Physical drivers of phytoplankton production in the southern Benguela upwelling system

Investigations of primary production (PP) were undertaken in the southern Benguela ecosystem during two research surveys in October 2006 and May 2007. Significant differences in environmental conditions, as well as biomass and PP, were observed between October and May. During October, integrated biomass and PP were significantly higher, ranging from 20.43 to 355.01 mg m⁻², and 0.71 to 6.98 g C m⁻² d⁻¹, respectively, than in May, where the range was 47.92–141.79 mg m⁻², and 0.70–3.35 g C m⁻² d⁻¹, respectively. Distribution patterns indicated low biomass and PP in newly upwelled water along the coast, higher biomass and PP in the mid-shelf region, while lower values were observed at and beyond the shelf edge. Latitudinal variations showed consistently higher biomass and PP in the St. Helena Bay region compared to biomass and PP south of Cape Town. During both surveys, phytoplankton communities were comprised primarily of diatoms and small flagellates, with no significant differences. Phytoplankton adaptation to environmental variability was characterised by increased P_m^B and E_k under elevated temperatures and irradiance, while no clear relationships were evident for α^B. Generalised Additive Models (GAMs) showed that photosynthetic parameters were all significant predictors of photosynthesis rates (P_z), with P_m^B being the most important, accounting for 36.97% of the deviance in P_z. However, biomass levels and environmental conditions exerted a much greater influence on P_z, with irradiance explaining the largest proportion (68.24%) of the deviance. Multiple predictor GAMs revealed that 96.26% of the deviance in P_z could be explained by a model which included nitrate, chlorophyll a, and irradiance.     

Microheterotrophic pathways in the southern Benguela upwelling system

During the 1980s, there was a major shift in the understanding of the structure of marine foodwebs. As a result, the microbial loop has been incorporated into the classical concept of the planktonic food chain. Heterotrophic bacteria and Protozoa have been shown to be important components of the plankton biomass in many parts of the world's oceans, and their role in the trophic dynamics of pelagic foodwebs has been studied intensively. In the southern Benguela, field, laboratory and modelling studies have been combined to calculate carbon and nitrogen fluxes through the microheterotrophic portion of the pelagic foodweb. A size-based simulation model incorporating recent hypotheses on the structure and functioning of the pelagic foodweb after upwelling predicts rapid growth of a phytoplankton community dominated by netphytoplankton and chain-forming nanophytoplankton cells. After nitrate-depletion the bloom declines, to be followed by a bloom of single nanophytoplankton cells dependent upon regenerated nitrogen. During conditions of high nitrate availability and a netphytoplankton-dominated community, mesozooplankton ingest 44 per cent of the total primary production through herbivory and 1,4 per cent through carnivory. During periods of low nitrate availability and a nanophytoplankton-dominated community, mesozooplankton ingest 0,6 per cent of the total primary production through herbivory and 5 per cent through carnivory. Food chains are longer and microheterotrophs are an important link between primary producers and the larger heterotrophs. Simulation results show that microheterotrophs are an important component of the pelagic foodweb, primarily as regenerators of nitrogen, which sustains phytoplankton growth, and as a food source for larger heterotrophs of the metazoan foodweb during the nanophytoplankton-dominated bloom.     

Mesopelagic fish derivatives in the southern Benguela upwelling region

Discussion of mesopelagic fish within the southern Benguela system necessitates recognition of both oceanic and neritic realms, demarcated by the continental shelf-break. In the eastern South Atlantic (FAO Area 47), the stock size of two oceanic, mesopelagic families (Myctophidae, Sternoptychidae) have been estimated at 8–12 million and 750 000 metric tons respectively. Each possesses pseudoceanic species, populations of which constitute a possible alternative fishery resource to conventional pelagic stocks. Of these upper-slope/shelf derivatives, Lampanytodes hectoris (Myctophidae) and Maurolicus muelleri (Sternoptychidae) are most abundant. In the southern Benguela system (Orange River mouth to Cape Point), both species appear to be restricted mainly to the region east of the offshore divergence, corresponding roughly to the continental shelf-break. Maurolicus muelleri has not been taken by the fishery, but purse-seine landings of Lampanyctodes hectoris have accounted for up to some 10 per cent of the total South African pelagic catch. Anomalies in the commercial catch (sexual disparity, sexual immaturity) have been linked to expatriatism. This phenomenon is re-examined in the light of more recent data from research sampling in the southern Benguela region, and alternative biological hypotheses are presented to explain the anomalies. It is suggested that the energy "bonus" derived from vertical migration in summer is enhanced both by a physical component, termed "cycle riding", and by a biochemical component, linked to long-chain wax esters. Seasonal variation in percentage lipid concentration also appears to be associated with reproductive activity in the late winter-spring spawning period. Vertical migration may be reduced at this time, in tandem with the cessation of upwelling within the system and paucity of food in the upper water column. Comparative data for Maurolicus muelleri are presented.     

