The functioning of the ecosystem of the Banc d'Arguin,Mauritania: a review

The Banc d'Arguin, Mauritania, is an area of over 10 000 km² of shallow water and tidal flats between the Sahara and the upwelling system off the Mauritanian coast. The results of the Dutch-Mauritanian project Banc d'Arguin 1988 as well as the relevant literature are reviewed in order to develop a preliminary conceptual model of the functioning of this ecosystem. Hydrographically the Banc d'Arguin can be characterized as a large-scale negative estuary with higher salinities near the shore. It is concluded that ultimately the Banc d'Arguin ecosystem is fueled by nutrients and organic matter derived from the upwelling area. The inner part of the Banc d'Arguin system is dominated by a detritus-based benthic foodweb in which seagrasses are the principal primary producers. Little of the seagrass production seems to be exported to other parts of the system or other areas. Zooplankton and zoobenthos biomasses are relatively low, but nevertheless support high densities of consumers. Aquatic birds are especially numerous. The study identifies many research topics for the future.     

Food items of the grey mullet Mugil cephalus in the Banc d'Arguin area (Mauritania)

The stomach and gut content of 22 Mugil cephalus from the Banc d'Arguin shallows (Mauritanian coast, West Africa) is examined. Quartz grains are the predominant material found in the stomach together with small portions of benthic diatoms and flakes, aggregates of fine-grained inorganic and organic particles. As the composition of this material is modified by a sorting procedure during feeding it is difficult to conclude, whether seagrass stands (muddy and mixed sediments with sand fraction) or sandy flats are preferred as feeding habitats. The animals studied did not or not relevantly utilize the non-diatom microphytes, though the environment of the Banc d'Arguin is rich in microbial mats (cyanobacteria), Vaucheria beds and epiphytic vegetation of seagrass leaves.     

Hydrology and underwater climate of the Banc d'Arguin,Mauritania: a review

The northern coast of Mauritania between Cap Blanc and Cap Timiris is characterized by a large variety of hydrological situations. At the western slope of the Banc d'Arguin, at a depth of about 20 m, the underwater climate is essentially of a temperate character. This is a highly unstable area due to the occurrence of upwellings and the movements of the intertropical front. Close to the coast the isolation of water masses by the very shallow Banc d'Arguin results in an increase of temperature and salinity. These factors show a gradient towards the South from Cap, St. Anne to the islands. In this region the underwater climate is tropical, a feature which is most clearly expressed in land-locked bays such as the Baie d'Arguin, the area around the island of Tidra, and the Baie de St. Jean. In the latter bay the conditions are most extreme (salinity > 80). Due to the coastal topography the area of the islands around Tidra is highly isolated. This area receives no fertilization from the upwellings offshore.     

Cockle emergence at the sediment surface: 'favourization' mechanism by digenean parasites?

The aim of the present work was to assess the effect of digenean trematodes on indirect mortality of the cockle Cerastoderma edule, an infaunal bivalve. The tested hypothesis was that parasites altered the burrowing capacity of cockles and thus exposed them at the sediment surface, where they are more vulnerable to predators. If the predator is the final host, this mechanism, which drives the cockle out of the sediment, is considered as a 'favourization'. Cockle populations from 2 stations in Arcachon Bay (France)-Banc d'Arguin (oceanic situation) and La Canelette (lagoonal situation)--were sampled for 1 yr. At La Canelette, monitoring every 2 d showed that 50% of adult cockles regularly migrated to the sediment surface at a rate of 5 cockles m(-2) yr(-1) and disappeared in a few days. In the laboratory, 67% of these 'surface cockles' did not burrow again, suggesting that they would die in the field. Moreover, mortality measured after 7 d in the laboratory was 2 to 5 times higher than mortality of 'buried cockles', at both stations and particularly during summer. Species richness and abundance of digeneans from both stations were compared in 'buried' and 'surface' individuals to determine whether parasites played a role in cockle migration and mortality. Ten and 9 digeneans were found at Banc d'Arguin and La Canelette, respectively, with Himasthla quissetensis and Labratrema minimus being the most prevalent and abundant species at both stations. The abundance of H. quissetensis was slightly higher in surface cockles at Banc d'Arguin, but the difference fluctuated with station and cockle age (or size). L. minimus prevalence was only higher in surface cockles at La Canelette. In the latter station, we estimated that L. minimus and H. quissetensis were responsible for the emergence of 9 and 2%, respectively, of the buried cockles. Although this favourization mechanism may induce some mortality in cockles, it does not alone explain the magnitude of the observed mortalities (41 and 57% at La Canelette and Banc d'Arguin, respectively). A correspondence analysis did not show the presence of a particular parasite community in buried or surface cockles, which could explain these high surface cockle mortalities in association with the 2 dominant digeneans.     

