The influence of the Benguela upwelling system on Namibia's marine biodiversity

Namibia's marine environment falls within the Benguela system, an eastern boundary current upwelling system in the south eastern Atlantic Ocean. Conditions within much of this environment change continuously as a consequence of the upwelling of nutrient-rich water into the surface zone. In addition, irregular anomalies in temperature, oxygen concentration and salinity occur, particularly in shelf waters. These fluctuations, which are inherent in the functioning of the Benguela system, tend to favour the persistence of few, generalist species, while at the same time high productivity supports large abundances. This trend is evident in all the major marine habitats off Namibia, where diversity is often lower than in comparable habitats in the southern Benguela system off the west coast of South Africa. Namibia's marine environment is considered 'relatively pristine', although threats to biodiversity are posed by both natural and anthropogenic phenomena.     

Zooplankton of the northern Benguela region in a quiescent upwelling period

The general features of the horizontal and vertical distributionof the main zooplankton groups in the northern Benguela upwellingregion during a period of abated upwelling in April 1986 arediscussed. Three different types of water were detected: (i)an inshore strip of recently upwelled water; (ii) oceanic waterover the slope; and (iii) Angola Current water penetrating fromthe north. Abundance of the most representative zooplanktongroups, as well as of the total volume of zooplankton, was highestin the shelf region, particularly south of Walvis Bay. Zooplanktonwas most abundant in areas where phytoplankton concentrationswere high and offshore transport was low. There were significantdifferences between the abundance levels of copepods, euphausiids,chaetognaths, and fish eggs and larvae in the different layersof the water column sampled, with the highest concentrationsin the surface layers. The sharpest vertical gradients wereobserved at stations affected by Angolan water, where the thermoclinewas very strong. In contrast, inshore, where stratificationwas low, no such vertical gradients existed.     

A model of copepod population dynamics in the southern Benguela upwelling region

A simple population dynamics model was constructed to simulate temporal variability in the biomass of a dominant copepod Calanoides carinatus (Copepoda: Calanoida) along the West Coast region of South Africa. Calanoides carinatus is extensively preyed upon by the commercially important anchovy Engraulis capensis, thus variability in zooplankton production may serve as a useful predictor of variability in anchovy recruitment levels. The model developed here circumvents the need to include a large number of parameters because it uses satellite-derived estimates of chlorophyll a concentration and sea surface temperature as primary inputs. Abundance estimates necessary to initialize the model are readily obtainable from biannual research cruises. The model successfully simulates observed features of a copepod population's response to pulses of upwelling and is robust with respect to most of its parameters because minor changes in their values result in predictable changes in model output. The model showed greatest sensitivity to parameters that are difficult to determine empirically, such as predator-induced mortality rates. Gaps in our present understanding of the nature and scale of processes affecting copepod egg abundance, survival and viability in the southern Benguela system were identified as the dominant impediment to simulating copepod population dynamics in the region.      

Genetic structure of Sufflogobius bibarbatus in the Benguela upwelling ecosystem using microsatellite markers

The bearded goby Sufflogobius bibarbatus is an abundant endemic small fish species on the continental shelf of the northern Benguela. The goby habitat is characterised by generally low bottom oxygen concentrations that vary spatially and seasonally. In the present study of population structure, 13 samples of S. bibarbatus from inner and outer shelf areas between 19°S and 32°S were screened using ten microsatellite loci. The genetic data were analysed in relation to isolation by distance and depth. Furthermore, for the first time, this study examined genetic data in relation to bottom oxygen concentration at the sampling locations. The data show low but significant genetic heterogeneity (G-test; F_ST = 0.007, p < .05). There was weak but significant genetic differentiation along a latitudinal gradient across all sampling sites from 19.50°S to 32.37°S (Mantel test; r = .464, p = .001), but this disappeared when the southernmost sample was removed. On the other hand, a positive correlation of bottom oxygen concentration with pairwise F_ST (r = .336; p = .017) was observed among the sampling sites from the Northern Benguela shelf area. Overall, the data are complex but suggest that isolation by distance and bottom oxygen concentration may play a role in the genetic structuring of S. bibarbatus. The findings are discussed in relation to the species’ life history features and oceanographic characteristics of the Benguela upwelling ecosystem.     