Composition of elasmobranch landings in Bahrain

The first detailed elasmobranch fisheries data for the Kingdom of Bahrain are presented, based on surveys of fish markets in April 2012. At least 25 species were recorded, including undescribed taxa. The milk shark Rhizoprionodon acutus was the most frequently recorded species; together with the Arabian smoothhound Mustelus mosis and banded eagle ray Aetomylaeus nichofii, these species comprised 53% of individual abundance. Sharks were almost entirely small individuals <1 m total length (TL). Males of small shark species were largely mature, whereas nearly all individuals of larger sharks were immature. For several elasmobranch species, landings were significantly biased towards males, which were largely mature. The species assemblage showed some notable differences in composition to that of adjacent Qatar, sampled at the same time of year, highlighting the importance of local data collection.     

Phytoplankton composition and biomass across the southern Indian Ocean

Phytoplankton composition and biomass was investigated across the southern Indian Ocean. Phytoplankton composition was determined from pigment analysis with subsequent calculations of group contributions to total chlorophyll a (Chl a) using CHEMTAX and, in addition, by examination in the microscope. The different plankton communities detected reflected the different water masses along a transect from Cape Town, South Africa, to Broome, Australia. The first station was influenced by the Agulhas Current with a very deep mixed surface layer. Based on pigment analysis this station was dominated by haptophytes, pelagophytes, cyanobacteria, and prasinophytes. Sub-Antarctic waters of the Southern Ocean were encountered at the next station, where new nutrients were intruded to the surface layer and the total Chl a concentration reached high concentrations of 1.7 ??g Chl a L⁻¹ with increased proportions of diatoms and dinoflagellates. The third station was also influenced by Southern Ocean waters, but located in a transition area on the boundary to subtropical water. Prochlorophytes appeared in the samples and Chl a was low, i.e., 0.3 ??g L⁻¹ in the surface with prevalence of haptophytes, pelagophytes, and cyanobacteria. The next two stations were located in the subtropical gyre with little mixing and general oligotrophic conditions where prochlorophytes, haptophytes and pelagophytes dominated. The last two stations were located in tropical waters influenced by down-welling of the Leeuwin Current and particularly prochlorophytes dominated at these two stations, but also pelagophytes, haptophytes and cyanobacteria were abundant. Haptophytes Type 6 (sensuZapata et al., 2004), most likely Emiliania huxleyi, and pelagophytes were the dominating eucaryotes in the southern Indian Ocean. Prochlorophytes dominated in the subtrophic and oligotrophic eastern Indian Ocean where Chl a was low, i.e., 0.043-0.086 ??g total Chl a L⁻¹ in the surface, and up to 0.4 ??g Chl a L⁻¹ at deep Chl a maximum. From the pigment analyses it was found that the dinoflagellates of unknown trophy enumerated in the microscope at the oligotrophic stations were possibly heterotrophic or mixotrophic. Presence of zeaxanthin containing heterotrophic bacteria may have increased the abundance of cyanobacteria determined by CHEMTAX.     

