced terrigenous input via the Orange River after the Mi5 event. At about 11.1 Ma, the dinoflagellate cyst productivity increases again. The polar cyst species Caracomia arctica occurs here for the first time. This implies an influence of subantarctic mode water and therefore a change in the quality of the upwelling water which allowed the Benguela upwelling to develop into modern conditions. From about 10.4 Ma, C. arctica forms a permanent part of the association, pointing to an establishment of the upwelling regime.

Pliocene-Pleistocene alkenone, nitrogen and opal record of ODP Site 175-1082

In this study we present combined high-resolution records of sea surface temperature (SST), phytoplankton productivity, and nutrient cycling in the Benguela Upwelling System (BUS) for the past 3.5 Ma. The SST record provided evidence that upwelling activity off Namibia mainly intensified ca. 2.4-2.0 Ma ago in response to the cooling of the Southern Ocean and the resultant strengthening of trade winds. As revealed by productivity-related proxies, BUS intensification led to a major transition in regional biological productivity when considering the termination of the Matuyama Diatom Maximum (a diatom high-production event). Major oceanic reorganization in the Benguela was accompanied by nutrient source changes, as indicated by a new nitrogen isotopic (delta15N) record that revealed a stepwise increase at ca. 2.4 and ca. 1.5 Ma ago. The change in source region likely resulted from significant changes in intermediate water formation tied to the reorganization of oceanic conditions in the Southern Ocean, which may have in turn mainly controlled the global ocean N cycle, and therefore the N isotopic composition of nutrients since 3.5 Ma ago.

Late Quaternary planktonic foraminifera of ODP Hole 175-1087A

Planktonic foraminiferal assemblages from the upper Pleistocene part of Hole 1087A (0 to 12.1 meters below seafloor) are investigated to assess the role of global and local climate changes on surface circulation in the southern Benguela region. The benthic stable isotope record indicates that the studied interval is representative of the last four climatic cycles, that is, down to marine isotope Stage (MIS) 12. The species assemblages bear a clear transitional to subpolar character, with Neogloboquadrina pachyderma (d), Globorotalia inflata, and Globigerina bulloides, in order of decreasing abundance, as the dominant taxa. This species association presently characterizes the mixing domain of old upwelled and open ocean waters, seaward of the Benguela upwelling cells. Abundance variation of the dominant foraminiferal species roughly follows a glacial-interglacial pattern down to MIS 8, suggesting an alternation of upwelling strength and associated seaward extension of the belt of upwelled water as a response to global climate changes. This pattern is interrupted from ~250 ka down to MIS 12, where the phase relationship with global climate is ill defined and might be interpreted as a local response of the southern Benguela region to the mid-Brunhes event. Of particular interest is a single pulse of newly upwelled waters at the location of Site 1087 during early MIS 9 as indicated by a peak abundance of sinistral N. pachyderma (s). Variable input of warm, salty Indian Ocean thermocline waters into the southeast Atlantic, a key component of the Atlantic heat conveyor, is indicated by abundance changes of the tropical taxon Globorotalia menardii. From this tracer, we suggest that interocean exchange was hardly interrupted throughout the last 460 k.y., but was most effective at glacial terminations, particularly during Terminations I and II, as well as during the upper part of MIS 12. This maximum input of Indian Ocean waters around the southern tip of Africa is associated with the reseeding of G. menardii in the tropical Atlantic.

Respiration rates and electron transport system activities of copepod species obtained during Maria S. Merian cruise MSM17/3

Respiration rates and electron transport system (ETS) activities were measured in dominant copepod species from the northern Benguela upwelling system in January-February 2011 to assess the accuracy of the ETS assay in predicting in vivo respiration rates. Individual respiration rates varied from 0.06 to 1.60 µL O2/h/ind, while ETS activities converted to oxygen consumption ranged from 0.14 to 4.46 µL O2/h/ind. ETS activities were significantly correlated with respiration rates (r**2 = 0.79, p = 0.0001). R:ETS ratios were lowest in slow-moving Eucalanidae (0.11) and highest in diapausing Calanoides carinatus copepodids CV (0.76) while fast-moving copepods showed intermediate R:ETS (0.23-0.37). 82% of the variance of respiration rates could be explained by differences in dry mass, temperature and the activity level of different copepod species. Three regression equations were derived to calculate respiration rates for diapausing, slow- and fast-moving copepods, respectively, based on parameters such as body mass and temperature. Thus, knowledge about the activity level and behavioral characteristics of copepod species can significantly increase the predictive accuracy of metabolic models, which will help to better understand and quantify the impact of copepods on nutrient and carbon fluxes in marine ecosystems.