Phytoplankton biomass and production in the upwelling region of NW Africa. Relationships with hydrographic parameters

Phytoplankton biomass and production in the upwelling region of NW Africa and relationships with hydrographic parameters were studied. During the cruise of Atlor VII carried out in November 1975 in the upwelling region of NW Africa, measurements of chlorophyll a and primary production as derived from ¹⁴C uptake experiments were made at a total of 40 stations. Biomass and production showed the higher values on the shelf in the area of Banc d'Arguin and north of Cap Blanc. Production estimates in this area ranged between 1.4 and 3.2 g C m^-2 d^-1. There was a marked minimum in biomass and production at Lat. 21 ° N, in the zone of maximum upwelling intensity. With the exception of this minimum, the productive area coincided with the zone where surface temperature was lower than 18 ° C (indicating dominance of upwelled Central Waters) and nutrients were detectable in the upper layers. In the poorer offshore area there was a distinct subsurface chlorophyll maximum. The results are compared with those of previous cruises and some features of the seasonal cycle in the studied area are discussed.     

Formation and potential trophic significance of marine foam near kelp beds in the benguela upwelling system

The present study investigates the importance of foam in nearshore waters on the west coast of the Cape Peninsula (South Africa) as a possible food resource for consumer organisms. A bacterial density of 3.45x10⁹ cells ml^-1 foam suspension was recorded. Calorific values of up to 15.39 kJ g^-1 ash-free dry weight were noted, and a biochemical analysis of the drained foam suspension gave a composition of 22.85% protein, 10.76% lipid and 3.07% carbohydrates. Field data showed a correlation between peaks of phytoplankton up to 510 mg C m^-3 water temperatures and days of intense foam formation during periods of strong onshore winds. Experimental foam formation in the laboratory indicates that kelp mucilage and phytoplankton contain surface-active agents. An additional feature of kelp mucilage is its capability to improve foam stability. The comparison of the chemical composition of 12 and 120 h-old foam suspension indicates a loss of easily metabolizable components such as trichloroacetic-acid precipitated protein and neutral lipid with time. It is hypothesised that the losses are due to utilisation of these components by consumers.     

The role of fungi in processing marine organic matter in the upwelling ecosystem off Chile

In a study that spanned from March 2007 through November 2009, we report high fungal biomass and over 90% of extracellular enzymatic activity occurring in the size classes dominated by fungi during periods of high autotrophic biomass in surface waters of the upwelling ecosystem off central-southern Chile (36°30.80′S–73°07.70′W). Fungal biomass in the water column was determined by the abundance of hyphae and was positively correlated with the concentration of the fungal biomarker 18:2ω6. High fungal biomass during active upwelling periods was comparable to that of prokaryotes (bacteria plus archaea) and was associated with an increase in phytoplankton biomass and in extracellular enzymatic hydrolysis in waters from the depth of maximum fluorescence. We show fungi as a new microbial component in the coastal upwelling ecosystem of the Humboldt Current System off central Chile. Our results suggest that the temporal pattern in fungal biomass in the water column during a year cycle is a reflection of their capacity to hydrolyze organic polymers and, in consequence, fungal biomass and activity respond to a seasonal cycle of upwelling in this ecosystem.     

Copepod carcasses in the upwelling region south of cap blanc,N.W. Africa

High numbers of copepod carcasses were found in zooplankton samples taken in the upwelling system off N.W. Africa. The validity of the measurements is discussed. It is suggested that rapid changes in the environment, caused by the pulsation of violent mesoscale upwelling at the shelf edge, result in an increased mortality in zooplankton. Considerations are made on the role of copepod detritus in the upwelling region.     

Vertical migration and feeding of Euphausia lucens at two 72 h stations in the southern Benguela upwelling region

Diel vertical migration of Euphausia lucens was studied over 72 h at an inshore and at a midshelf station in the southern Benguela upwelling region during February 1991, using stratified net tows and by employing gut contents (chlorophyll/copepods) as tracers of historic position. Soon after population arrival in the surface layers of the inshore station (abundant chlorophyll), E. lucens became dispersed and redistributed over depth. This was accompanied by a rapid downward transfer of surface-collected chlorophyll. A continuous exchange of individuals between deep and shallow water was subsequently inferred, prior to population descent at around dawn. These data are in agreement with the hunger-satiation hypothesis and were shared by individuals at the midshelf station (low chlorophyll). Interestingly, the major part of the E. lucens population at the midshelf station was confined to deep water throughout 24 h. While this conflicts with the hunger-satiation hypothesis, such behaviour is required to both ensure population maintenance and avoid surface predation within the Benguela region. Although the examination of some recent literature suggests that asynchronous, hunger-satiation behaviours may be common amongst euphausiids in single-species assemblages, responses to the food environment may be constrained by other factors.     