First record and description of metacercariae of Curtuteria arguinae n. sp. (Digenea: Echinostomatidae), parasite of cockles Cerastoderma edule (Mollusca: Bivalvia) in Arcachon Bay, France

A new Himasthlinae species, Curtuteria arguinae, is described as metacercariae from the cockle Cerastoderma edule (L.), collected at Banc d'Arguin (southwestern France). These metacercariae encysted preferentially in the mantle and also in the foot of cockles. Encysted and chemically excysted metacercariae were studied by light microscopy and scanning electron microscopy, respectively. Excysted metacercariae were elongated and curved ventrally. They bore a 33-spine circumoral collar. Sensory papillae were arranged around the oral sucker and also symmetrically along the ventral surface body, from the collar to the acetabulum. The dorsal and ventral tegument surfaces were densely packed with similar pointed spines. The posterior end of the body was without any spines. Among the Curtuteria species described previously, only Curtuteria haematopodis Smogorjewskaja and Iskova, 1966 had the same number of circumoral collar spines. A 6-yr field survey showed that the cockle population at Banc d'Arguin was subjected to a summer infection of C. arguinae. Curtuteria arguinae phenology of infection is characterized by interannual variability and seasonality (beginning in July-August and maximum in autumn). The first intermediate and final hosts remain unknown.     

Hydrographical observations west of the Banc d'Arguin,Mauritania, in May 1988

The present paper discusses the distribution of hydrographical parameters in Mauritanian waters between 19° 20 N and 20° 40 N. Data were collected during the first leg of the Dutch Mauritania Expedition in May 1988 with the R/V TYRO and comprise measurements of water temperature (T), salinity (S) and dissolved oxygen contents (O₂) in the area off the Banc d'Arguin. The time of investigation fell in the transitional period between the cold and warm season, which in this area is characterised by a peak in the trade wind speed and progressive heating of waters. Wind speed and sea surface temperature distribution in May 1988 were close to the seasonal averages which suggests that the observed hydrographical conditions were normal for this period as well. The highest spatial variability of T, S and O₂ was found near the southern end of the Banc d'Arguin, where the combined effect of coastal upwelling and seasonal heating made the thermocline upper limit rise towards the surface and upwelling waters reach the shallowest parts of the shelf. In the northern part of the study area upwelling phenomena were observed in the subsurface layers below 50 m depth; upwelling water did not reach beyond the outer edge of the shelf. The analysis of the TS diagram showed that water masses in the upper 300 m layer were mainly composed of South Atlantic Central Water with an admixture of warm, low-salinity surface water of Guinean origin. Traces of North Atlantic Central Water were found in the subsurface layer and at the deepest levels of measurements. Almost no traces of warm, saline water originating from the shallow Banc d'Arguin were registered during the survey.     