Late Quaternary development of the Java upwelling system (eastern Indian Ocean) as revealed by coccolithophores

The paleoceanographic potential of coccolithophores was used to decipher the paleoproductivity changes in the eastern Indian Ocean during the past 300,000 years. Core SO139-74KL was taken at the seaward limit of a fore-arc basin of the Indonesian continental shelf located beneath the Java upwelling system. Coccolithophores occur in all samples, and total coccolith concentration exhibit distinct variations over the entire section. Peak abundances occur every 20,000 to 25,000 years with the highest peak at isotope stage 7. Abundances increase during the glacials but peak abundances also occur during interglacials. The preservation of coccoliths is good to moderate in most of the samples. The most abundant species is Florisphaera profunda with a mean relative abundance of 41.5% followed by Gephyrocapsa ericsonii and Emiliania huxleyi (EhuxGeric) and Gephyrocapsa oceanica. These four taxa dominate the assemblage throughout the core, forming on average 90.5% of the total assemblage. The species composition suggests that warm tropical conditions prevailed throughout the investigated time period indicating that temperature was not the driving force for the assemblage variations at this site. The geologic record for present-day and Holocene oceanographic conditions seemed to be predominantly characterised by high productivities in combination with an unstable water column. Indications for oligotrophic open ocean conditions were sparse. However, during most of the year oligotrophic conditions prevail and upwelling recurs only for a short time period but upwelling indicating proxies dominate the geological record. A contrasting fully oligotrophic scenario characterised by peaks in the abundances of total coccolithophores, Umbellosphaera irregularis, and in the percentage ratio of EhuxGeric to G. oceanica can be seen with a periodical recurrence every 20,000 to 25,000 years. Synchronously the records of the high productivity indicators total organic carbon and G. oceanica are characterised by distinct minima. We believe that upwelling was totally cut off during these times and oligotrophic conditions with a pronounced water column stratification prevailed throughout all seasons. An obvious correspondence between the shut down times of upwelling and insolation minima suggests that surface water conditions were driven by orbital forcing.     

Vertical migration and habitat partitioning of six euphausiid species in the northern Benguela upwelling system

The vertical distribution of the adults of six species of euphausiids,Thysanöessa gregaria, Nematoscelis megalops, Euphausiaamericana, E.gibboides, E.hanseni, Stylocheiron longicorne,as well as the larval stages of E.hanseni, were studied duringa 48 h cycle at a fixed station in the northern Benguela upwellingsystem. All the species, except T.gregaria and S.longicorne,proved to perform diel vertical migration, but both migratoryand non-migratory species appeared to be segregated in spaceduring night-time hours, regardless of potential prey (phytoplanktonand copepods). It is suggested that water column structure andhydrographic discontinuities caused by a warm, depth-localizedintrusion, as well as the reproductive strategies of particularspecies, are responsible for this pattern. Eggs and larval stagesof E.hanseni were concentrated near the surface, and the effectof the short-term pulses of the intrusion on their abundanceis discussed.     

Effect of food composition on egg production and hatching success rate of two copepod species (Calanoides carinatus and Rhincalanus nasutus) in the Benguela upwelling system

We have analysed the daily egg production (EPR) and hatchingsuccess rates of the calanoid copepods Calanoides carinatusand Rhincalanus nasutus as a function of nano- and microplanktonconcentration and composition in the northern Benguela upwellingsystem off Namibia. Food concentration explained 55% (R. nasutus)to 62% (C. carinatus) of the EPR variability. We found no relationbetween the residuals of the food concentration–EPR regressionand the percentage of the different taxonomic components ofthe nano- and microplankton. Nor was there a relation with theproportion of the diatom Skeletonema costatum that dominatedthe major blooms or with the number of nano- and microplanktonspecies. We conclude that food quality differences could notbe attributed to the relative composition of microplanktonicparticles of the different groups (i.e. taxonomic composition).     

Microbial communities associated with phosphogenic sediments and phosphoclast‐associated DNA of the Benguela upwelling system

The processes that lead to the precipitation of authigenic calcium phosphate minerals in certain marine pore waters remain poorly understood. Phosphogenesis occurs in sediments beneath some oceanic upwelling zones that harbor polyphosphate‐accumulating bacteria. These bacteria are believed to concentrate phosphate in sediment pore waters, creating supersaturated conditions with respect to apatite precursors. However, the relationship between microbes and phosphorite formation is not fully resolved. To further study this association, we examined microbial community data generated from two sources: sediment cores recovered from the shelf of the Benguela upwelling region where phosphorites are currently forming, and DNA preserved within phosphoclasts recovered from a phosphorite deposit along the Benguela shelf. iTag and clone library sequencing of the 16S rRNA gene showed that many of our sediment‐hosted communities shared large numbers of phylotypes with one another, and that the same metabolic guilds were represented at localities across the shelf. Sulfate‐reducing bacteria and sulfur‐oxidizing bacteria were particularly abundant in our datasets, as were phylotypes that are known to carry out nitrification and the anaerobic oxidation of ammonium. The DNA extracted from phosphoclasts contained the signature of a distinct microbial community from those observed in the modern sediments. While some aspects of the modern and phosphoclast communities were similar, we observed both an enrichment of certain common microbial classes found in the modern phosphogenic sediments and a relative depletion of others. The phosphoclast‐associated DNA could represent a relict signature of one or more microbial assemblages that were present when the apatite or its precursors precipitated. While these taxa may or may not have contributed to the precipitation of the apatite that now hosts their genetic remains, several groups represented in the phosphoclast extract dataset have the genetic potential to metabolize polyphosphate, and perhaps modulate phosphate concentrations in pore waters where carbonate fluorapatite (or its precursors) are known to be precipitating.     