Diet of sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The diet of sardine Sardinops sagax in the southern Benguela was investigated by microscopic examination of stomach contents. The relative dietary importance of prey size and prey type was assessed by calculating the carbon content of prey items. Sardine is an omnivorous clupeoid, ingesting both phytoplankton and zooplankton, with the relative importance of these two food types varying both spatially and temporally. Stomach contents were numerically dominated by small prey items, principally dinoflagellates, followed by crustacean eggs, cyclopoid copepods, calanoid copepods and diatoms. Virtually all prey items ingested by sardine were <1.2 mm maximum dimension, the particle size below which sardine only filter-feed. Despite the numerical dominance by phytoplankton, zooplankton contributed the major portion to sardine dietary carbon, small calanoid and cyclopoid copepods, anchovy eggs and crustacean eggs being the primary prey types. These results indicate that, like anchovy Engraulis capensis, sardine in the southern Benguela are primarily zoophagous, and contrast with earlier dietary studies on sardine in the region. However, the two species appear to partition their prey on the basis of size; sardine consume small zooplankton, whereas anchovy consume large zooplankton. This difference has been observed in other upwelling ecosystems where the two genera co-exist and is likely to contribute to the regime shifts observed between sardine and anchovy.     

热带西南印度洋上升流区季节和年际变化研究及与印太系统海气相互作用

用59年Ishii再分析温度资料,讨论了热带西南印度洋(SWTIO)上升流区的季节和年际变化以及与上升流区有关的温度距平的变化,同时分析了其与热带印太海气系统的关系,结果显示SWTIO 上升流在南半球冬、夏季比较强,春季最弱。它的范围在5°~1°S,在东西向从50°E可以伸展到90°E。该上升流区的变化与温跃层的温度距平有密切的关系,并存在明显的5 a振荡周期。SWTIO上升流区温度距平的5 a周期振荡是由热带东印度洋温度距平在最大垂直温度距平曲面(MTAL)上向西沿着11.5°~6.5°S传播过来的,它与热带太平洋的温度距平传播方式不同。SWTIO上升流是热带印太海气系统的一个重要组成部分,印度洋偶极子 超前SWTIO上升流区温度变化5个月,最大相关系数达到0.57,NINO3区指数超前SWTIO上升流区指数2个月达到0.49。当热带印太区域的大气风场改变,影响热带太平洋和印度洋表层SSTA,出现ENSO和DIPOLE,进一步向西传播到SWTIO次表层,导致SWTIO上升流区出现改变。     

Kinematics of the upwelling front off southern Africa

Mesoscale circulation features have been shown to play an important role in the cross-frontal mixing of upwelling cells, their frontal morphology and in their interaction with oceanic water masses. With three years of detailed thermal infra-red satellite information on the South-East Atlantic upwelling system available, it proved possible to present a preliminary study of four prevalent frontal features intrinsic to the short-term behaviour of upwelling in this area. Upwelling filaments are shown to extend between 50 and 600 km seawards of the main front and are found, as are upwelling plumes, predominantly off the recognized major upwelling cells. Frontal eddies have a range of diameters and are found distributed over the full area of upwelling and on both sides of the main upwelling front. Warm filaments of Agulhas Current origin are advected preferentially along the western border of the Agulhas Bank and follow closely the front of the southernmost upwelling cells, where they may play a catalytic role in the creation of frontal turbulence.     

Upper ocean high resolution regional modeling of the Arabian Sea and Bay of Bengal

In this paper, effort is made to demonstrate the quality of high-resolution regional ocean circulation model in realistically simulating the circulation and variability properties of the northern Indian Ocean(10°S–25°N,45°–100°E) covering the Arabian Sea(AS) and Bay of Bengal(BoB). The model run using the open boundary conditions is carried out at 10 km horizontal resolution and highest vertical resolution of 2 m in the upper ocean.The surface and sub-surface structure of hydrographic variables(temperature and salinity) and currents is compared against the observations during 1998–2014(17 years). In particular, the seasonal variability of the sea surface temperature, sea surface salinity, and surface currents over the model domain is studied. The highresolution model's ability in correct estimation of the spatio-temporal mixed layer depth(MLD) variability of the AS and BoB is also shown. The lowest MLD values are observed during spring(March-April-May) and highest during winter(December-January-February) seasons. The maximum MLD in the AS(BoB) during December to February reaches 150 m (67 m). On the other hand, the minimum MLD in these regions during March-April-May becomes as low as 11–12 m. The influence of wind stress, net heat flux and freshwater flux on the seasonal variability of the MLD is discussed. The physical processes controlling the seasonal cycle of sea surface temperature are investigated by carrying out mixed layer heat budget analysis. It is found that air-sea fluxes play a dominant role in the seasonal evolution of sea surface temperature of the northern Indian Ocean and the contribution of horizontal advection, vertical entrainment and diffusion processes is small. The upper ocean zonal and meridional volume transport across different sections in the AS and BoB is also computed. The seasonal variability of the transports is studied in the context of monsoonal currents.     