The benthic infauna and benthic respiration off the Banc d'Arguin (Mauritania,Northwest Africa)

In May 1988, a study was made of the benthic community structure and benthic respiratory activity along two transects across the Mauritanian shelf off Banc d'Arguin. The main emphasis of the present paper is put on the macrobenthos, but results of a gross analysis of the meibenthos are discussed as well. Macrofaunal and meiofaunal density showed no consistent decrease with distance from the shore. The composition of the macrofauna taxa appeared to be closely correlated with sediment parameters and less so with depth. The highest macrofaunal biomass was found at the northern edge of the Banc d'Arguin, and the lowest biomass along the outer shelf. The biomass levels on the shelf match the ones reported for the northern Cap Blanc area where, in contrast to the seasonal upwelling in the investigated area, upwelling takes place all year round.Benthic respiration rates on the offshore shelf were relatively high at the nearshore stations and low near the shelf break. Experimental evidence suggests that the low O₂ levels in the upwelling water covering part of shelf, inhibited benthic respiration. The high respiratory activity at some northern inshore stations coincided with the presence of oxygen-rich coastal water. In this area benthic respiration surpassed the level previously reported for the enriched Cap Blanc area. On the basis of our respiration data, an estimate is made of the total carbon demand of the benthic community on the shelf in May and this quantity is compared with the measurements of daily primary production.     

The trawlfauna of the Mauritanian shelf (Northwest Africa): density,species composition,and biomass

In May 1988, epifaunal trawlsamples were collected at 17 stations on the Mauritanian shelf off the Banc d'Arguin. The species composition and biomass of the invertebrates and demersal fish were analysed. On the basis of species abundances, three station groups could be distinguished each corresponding to a subregion of the shelf: an inner shelf, a midshelf and an outer shelf station group. The average depth increased from 20 m for the inner to more than 100 m for the outer shelf stations. The biomass of mobile epifauna and fish was lowest on the inner shelf stations where the trawlfauna was dominated by large sessile filterfeeders. The trawlfaunas of the mid- and outer shelf were more similar in species composition. Demersal fish and crustaceans appeared to be more abundant on the outer shelf. The increase in density and biomass from inner to outer shelf is discussed in relation to the distribution of the infauna and the hydrographical conditions on the Mauritanian shelf. Being a demersal fishing area, density and biomass of the epifauna on the shelf are compared with that of the demersal fishing grounds in the southern North Sea. The result of this comparison shows that, except for echinoderms, all taxa attain higher densities and biomasses at the Mauritanian shelf.     

Seabirds in the shelf edge waters bordering the Banc d'Arguin,Mauritania, in May

Seabird distribution was mapped over the shelf off the Banc d'Arguin, Mauritania, in May 1988 at the end of the upwelling season. Storm-petrels (Oceanitidae), were the most numerous with an average density of 14.5 km^–2. They were most abundant over water-patches rich in primary production or zooplankton, but the simultaneous presence of trawlers in these areas made it impossible to relate seabird density directly to hydrography. The trawlers attracted the seabirds and their waste served as an important source of food. The most numerous species in the area was the Wilson's Storm-petrel Oceanites oceanicus. Northern-hemisphere migrants, including summering sub-adults also used the area in numbers, and they too, profited from the fisheries. The Royal Tern Sterna maxima was the only local breeding seabird reaching the shelf slope area.     

Patches of barnacles and ascidians in soft bottoms: Associated motile fauna in relation to the surrounding assemblage

Epibenthic patches dominated by barnacles Balanus crenatus Bruguiere and solitary ascidians Styela spp., Bolthenia echinata (L.) and Molgula spp. in the White Sea shallow subtidal develop on bivalve shells and small stones surrounded with muddy sand. The space between barnacles and ascidians is filled with muddy sediment inhabited by motile taxa. We hypothesized that (i) epibenthic patches and unstructured sediment would attract different motile fauna and (ii) motile fauna of the patches would be affected by local abundances of epibenthic foundation species. Most dominant motile species demonstrated a significant difference in abundance between the two microhabitats. In contrast to the fauna of the sediment, species composition observed in aggregations of barnacles and ascidians was stable across different locations. In the field experiment initially clear bivalve shells after 5 years of exposure developed barnacle clusters with motile fauna similar to that observed in natural aggregations. Amphipods, isopods and bivalves, capitellid polychaets, Cirratulus cirratus (Müller) and Pholoe minuta Fabricius (Polychaeta) dominated in the sediment inside epibenthic patches. The proportion of capitellids, known to be sensitive to organic enrichment, was much higher within the patches than outside. The abundances of motile taxa found in aggregations were mostly determined by the number of barnacles of different size and of their empty shells, the biomass of ascidians, and the effect of location. Different dominant species demonstrated sensitivity to different parameters.Physical structure of the habitat, provided by barnacles and ascidians, as well as their biodeposition activity are regarded as the main factors structuring the motile fauna in the community studied. The spatial pattern observed seems to imply a range of pattern-generating biogenic processes, similar to those previously revealed in patches of filter-feeding bivalves, tube-building worms and seagrass.     