Behavioural inertia places a top marine predator at risk from environmental change in the Benguela upwelling system

In variable environments, organisms are bound to track environmental changes if they are to survive. Most marine mammals and seabirds are colonial, central-place foragers with long-term breeding-site fidelity. When confronted with environmental change, such species are potentially constrained in their ability to respond to these changes. For example, if environmental conditions deteriorate within their limited foraging range, long-lived species favour adult survival and abandon their current breeding effort, which ultimately influences population dynamics. Should poor conditions persist over several seasons, breeding-site fidelity may force animals to continue breeding in low-quality habitats instead of emigrating towards more profitable grounds. We assessed the behavioural response of a site-faithful central-place forager, the Cape gannet Morus capensis, endemic to the Benguela upwelling system, to a rapid shift in the distribution and abundance of its preferred prey, small pelagic shoaling fish. We studied the distribution and the abundance of prey species, and the diet, foraging distribution, foraging effort, energy requirements, and breeding success of gannets at Malgas Island (South Africa) over four consecutive breeding seasons. Facing a rapid depletion of preferred food within their foraging range, Cape gannets initially increased their foraging effort in search of their natural prey. However, as pelagic fish became progressively scarcer, breeding birds resorted to scavenging readily available discards from a nearby demersal fishery. Their chicks cannot survive on such a diet, and during our 4-year study, numbers of breeding birds at the colony decreased by 40% and breeding success of the remaining birds was very low. Such behavioural inflexibility caused numbers of Cape gannets breeding in Namibia to crash by 95% following over-fishing of pelagic fish in the 1970s. In the context of rapid environmental changes, breeding-site fidelity of long-lived species may increase the risk of local or even global extinction, rendering these species particularly vulnerable to global change.     

Carbon and nitrogen exchange between sandy beach clams (Donax serra) and kelp beds in the Benguela coastal upwelling region

Carbon consumption and nitrogen requirements were estimated for populations of the sandy beach bivalve Donax serra on nine beaches of the west coast of South Africa. Subtidal populations composed mainly of adult clams were responsible for the bulk of standing stock (3538 g C m⁻¹), annual carbon consumption (13 444 g C m⁻¹ yr⁻¹), faeces production (6478 g C m⁻¹ yr⁻¹ ) and nitrogen regeneration (2525 g N m⁻¹ yr⁻¹). Kelp detritus, bacteria and kelp consumers' faeces available in the water column surpass several times the carbon and nitrogen requirements of intertidal and subtidal clam populations. Individual Donax serra pop ulations, in turn, may regenerate up to 3.2% of the total nitrogen requirements of all primary producers from kelp beds and 14% of the requirements of phytoplankton. These high standing stocks of clams are presumably supported mainly by organic matter originating from kelp which, in contrast to phytoplankton, is in constant supply and comprises the largest proportion of the annual production of particulate organic matter on this coast. Wide and shallow continental shelves with gentle slopes probably limit the penetration of upwelled waters to the nearshore waters, decreasing the influence of external inputs and increasing the importance of internal flows of nutrients and carbon within the nearshore zone. In this context, sandy beaches, rocky shores and kelp beds may be more closely interlinked compartments of a larger ecosystem encompassing the whole nearshore than traditionally thought. Received: 28 August 1996 / Accepted: 7 October 1996     

Trophic impact, metabolism, and biogeochemical role of the marine cladoceran Penilia avirostris and the co-dominant copepod Oithona nana in NW Mediterranean coastal waters

In this work we studied the trophic ecology and feeding impact of the cladoceran Penilia avirostris and the cyclopoid copepod Oithona nana, the two dominant zooplankters in the summer communities of the coastal NW Mediterranean, on the naturally occurring microbial communities. In order to ascertain carbon surplus for growth and reproduction and the contribution to carbon and nitrogen recycling of these two species, we also determined their basal metabolism and excretion rates. The experiments conducted during summers 2002, 2003, and 2004 indicate that P. avirostris grazed mostly upon small flagellates, dinoflagellates, and diatoms, whereas O. nana had a narrower prey range, selecting motile organisms such as ciliates and occasionally dinoflagellates. The grazing impact of both species accounted, on average, for <10% of the standing stock of the microbial groups considered. In spite of the oligotrophic conditions, the feeding activity of P. avirostris is in general sufficient to compensate basal metabolism and allows a surplus for growth and reproduction. This was not the case for O. nana, its daily rations being often lower than the carbon basal demands. Regarding excretion rates, both species presented different N:P excretion ratios, the ones of O. nana falling within values typical for copepods, whereas the absence of detectable phosphorus excretion by P. avirostris implied an unbalance recycling with respect to typical Redfield ratio composition of marine seston.     