Wind-Wave Characteristics and Climate Variability in the Indian Ocean Region Using Altimeter Data

The significant wave height and wind speed derived for the period 1993–2010 from altimeter data sets over the Arabian Sea, Bay of Bengal, and the Indian Ocean categorized as six zones has been analyzed. The average variation of both significant wave height and wind speed is found to be almost stable for the period of study. The study reveals that the average wind speed increases by about 6cm/sec/year during monsoon and post monsoon in the southern Indian Ocean. The distribution of wind and waves was studied in the context of seasonal variations. In addition, the average inter-annual and intra-annual variations along with the statistical parameters such as standard deviation, and root mean square wave height for the six zones are also reported in this paper.     

Comparing trophic flows in the southern Benguela to those in other upwelling ecosystems

A balanced trophic flow model of the southern Benguela ecosystem is presented, averaging the period 1980–1989 and emphasizing upper trophic levels. The model is based largely on studies conducted within the framework of the Benguela Ecology Programme and updates the results of an expert workshop held in Cape Town in September 1989. Small pelagic fish other than anchovy Engraulis capensis and sardine Sardinops sagax, mainly round herring Etrumeus whiteheadi and mesopelagic fish, were important components of the food web in the southern Benguela. Severe balancing difficulties were encountered with respect to the semi-pelagic resources (hake Merluccius spp.) and demersal top predators (sharks), indicating the need for further research on the interaction of these groups with their ecosystem. The model is compared to other existing trophic flow models of ecosystems in major upwelling areas, i.e. the northern Humboldt Current (4–14°S), the California Current (28–42°N) and the southern Canary Current (l2–25°N), and to two independently constructed models of the northern Benguela ecosystem. These models are compared using network analysis routines of the ECOPATH software, focusing on the interactions between the five dominant fish species (anchovy, sardine, horse mackerel Trachurus trachurus capensis, chub mackerel Scomber japonicus and hake) that support important fisheries in all systems. The upwelling systems rank by size rather than species dominance. The ratio of catches and primary production differs between systems, partly because of differences in fishing regimes. Predation on the five dominant fish groups by other fish in the system was the most important cause of fish mortality in all models. Fishery catches are generally a larger cause of mortality for these groups than predation by mammals. The ecological cost of fishing appears to be comparatively low in the southern Benguela, because catches are low compared with the primary production, but also because the fishery is relatively low in the foodweb. However, in view of the very tight foodweb demonstrated in the model. it is likely that an increase in fishing pressure would cause severe trade-offs with respect to other components of the southern Benguela ecosystem.     

Identification and classification of vertical chlorophyll patterns in the Benguela upwelling system and Angola-Benguela front using an artificial neural network

Information on the vertical chlorophyll structure in the ocean is important for estimating integrated chlorophyll a and primary production from satellite. For this study, vertical chlorophyll profiles from the Benguela upwelling system and the Angola-Benguela front were collected in winter to identify characteristic profiles. A shifted Gaussian model was fitted to each profile to estimate four parameters that defined the shape of the curve: the background chlorophyll concentration (B ₀), the height parameter of the peak (h), the width of the peak (σ) and the depth of the chlorophyll peak (z_m ). A type of artificial neural network called a self-organizing map (SOM) was then used on these four parameters to identify characteristic profiles. The analysis identified a continuum of chlorophyll patterns, from those with large surface peaks (>10 mg m^?3) to those with smaller near-surface peaks (<2 mg m^?3). The frequency of occurrence of each chlorophyll pattern identified by the SOM showed that the most frequent pattern (～12%) had a near-surface peak and the least frequent pattern (～2%) had a large surface peak. These characteristic profile shapes were then related to pertinent environmental variables such as sea surface temperature, surface chlorophyll, mixed layer depth and euphotic depth. Partitioning the SOM output map into environmental categories showed large peaks of surface chlorophyll dominating in water with cool temperature, high surface chlorophyll concentration and shallow mixed layer and euphotic depth. By contrast, smaller peaks of subsurface chlorophyll were in water with warmer temperature, lower surface chlorophyll concentration, intermediate mixed layer and deep euphotic depth. These relationships can be used semi-quantitatively to predict profile shape under different environmental conditions. The SOM analysis highlighted the large variability in shape of vertical chlorophyll profiles in the Benguela. This suggests that an ideal typical chlorophyll profile, as used in the framework of biogeochemical provinces, may not be applicable to this dynamic upwelling system.     