Evolutionary ecology of nine sympatric species of the pelecypod Limopsis in Cretaceous chalk

The ecology and functional morphology of nine species of Limopsis that lived together in muddy bryozoan bioherms in the Cretaceous of Denmark are discussed. Two independent lines of limopsids entered the white-chalk environment: a non-byssate line represented by one flat circular burrowing species and abyssateline which gave rise to the formation of eight species. Seven of these, ranging from a large globose infaunal species to a small mytiliform epibyssate monomyanan, constitute a morphocline. The morphocline reflects adaptations along an environmental gradient, possibly related to substrate size and water flow. The morphocline contains two dominant species. One of these is epifaunal and the other infaunal, which demonstrates a trophic-group distribution sensu Turpaeva, reducing competition. A third, infaunal, species - not a member of the cline - reached a level of dominance, too, which underlines the evolutionary distance between this species and those of the morphocline. The sympatrical formation of the species appears to have taken place through specific habitat selection, resulting in non-competitive units. The morphological modifications resulted in a series of species ranging from near isomyarian forms over increasingly heteromyarian species to a monomyarian species, a unique case within the arcids.     

Polychaete assemblages of a sub-arctic Newfoundland fjord: habitat, distribution, and identification

This study explores the association of 24 polychaete species with sandy and muddy habitats located in a sub-arctic fjord, and across Atlantic Canada including Labrador, Newfoundland, and the Gulf of St. Lawrence. Key characters used to facilitate species identification are also summarized. Within Bonne Bay, distinctive polychaete assemblages were associated with specific sediment types and polychaete species richness and density were significant both predictors of corresponding total (infaunal) density and species richness. Polychaetes were denser and more diverse in sandy sediments, partly because sandy locales were associated with the outer portion of the bay, and therefore were closer to the more productive and diverse Gulf of St. Lawrence. In general, species that occupied both sediment types were more widely distributed within Bonne Bay and across the region. The biogeography of most species also suggests that the Bonne Bay fauna is transitional between Labrador and Acadian biogeographic provinces.     

Zooplankton distribution in the coastal upwelling system along the Banc d'Arguin,Mauritania

In the framework of the Dutch Mauritania Expedition 1988, zooplankton was sampled in the beginning of the spring upwelling season at 44 stations along the Banc d'Arguin, Mauritania. Distribution patterns of oceanic and neritic copepod species and especially of the upwelling indicator Calanoides carinatus (Kröyer) were analysed in search for direct proof of circulation of ocean upwelling water over the inshore banks. The vertical temperature distribution at the stations pointed to upwelling in two different parts of the shelf area. According to the copepod data the two upwellings were of a different nature. A restricted area at the shelf edge near Cap Timiris appeared to represent a relatively stable coastal upwelling situation. Upwelled water present near Cap Blanc was found to have traveled quite recently over a considerable part of the shelf. In the central area between the two upwellings, Banc-water extended from the shelf into the ocean. The possible impact of the observed circulation of ocean water on productivity of the shelf benthic system is discussed.     