Early Devonian (~410 mya) microfossils resembling Characiopsis (Tribophyceae) and Characium (Chlorophyceae)

Unusual microfossils that occurred associated with fungal spores in the Lower Devonian (~410 mya) Windyfield chert from Scotland were composed of a narrow stipe (2.5–9 μm long) to which was attached an obovoid or elongate drop‐shaped cell up to 14 μm long; a basal attachment pad was present in several specimens. The fossils were strikingly similar morphologically to certain present‐day unicellular freshwater Tribophyceae and Chlorophyceae, but affinities to the fungal phylum Chytridiomycota also cannot be ruled out. This discovery adds to the inventory of distinctive microbial morphologies in the early non‐marine paleoecosystems.     

Ammonium formation in cape timiris (Mauritania) upwelling

Ammonium formation was studied in two consecutive years by following a newly upwelled ammonium-free ‘parcel’ of water with a drogue. High concentrations (> 2 μg-at. NH₄-N l^?1) were found at the end of each study. The relative importance of WP-2 net mesozooplankton excretion on the ammonium values is greater before the phytoplanktonic bloom (35–48%) than during it (10–16%). From carbon grazing estimates, it is shown that other herbivore excretion does not take the place of WP-2 mesozooplankton and because of synchronous variations of chlorophyll with dissolved organic nitrogen, and bacterial activity with ammonium, it is suggested that much of the ammonium is produced from phytoplankton decay or nitrogen excretion during the bloom.     

The relative importance of food and temperature to copepod egg production and somatic growth in the southern Benguela upwelling system

The fecundity and somatic growth rates of Calanus agulhensisand Calanoides carinatus, the dominant large calanoid copepodsin the southern Benguela upwelling system, as well as the fecundityof several other common copepods, were measured between Septemberand March of 1993/94 and 1994/95. Mean egg production of mostcopepods was low at >30 eggs female^-1 day^-1 {Calanoides carinatus23.7, Calanus agulhensis 19.0, Neocalanus tonsus 16.1 and Rhincalanusnasutus 26.1), whereas the mean fecundity of Centropages brachiatuswas significantly greater (83.6 eggs female^–1 day^-1).This study also presents the first comprehensive field estimatesof the fecundity of Nanno-calanus minor (mean: 26.1 eggs female^–1day^–1, range: 0.0–96.2 eggs female^–1 day^–1)and of somatic growth of N6 and all copepodite stages of Calanoidescarinatus (decreasing from 0.58 day^–1 for N6 to 0.04 day^–1for C5). Somatic growth rates of Calanus agulhensis also declinedwith age: from 0.57 day1 for N6 to 0.09 day1 for C5. Data ongrowth rates were used to assess the relative importance offood [as measured by total chlorophyll (Chi) a concentration],phytoplankton cell size (proportion of cells >10 µm)and temperature to the growth of copepods. Multiple regressionresults suggested that fecundity and somatic growth rates werepositively related to both Chi a concentration and phytoplanktoncell size, but not to temperature. Although it was not possibleto separate the effects of Chi a concentration and phytoplanktoncell size, data from previous laboratory experiments suggestthat copepod growth is not limited by small cells per se, butby the low Chi a concentrations that are associated with theseparticles in the field. Despite growth not being directly relatedto temperature, a dome-shaped relationship was evident in somespecies, with slower growth rates at cool (<13°C) andwarm (>18°C) temperatures. The shape of this relationshipmirrors that of Chi a versus temperature, where poor Chi a concentrationsare associated with cool and warm temperatures. It is concludedthat the effect of food limitation on growth of copepods outweighsthat of temperature in the southern Benguela region. Sourcesof variability in relationships between growth and Chi a concentrationare discussed.     