Comparisons of surface Chl a and primary productivity along three transects of the southern South China Sea,northern Java Sea and eastern Indian Ocean in April 2011

Results are presented about the changes in chlorophyll a density, carbon fixation and nutrient levels in the surface waters of three transects of the southern South China Sea (SCS), northern Java Sea (...     

季风上升流对印尼爪哇岛南部海域pH影响的初步研究

基于2013年9月底至10月初在印尼爪哇岛南部海域调查得到的碳酸盐参数和相关水文数据,首次报道了该海域pH的分布,并重点探讨了南爪哇上升流(季风上升流)对其影响。结果显示上升流影响区表层pH低于周围非上升流影响区。通过两端元混合模型,定量讨论了上升流的物理输运和生物活动对pH的影响。研究表明,上升流的物理输运至少造成了海表层盐度增加0.4个单位,溶解无机碳(DIC)增加110μmol/kg,pH降低约0.2个单位;同时,在上升流区,强烈的生物活动(叶绿素a浓度大于0.4mg/m3)使得DIC的降低量达70μmol/kg,pH的增加量达0.15个单位。总体来看,该研究区域的物理输运作用大于生物作用,综合效应表现为DIC的增加和pH的降低。另外,同上升流的物理输运作用和生物作用相比,上升流引起的表层冷却和增盐对pH的影响较小(热力学作用)。     

On vertical velocity fluctuations and internal tides in an upwelling region off the west coast of India

Hourly fluctuations of vertical velocity in relation to components of flow and wind and temperature oscillations at a morring site in the shelf waters off the west coast of India are discussed. The vertical velocities were computed from a time series of vertical temperature profiles assuming that horizontal advection of temperature is negligible. The computed values at a depth of 40 m during the 72-h period of observation were of the order of 10⁻¹ to 10⁻²cm s⁻¹, with a mean value of −2·77 × 10⁻² cm s⁻¹ indicating a net upward movement of water. The computed vertical velocity showed fluctuations of about 2–3 h, in addition to weaker signals of about 12 h. Based on the spectral estimates, we speculate that these fluctuations of 2–3 h in the vertical velocity may be caused by the fluctuations in the along-shore wind. The oscillations of isotherms found in the temperaturedepth time series and the spectral estimates of temperature and cross-shore flow component showed a periodicity of about 12 h, which indicated the presence of semi-diurnal internal waves. The fact that these internal wave troughs were associated with the measured onshore flow suggested that the waves were propagating offshore. The computed stability parameters showed little evidence of instability or mixing. It was found that the isotherm troughs in the temperaturedepth time series at about 12-h period coincided with high vertical shear in the cross-shore direction and low values of Brunt Vaisälä frequency.     

Regime shifts in upwelling ecosystems: observed changes and possible mechanisms in the northern and southern Benguela

A regime shift is considered to be a sudden shift in structure and functioning of a marine ecosystem, affecting several living components and resulting in an alternate state. According to this definition, regime shifts differ from species replacement or alternation of species at similar trophic levels, whereby the ecosystem is not necessarily significantly altered in terms of its structure and function; only its species composition changes. This paper provides an overview of regime shifts, species replacements and alternations that have been observed in the northern and southern Benguela ecosystems over the past few decades. Bottom-up control, initiating and sustaining regime shifts or species replacements via environmental forcing, is documented for both the southern and the northern Benguela ecosystems. Fishing (a case of top-down control) appears to have played an important role in regime shift processes in the Namibian ecosystem. Very low biomass levels of exploited fish stocks associated with less efficient energy transfer in the northern Benguela are indicative of a regime shift. Very high biomass levels have been reached in the southern Benguela in the 2000s. However the alternation between sardine and anchovy that has been observed in the southern Benguela over the last two decades appears not to have had major effects on the overall functioning of the ecosystem. The consequences of regime shifts for exploitation are highlighted, suggesting that fisheries managers should move towards a more effective ecosystem approach to fisheries.     