Periphyton density and shading in relation to tidal depth and fiddler crab activity in intertidal seagrass beds of the Banc d'Arguin (Mauritania)

Periphyton development was studied on microscopic glass slides and leaves of Zostera noltii Hornem. in an intertidal area in the Banc d'Arguin (Mauritania). The effects of shading, tidal depth and grazing activities by the fiddler crab Uca tangeri Eydoux were evaluated. For all experiments, periphyton ash content was high (52–93%) and ash-free dry weight ranged between 0.10–0.63 mg cm^–2. Slides accumulated more periphyton than leaves.Artificial shading (62–99%) for 13 days had no effect on periphyton densities on leaves. Increased tidal depth resulted in higher ash-free dry weight on slides, but in lower ash-free dry weight on leaves. Significant variation along the coastline also existed. The effect of fiddler crabs was studied using exclosures. Presence of fiddler crabs reduced periphyton density on slides, whereas light transmittance was increased. On leaves, no significant fiddler crab effect was found. This difference between leaves and slides was probably caused by a storm at the day before the end of the experiment, and by the higher periphyton density on slides as compared with leaves. As visual inspection during the experiment showed clear differences in appearance of leaves inside and outside the exclosures, the storm probably sloughed off mainly the older leaves, i.e. those on which the differences in periphyton cover were the highest.It is hypothesized that periphyton accumulation is higher with increased tidal depth, whereas fiddler crab grazing pressure also increases in this direction. The result is a decreased periphyton density with increased tidal depth.The presently found light extinction coefficients (mean 0.8 m^–1) and periphyton light attenuance (up to 25%) in Banc d'Arguin are not likely to affect seagrass leaf growth.     

The distribution and abundance of young fish in the Banc d'Arguin,Mauritania

The importance of the Banc d'Arguin (Mauritania) for juvenile fish has been investigated in September 1988. Fish have been sampled with a 2 m beam trawl at 50 locations in the coastal zone between Cap Sainte Anne and Cap Timiris. Water depth varied between 1 and 16 metres. Temperature and salinity at each station have been measured, as well as the weight of the catch and total length of the individual fish. The largest catches — in numbers as well as in weight — were obtained around the shallows of Tidra and Arel. This region has a large diversity in species, and contains large numbers of juvenile fish and cephalopods. The deeper areas seem to be much poorer in species, but average length and weight of the sampled fish are higher than around Tidra and Arel.     

Life in the arena: infaunal gastropods and the late Phanerozoic expansion of marine ecosystems into sand

Marine ecosystems have expanded into the infaunal realm below the surface of soft sediments throughout the Phanerozoic eon. During the Palaeozoic era, this expansion largely involved sedentary animals living in permanent resting places. Active sand‐burrowing animals colonized the infaunal environment later, but when this happened and when specialization for infaunal life evolved remain open questions. Here, phylogenetic evidence, fossil occurrences and previously established criteria for recognizing the sand‐burrowing habit in marine gastropods are used to determine how many gastropod clades became infaunal and when the transitions from surface‐dwelling to infaunal life occurred. At least 20, and as many as 35, clades (all but one of post‐Palaeozoic age) contain actively infaunal species. The overwhelming majority (15 of 20 clades) became infaunal during the Cenozoic, and clades with hundreds of infaunal species in the living fauna diversified beginning in the Early Miocene. The repeated evolution of, and specialization to, the sand‐burrowing habit by gastropods and other animals was enabled by increased habitat availability and higher marine productivity, and was necessitated by intensifying predation. As a result, the infaunal realm was transformed from a marine refuge to an integrated part of the marine biosphere in which high performance in locomotion and defence has become the norm.     