Variability of particulate organic carbon and nitrogen in the Namibian upwelling system

Particulate organic carbon (POC) and nitrogen (PON) were analyzed during cruises undertaken in September 1985 and April 1986 along the Namibian upwelling system. The main objectives were to provide estimates of standing stocks of particulate organic matter (POM) and analyze its temporal and spatial variability. Average estimates of total carbon standing stock (0 to 100 m depth) accounted for 1.2 g-at C m^–2 during active and 1.32 g-at C m^–2 during abated upwelling. Zooplankton carbon was estimated as 0.22 and 0.27 g-at C m^–2, respectively, for both periods. POM was generally concentrated near the surface, especially during abated upwelling. During abated upwelling, POM was not only abundant inshore but also over the shelf, as a response of a more diffuse frontal system and a more strongly stratified water column. Cross-shelf gradients were more significant during active upwelling, while alongshore gradients accounted for most of the variance of particulate organic matter during abated periods. This result was a consequence of the seasonal intrusion of warm, Angolan water from the north during the period of minimum upwelling, and resulted in poorer POM concentrations and higher consumer: producer ratios (24.4%). Nevertheless, this last conclusion should be regarded with caution due to the lack of comparative interannual variability. A 48 h study at a fixed station permitted analysis of the daily variability in POM during the intrusion process. Changes in the thickness of the surface mixed layer due to irregular time-spaced pulses of non-homogeneous water masses resulted in sudden enrichments and renewals of phytoplankton and zooplankton populations in a matter of hours.     

Release of nitrite by marine dinoflagellates: development of a mathematical simulation

The dinoflagellates Scrippsiella trochoidea (Stein) and Alexandrium minutum (Halim) were grown in a light–dark cycle with nitrate or nitrate plus ammonium under three different nutrient-supply regimes (dilution with fresh media in dark phase only or during the entire light–dark cycle at the same daily dilution rate, or with a faster continuous dilution). When supplied with nitrate + ammonium, A. minutum released a proportion (as much as 100% from dark-fed cells) of the nitrate taken up during the dark phase as nitrite, reflecting a rate-limiting step at nitrite reduction and poor regulation of inorganic-N uptake and assimilation. S. trochoidea released much smaller amounts of nitrite, if any. Nitrate and ammonium were not accumulated to any extent by either species in darkness, and the transient increases in the size of the free amino acid pool were too small to explain the fate of the newly assimilated N. Thus uptake through to incorporation of N into macromolecules appeared to be coupled in these species, even in darkness when increasing glutamine:glutamate (Gln:Glu) ratios suggested rising C-stress. A mechanistic model was developed from an earlier ammonium–nitrate interaction model (ANIM) by the inclusion of an internal nitrite pool, with control over the supply of reductant for nitrite reduction linked to photosynthetic and respiratory components. The model can reproduce the release of nitrite seen in the experiments, and also the release of nitrite in response to nitrate-feeding of N-stressed cells reported elsewhere. Received: 22 August 1997 / Accepted: 26 September 1997     

GPS tracking a marine predator: the effects of precision,resolution and sampling rate on foraging tracks of African Penguins

We used a prototype GPS logger to track the movements of breeding African Penguins (Spheniscus demersus). The loggers also recorded temperature and water depth, which allowed us to reconstruct foraging tracks in three dimensions, although GPS signals are interrupted when the birds dive. Here we report the loggers performance in the field and assess the effects of GPS error, resolution and sampling rate on estimates of foraging track length and speed. There is a trade-off between sampling rate and battery lifespan. We tested loggers at sampling intervals of 1 s, 10 s, 1 min, 2 min and 10 min. Sampling less frequently increases the chance of tracking an entire foraging trip, but it slows uplink times, slightly decreases the accuracy of positional fixes, and significantly reduces the ability to measure fine-scale aspects of foraging behaviour. Compared with radio or satellite tracking, GPS loggers offer unprecedented detail about animal movements. The results of our analysis suggest that techniques that sample relatively infrequently, such as satellite tracking, underestimate actual track lengths by up to 50%. However, caution is needed when interpreting fine-scale sampling for relatively slow-moving organisms. Re-sampling 1-s tracks suggests that c. 35% of apparent movements at this scale are due to measurement error and, more importantly, the limited spatial resolution of GPS (1.85×1.54 m at the study area). We recommend that researchers use a 1-s sampling rate for fine-scale studies, but resample at less frequent intervals to remove spurious noise for slow-moving animals. At current levels of resolution, animals should move at least 4 m per sampling interval. We provide empirical correction factors to compare inferred track length sampled at different rates, but caution that these are idiosyncratic and strongly dependent on the animals behaviour. Overall, GPS loggers offer a significant advance for studies of fine-scale animal movement patterns.Communicated by O. Kinne, Oldendorf/Luhe     