The use of sea temperature in characterizing the mesoscale heterogeneity of phytoplankton in an embayment of the southern Benguela upwelling system

The role of meteorological events and hydrography in determiningchanges to the phytoplankton community was investigated in anembayment exposed to a narrow band of coastal upwelling. Dailysampling demonstrated the importance of advective processesdriven by meteorological forcing in controlling rapid shiftsin the biomass and species composition of the phytoplanktoncommunity. Samples of similar phytoplankton composition wereassociated with different stages of the upwelling cycle, asdefined by an index of biological ageing of upwelled waters.Relationships between the physical, chemical and biologicalfields were defined from time-senes measurements. The time elapsedfollowing upwelling, required for the determination of biologicalrates, was estimated from the rate of heating. A primary productionestimate of 2.14 g C m^–2 day^–1 was derived fromdetermination of the rate of nutrient depletion, whereas a phytoplanktonbiomass-nutrient consumption equation provided an estimate of3 92 g C m ^–2 day^–1. Both rates were within therange of estimates obtained from in vitro tracer methods.     

Reproductive response of the copepods Calanoides carinatus and Calanus agulhensis to varying periods of starvation in the southern Benguela upwelling region

Egg production by the calanoid copepods Calanoides carinatusand Calanus agulhensis fed excess Thalassiosira weissflogiiwas monitored in the laboratory following starvation periodsof 1, 3, 5, 7 and 9 days. Following short (1–3 day) periodsof starvation, egg production by C.agulhensis returned to thesatiated rate (51.1 eggs {female} day^–1) more rapidly(after 0.9–2.4 days of excess food) than that of Ca. carinatus(after 2.8–3.1 days). However, following longer (5–9day) periods of starvation, Ca. carinatus regained satiatedlevels of egg production (55.8 eggs {female}^–1 day^–1)more rapidly (after 3.1–4.0 days of excess food) thanC. agulhensis (after 3.8–5.2 days of feeding following5–7 days of starvation). Moreover, many C. agulhensisfemales did not regain normal rates of egg production after9 days of starvation. For both species, the time required foregg production to recover was proportional to the starvationperiod, although only up to 7 days for C. agulhensis, and wasthe same following 4.25 days of starvation. Previously fed Ca.carinatus terminated egg production more rapidly than C. agulhensiswhen starved. The ability of Ca. carinatus to tolerate, andrecover rapidly from, prolonged periods of starvation, combinedwith a comparatively fast development time and high rate ofegg production, provides this species with a strong competitiveadvantage over C. agulhensis in the highly pulsed food environmentof the southern Benguela upwelling region.     

Vertical migration and feeding of Euphausia lucens (Euphausiacea) in the Southern Benguela

Diel migration and feeding were examined for populations ofEuphausia lucens living in the near- and offshore waters ofthe southern Benguela. Euphausiids at both stations displayednocturnal diel vertical migration patterns. Animals inshoreascended in a slow-fast-slow manner which seems to be relatedto differential food concentrations throughout the water columnPronounced quantitative and qualitative changes in the dietof E.lucens were observed offshore but not inshore. These differencesare discussed in relation to different ambient food environments     

Mississippian (Lower Carboniferous) microfossils from the Chios Mélange (Chios Island, Greece)

The Chios Mélange is a thick Paleozoic wildflysch sequence that crops out on the Greek island Chios. It is composed of chert, limestone and volcanic blocks floating in a siliciclastic turbiditic matrix. New data suggest that the youngest blocks within the Chios Mélange are clasts of a breccia from the Kouramia-Nenitouria area that contain conodonts of late Visean or early Serpukovian age. The conodont fauna from the breccias is characterized by the genera Gnathodus and Lochriea, which favored deeper-water, open-marine facies. Elsewhere on the island, in the Papalia-Nagos area, calcareous microfossils of middle to late Visean age have been recovered from lime grainstone beds within the Chios Mélange. Allochems making up the grainstones are interpreted to have been transported into deeper-water turbiditic facies, but there is no evidence of stratigraphic reworking. Age-diagnostic calcareous microfossils include the alga Koninckopora inflata and foraminifers in the genus Paraarchaediscus. In situ Mississippian microfossils indicate that the Chios Mélange is older than Pennsylvanian (Upper Carboniferous) - Permian, as previously thought. The revised age suggests that the origin of the Chios Mélange may be related to the development of an accretionary prism during the Hercynian Orogeny.© 2003 Éditions scientifiques et médicales Elsevier SAS. All rights reserved.     