Heavy metals (Cu,Zn, Pb,Cd) in sediment,zooplankton and epibenthic invertebrates from the area of the continental slope of the Banc d'Arguin (Mauritania)

The concentrations of copper, zinc, lead and cadmium in the surface sediment (upper 5 mm) were generally higher in the silt fraction than in the bulk sediment. No significant geographical trend in the metal concentrations of the surface sediments was found, nor a correlation between concentrations in bulk sediment as well as in the silt fraction and the % silt could be established. In general, the metal concentrations in both bulk sediment and silt are lower, when compared to marine environments in other climatological regions.In zooplankton, the metal concentrations were relatively high: expressed in µg g^–1 on a dry weight (D.W.) basis, they ranged from 15–90 for copper, 70–580 for zinc, 12–55 for lead and 4–10 for cadmium.In epibenthic invertebrate species, both in crustaceans and bivalve molluscs, the concentrations of copper, zinc, and lead were in the same order of magnitude as compared to corresponding species from other geographical latitudes. Cadmium concentrations were relatively low, ranging from 0.13–0.42 µg g^–1 D.W. in the bivalve molluscs Pitaria tumens and from 0.04–0.27 µg g^–1 D.W. in the shrimp Processa elegantula. Also in the crab species Ilia spinosa, Inachus sp. and Pagurus sp., the cadmium concentrations were low, varying between 0.1 and 0.2 µg g^–1 D.W.No significant relation between the metal concentration in whole-body samples and sediment (either bulk or silt) was present. Also no gradient was apparent in concentrations in organisms sampled at different depths (5 to 200 m) along two off-shore transects perpendicular to the Banc d'Arguin. Data indicated lower metal concentration in epibenthic organisms from sampling stations along a northern transect (southwest of Cap Blanc) than in organisms from the southern transect, off Cap Timiris.Evidence was obtained for a considerable atmospheric input of heavy metals, in particular zinc and lead, in a certain area along the continental slope of the Banc d'Arguin.     

Shell size and shape in Madeiran land snails: do niches remain unfilled?

The distribution of shell heights and diameters in the mainly endemic Madeiran land snail fauna shows the bimodal pattern of high- and low-spired shells found in many other faunas. Field and laboratory studies show that shell shape is associated with the angle of substrate on which the snails crawl; as elsewhere, tall spired species use vertical surfaces or burrow in soft material. Flattened species predominate on horizontal surfaces, while globular species are less specific in their preferences. Detailed comparisons with the fauna of N.W. Europe show that the proportion of high-spired species in the Madeiran fauna is low, and large high-spired species associated with vertical surfaces are very few in number despite an apparent abundance of suitable habitats. Amongst low-spired species, one family, the Helicidae, dominates the Madeiran fauna. While the overall distribution of size in these species is much as in Europe, Madeiran helicids extend into smaller size classes than do those in Europe, and they appear to fill a gap in the scatter created by the absence of other families.
Non-endemic species, other than those strictly associated with man-made environments, are generally small in size. In the upper scatter, their size distribution parallels that of endemics, but in the lower scatter they constitute the whole of the smallest size classes.
The role of interspecific competition in determining these distributions is discussed. The range of helicid sizes is compatible with a relaxation of competition or predator pressure relative to other areas, but in the upper scatter there appear to be gaps in the range of size and shape expected despite a long period in which the fauna could evolve. This could indicate the existence of adaptive troughs blocking, or delaying, radiation over the full spectrum of size and shape.     

Ecological aspects of a silicified bivalve fauna from the Silurian of Gotland

The silicified Wenlockian (Silurian) bivalve fauna from MÖllbos, Gotland, is part of a life assemblage. The vast number of shells show unusual phenomena, e.g. shell repair, pearl and tumour formation, etc. A number of shells contain epibionts and bored, round holes. Presumptive predators of the bivalve community are discussed. Size-frequency distribution of the two most abundant species possibly reflects age classes. The fauna, comprising eleven species, is dominated by deposit-feeders (90 %). They exhibit niche diversification, including at least three different feeding levels within the sediment.     

The consortium of the sponge Lubomirskia baicalensis in Lake Baikal,East Siberia

The spatial distribution of the fauna associated with a branched sponge, Lubomirskia baicalensis, endemic of Lake Baikal has been quantitatively studied. The biomass and numbers of three amphipod species which inhabit the sponge correlate (linearly or non-linearly) with the weight of the sponge.