Vertical and horizontal structure in the picoplankton communities of a coastal upwelling system

The distribution of prokaryotic and eukaryotic picoplankton in the west coast upwelling-region off the South Island of New Zealand was investigated during midwinter (1988) the time of year when several commercially important fish species migrate into the region to breed. Picoplanktonic cells were major contributors to the autotrophic biomass, with > 80% of the particulate nitrogen and 39 to 55% of the total chlorophylla contained in the <2µm size-fraction. The prokaryotic picoplankton concentrations ranged from 6.3 × 10⁵ to 2.1 × 10⁷ cell l^–1, and the eukaryotic picoplankton between 3.9 × 10⁵ to 1.2 × 10⁷ cells l^–1. Picoplankton numbers increased with distance offshore to a maximum of ~ 3.0 × 10⁷ cells l^–1 at ~ 35 km from the coast, and then diminished towards the outer shelf and open ocean. The ratio of prokaryotic to eukaryotic cells varied between 1.01 and 4.71 in the mixed layer. Both groups declined substantially beneath the pycnocline, with no evidence of deep maxima. Prokaryotic cells dominated the planktonic cell concentrations at all but two stations, but eukaryotic cells dominated picoplankton biovolume as a result of their larger average cell size. The prokaryotic to eukaryotic picoplankton cell-number ratios in this system were considerably lower than often recorded elsewhere, and were inversely correlated with nitrate concentration. These observations show that a eukaryoticdominated picoplankton community makes a substantial contribution to autotrophic biomass in this nutrient-rich upwelling system, and may thereby play a major role in the food-web dynamics of this coastal fishery.     

Appropriate use of regression analysis in marine biology

The Model I linear regression theory is often used in the analysis of data under conditions when the Model II theory is clearly needed. Implications derived from the use of the two theories can differ greatly when there is not a high degree of correlation between the X and Y variables. The geometric mean Model II method is easy to use, and is probably needed in the analysis of most field data, since the X variable in field data is rarely under the control of the investigator.     

Growth in urban extent and allometric analysis of West African cities

Urban area expansion is happening at much faster rates in Asian and African cities than elsewhere in the world. This study uses multi-temporal Landsat images to map the urban extent of six small to large cities in West Africa at four different time steps from the early 1970s–2010. The selected cities are Kumasi of Ghana, Daloa of Cote d’Ivoire, Abuja and Kano in Nigeria, Kindia of Guinea, and Ouagadougou of Burkina Faso. All cities revealed significant urban growth in both urban area and population; however, it was apparent that there was a lot of variability in urban area development. Exponential urban growth rates in the cities were measured as ranging between 0.026 and 0.077, with allometric scaling factors matching those of other countries.     

Bioengineers and their associated fauna respond differently to the effects of biogeography and upwelling

Temperature and primary production (often linked to nutrient supply) are two of the few factors influencing species diversity and abundances across mesoscale gradients, while at smaller scales the habitat complexity offered by bioengineers is important. Previous studies have illustrated effects of upwelling and biogeography on intertidal bioengineers, but it is not known if these processes influence assemblages associated with those bioengineers in a similar way. We examined the habitat structure offered by two species of mussels and their associated fauna in five regions across 3000 km and three biogeographic provinces of the South African coast, replicating upwelling and non-upwelling areas within each region. Upwelling and region influenced the structure of mussel beds (the density and size of mussels). In contrast, upwelling did not influence mesoscale differences in composition, abundance, and numbers of species of crustaceans, mollusks, or polychaetes in mussel beds. Regardless of trophic level or mode of reproduction, mussel bed fauna were influenced only by region. Regional differences were strongly influenced by biogeography. The associated fauna was, however, also strongly correlated with the structure of the habitats created by mussels. Our results support the importance of upwelling to a critical ecosystem engineer, but show that these effects do not extend directly to the assemblages of associated fauna, which are more influenced by regional-scale effects and biogeography. We suggest that mesoscale patterns in the associated fauna of this bioengineered habitat are driven by the direct effects of biogeography, combined with the influence of biogeography and upwelling on mussel bed structure.     