Late Cretaceous and Cainozoic bathyal Ostracoda from the Central Pacific (DSDP Site 463)

Cainozoic deep-sea ostracod assemblages from the summits of Mid-Pacific guyots point to high levels of endemism possibly as a result of their bathymetric separation from the surrounding sea floor. However, the interpretation of these fossil assemblages is hampered by the paucity of comparative material from surrounding non-guyot sites. Fifteen ostracod assemblages from DSDP Site 463 (Late Cretaceous-Pleistocene) were studied to compare with those from nearby guyots. Three distinct faunal assemblages are recognised at Site 463: Assemblage A (Maastrichtian-Eocene), Assemblage B (Oligocene-Upper Miocene) and Assemblage C (Upper Miocene-Pleistocene) although the palaeoenvironmental significance of these units is unclear. Sixty-two ostracod species are identified, the thirteen most abundant are discussed in the taxonomic section, five of which are described as new. Between 30 and 100% of the species encountered in each sample are considered as endemic to Site 463, while some of the remaining species were previously thought to be endemic to individual guyots. Similarly high levels of endemism on nearby guyots probably reflect an incomplete knowledge of deep-sea ostracod faunas rather than the establishment of geographically or bathymetrically restricted populations. The presence of globally pandemic and geographically widespread taxa on sites such as the Mid-Pacific Mountains, surrounded by abyssal depths which lie below the CCD, indicates that some faunal exchange or migration of ostracods does take place. This must be achieved within the intermediate waters and probably occurs passively.     

Calcareous microfossils from the Karakatty Formation (Lower Cambrian) of the Northern Tien Shan

Lower Cambrian calcareous microfossils that preserved monospores and basal attachment organs with rhizoids have been discovered in jaspers of the Karakatty Formation of the Northern Tien Shan. A new genus, Koroleviphyton Kolosov, is established in the red algae division Rhodophyta with the type species K. attenuescens (Kolosov, 1983).     

Organic geochemical changes in Pliocene sediments of ODP Site 1083 (Benguela Upwelling System)

The intensification of the Northern Hemisphere Glaciation (INHG) was a major event in the development of the current climate state, and as one of the most productive regions in the world's oceans, the behaviour of the Benguela Upwelling System (BUS) following the INHG is of wide interest. To investigate post-INHG changes in productivity and organic matter accumulation, total organic carbon and biomarker accumulation rates were determined for sediments from ODP Site 1083 and compared to alkenone-derived sea surface temperatures and nitrogen isotopic compositions. These data indicate that the interval between 2.6 and 2.4 Ma was characterized by dramatic changes in upwelling intensity and organic carbon export on the northern edge of the modern BUS. The upwelling is reflected by significant changes in alkenone-derived SST estimates between glacial and interglacial intervals, with a total variability of 16 °C. The studied interval is also characterized by large changes in organic matter export as reflected by changes in TOC and biomarker accumulation rates, which show maxima during OIS 98 and during the transition from OIS 97 to 96. Intervals of elevated TOC are also characterized by elevated concentrations of sedimentary microbial biomarkers and lower %CaCO₃, suggesting that enhanced delivery of labile organic matter to the seafloor resulted in enhanced remineralisation with released CO₂ being consumed by CaCO₃ dissolution. However, in apparent contrast to recent Pleistocene sediments at the same site, organic matter export after the INHG was not solely driven by upwelling intensity. Of the three Pliocene glacial–interglacial cycles examined (OIS 101 to 96), each is unique with respect to the timing and magnitude of changes in organic matter accumulation. Each is also characterized by different algal assemblages as inferred from biomarker distributions, with OIS 97 and 96 particularly dominated by diatoms. We suggest that these differences reflect the important but evolving role of Southern Ocean waters in the Pliocene BUS: nutrient depletion of SO waters occurred during parts of Pliocene glacial intervals such that even intense upwelling did not persistently result in enhanced organic matter accumulation rates.     

Rugose corals and associated carbonate microfossils from the Brigantian (Mississippian) of Castelsec (Montagne Noire, southern France)

The disused quarry east of Castelsec offers a view of shallow-marine carbonates of the poorly known Uppermost Mississippian of the Montagne Noire. At Castelsec, sections are studied in two characteristic facies types (bioclastic wackestone and microbial dominated boundstone) of the Upper Mississippian. The succession is rich in rugose corals and carbonate microfossils. Six genera with seven species belonging to a rugose coral fauna consisting of at least eight genera with several species are described herein; Dibunophyllum castelsecensis sp. nov. is described as new. Twenty-seven carbonate microfossils of different groups have been identified. The Castelsec succession is Brigantian in age, based on the stratigraphic occurrence of rugose corals, foraminifers, and calcareous algae observed in both sections. The rugose coral fauna shows relationships with the well-known fauna of northwestern Europe and the Ouralian-Asian Province. Typical elements of northwestern Europe are missing at Castelsec and vice versa. This differentiation between north and south is interpreted as responses to different palaeolatitudes and tectonic settings.