Cetacean research in the southern African subregion: a review of previous studies and current knowledge

Cetacean research, in terms of the number of papers, and areas for which data are available, has expanded considerably in the southern African subregion in the past decade, especially in the South-West Indian Ocean. We review cetacean research within this subregion from the 1800s to the present to provide an overview of findings, investigate trends and identify knowledge gaps. Data are presented separately for large whales (those subject to commercial whaling) and smaller cetaceans, and are separated by era and ocean basin. Over 550 peer-reviewed papers and books were identified relating to research on cetaceans within the subregion. More than half (284) have been produced since 1990 and 193 relate specifically to South African waters. The most-studied species are those that are most accessible due to their coastal distributions (southern right whale Eubalaena australis: 45 papers, humpback whale Megaptera novaeangliae: 31 papers, killer whales Orcinus orca: 27 papers, Indo-Pacific bottlenose dolphin Tursiops aduncus: 30 papers, Indo-Pacific humpback dolphin Sousa chinensis (plumbea form): 25 papers) and/or were hunted commercially (sperm whale Physeter macrocephalus: 25 papers). Identified conservation concerns vary throughout the subregion, but include bycatch and directed hunts, oil and gas development, ecotourism activities, shifts in prey resources, and noise and chemical pollution. The inshore stocks of Bryde's whales Balaenoptera edeni, the Indo-Pacific humpback dolphin and the Atlantic humpback dolphin S. teuszii were identified as the populations of highest conservation concern, although there are considerable knowledge gaps relating to deep-water species and almost no data (even on species occurrence) are available for several areas and countries.     

Remotely sensed variability of temperature and chlorophyll in the southern Benguela: upwelling frequency and phytoplankton response

High-resolution (1km) satellite data from the NOAA AVHRR (Advanced Very High Resolution Radiometer) and OrbView-2 SeaWiFS (Sea-viewing Wide Field-of-view Sensor) are used to investigate the upper layer dynamics of the southern Benguela ecosystem in more detailed space and time scales than previously undertaken. A consistent time-series of daily sea surface temperature (SST) and chlorophyll a concentration images is generated for the period July 1998–June 2003, and a quantitative analysis undertaken. The variability in SST, upwelling and phytoplankton biomass is explored for selected biogeographic regions, with particular focus on intra-seasonal time scales. The location and emergence of upwelling cells are clearly identified along the length of the southern Benguela, being distinct on the narrow inner and the mid-continental shelves. Most notable is the rapidly pulsating nature of the upwelling, with intense warm/cold events clearly distinguished. The phytoplankton response to this physical forcing is described. Chlorophyll concentration on the inner shelf largely mirrors the pattern of SST variability, similarly dominated by event-scale processes. Over the mid-shelf, higher chlorophyll is observed throughout all seasons, although low biomass occurs during winter. The variability of the offshore extent of SST and chlorophyll is identified at locations of differing shelf width. Cooler upwelled water is confined primarily to the narrow inner-shelf, with event-scale pulses extending considerable distances offshore. Agulhas Current influences are readily observed, even on the Cape Peninsula inner-shelf. Chlorophyll concentrations vary considerably between the locations of differing shelf width. SST, upwelling and phytoplankton indices are derived for selected locations to quantify the intra-seasonal variations. The SST indices show marked temperature changes associated with rapid pulsation on the event scale. No strong seasonal signal is evident. In contrast, the upwelling indices display a strong seasonal signal, with most intense upwelling occurring in spring/summer in the south. The phytoplankton response to the seasonal upwelling index differs between the selected locations. This study concludes that, although low-resolution SST and chlorophyll data may be useful for investigating general patterns over large scales, higher resolution data are necessary to identify finer scale spatial and temporal variability, especially in the inshore coastal zones.