Tools for comprehensive estimate of coastal region marine economy potential and its use for coastal planning

For elaboration of strategy and spatial planning of sustainable coastal regions development it??s very important to know a real potential of development of the region. As parameter, which is comprehensively characterized marine economic, political and military power of the coastal region, it is possible to use the marine economy potential with comprehensive estimations of socio-economic, political, environment and military profits (damages) from coastal planning decisions. Some results, including the European and Russian coastal regions are presented.     

A detailed analysis of carbohydrates in marine particulate matter

Detailed profiles of organic carbon, organic nitrogen, carbohydrate and proteinous amino acid have been determined in particulate matter from various depths at a station in Sagami Nada off eastern Honshu, Japan. The profiles suggest that carbohydrate is decayed from particulate matter more rapidly than proteinous amino acid in the euphotic zone, while the latter is removed to a higher extent than the former in deep waters,which results in the increase of the C/N values of particulate matter in the depths. All of the particulate matter gave D-galactose, D-glucose, D-mannose, D-xylose and D-glucuronic acid upon acid hydrolysis. Vertical change of the monosaccharide composition of this particulate matter indicates that only D-glucose and its polymers are preferentially removed from it during its descent. Carbohydrates from the particulate matter of 20 m depth were fractionated into water-soluble and insoluble carbohydrates. Detailed analysis of these fractions indicates that 1,3-glucan and its related low molecular weight carbohydrates from the water-soluble fraction decay between 50 and 300 m depth, to leave waterinsoluble carbohydrates, immune to biological attack during the course of further sinking. On the basis of these facts, the effects of the biochemical nature of these carbohydrates on the vertical change of the particulate carbohydrate distribution are discussed.     

Lack of genetically-subdivided population structure in Bullia digitalis, a southern African marine gastropod with lecithotrophic development

Larval shell morphology in fossil and present-day gastropods is often used to infer modes of larval development and levels of dispersal. Dispersal ability influences not only genetic population structure, but also is thought to influence a species' geographical range and evolutionary duration. We tested these predictions in Bullia digitalis, a sandy-beach whelk, by examining genetic variability at 33 protein-coding loci in nine samples (N=739) taken in 1984 to 1985 at localities extending over about three-quarters of the geographical range of this species in southern Africa. Females of this species deposit eggs into benthic or brooded capsules in which larvae develop through the trochophore and veliger stages to emerge as crawling juveniles. Scanning electron micrographs confirmed a protoconch morphology typical for gastropods with lecithotrophic larval development. Contrary to expectations, subpopulations of B.␣digitalis had high levels of variability (H=0.102) and lacked a genetically-fragmented structure (=0.013). The lack of a genetically-subdivided population structure would not have been correctly inferred, if this species were known only from well-preserved fossil shells. Indirect estimates of migration between populations based on and the island model of migration, which assumes drift-mutation equilibrium, ranged between 19 and 23 individuals per generation. Either an undescribed mechanism of dispersal facilitates gene flow between populations, or the geographical range of this species has recently expanded to produce the appearance of high levels of gene flow. Gene-frequency distributions showed that relative to four other species of Bullia, populations of B. digitalis were in mutation-drift disequilibrium, with a significant excess of low-frequency alleles that is consistent with a recent rapid expansion from a small population. Also contrary to expectations, this species has a large geographical range (2 400 km) and an apparently long evolutionary history extending 5 to 20 million years, as estimated from an allozyme phylogeny with four other species of Bullia. Received: 15 January 1997 / Accepted: 28 January 1997     

Some development patterns of plankton communities in the upwelling areas of the Pacific Ocean

The principal trophic levels, each subdivided into groups of organismic elements, are distinguished in the planktonic communities of the Eastern Equatorial and the Peruvian upwellings. Production intensity or metabolism have been determined experimentally for all elements. A scheme is suggested for computing production from data on metabolism for all the elements of a community, as well as for computing net and real production and other functional characteristics for definite trophic levels and the community as a whole. Based on the quantitative estimation of the efficiency of primary production and other functional characteristics, the development of communities is divided into production and destruction periods; they are, in turn, subdivided into steps associated with a certain degree of water trophicity. The balance of net production of the communities in the Peruvian upwelling indicates that the excess production of a community above the shelf is utilized completely in the narrow (100 to 150 sea miles) band of off-shore water. This paper describes an attempt to trace the changes taking place in the functional characteristics of plankton communities and to compare them with the changes observed in the communities of the Peruvian and East-Equatorial upwellings.