Macrofauna distribution and bioturbation on tidal confluences of the dutch Wadden Sea

Macrofauna density and bioturbation intensity (measured with X-ray radiography), were studied in the Dutch Wadden Sea near Ameland and in Mok Bay, Texel, in September 1988. The bivalveMacoma balthica and the polychaeteHeteromastus filiformis were widely distributed in the areas studied. The cockleCerastoderma edule and the polychaeteScoloplos armiger did not occur in areas with high clay content, while the molluscsHydrobia ulvae andMya arenaria preferred fine sediments. The polychaeteNereis diversicolor preferred the higher parts of the intertidal. In the Ameland area disturbance of primary sediment structures was highest near the shore and near the middle of the tidal confluence where physical reworking was low. The sheltered Mok Bay sediments were completely reworked by deposit-feeders. Bioturbation intensity and deposit-feeder (bioturbator) density were, however, not highly correlated. A number of stations showed higher bioturbation towards the surface and this may be related to the high abundance of near surface dwelling macrofauna, due to eutrophication and organic enrichment of the sediments in recent years.     

Macrofauna on flood delta shoals in the Wadden Sea with an underground association between the lugwormArenicola marina and the amphipodUrothoe poseidonis

Living conditions for macrofauna on flood delta shoals are determined by surf, strong currents and sediment mobility. Thus, a unique assemblage of invertebrate species colonize these far off-shore, low intertidal flats. We here describe the macrobenthic fauna of emerging shoals in the Wadden Sea between the islands of Römö and Sylt. Besides ubiquitous macroinvertebrates of the intertidal zone and species which attain their main distribution in the subtidal zone, the flood delta shoals are characterized by organisms adapted to live in these highly unstable sediments, like the polychaetesSpio martinensis, Streptosyllis websteri, Magelona mirabilis, Psammodrilus balanoglossoides, the pericarid crustaceansCumopsis goodsiri, Tanaissus lilljeborgi, Bathyporeia sarsi and a few others. Average abundance (1440 m^?2 of ind >1 mm) and biomass (12.9 g AFDW m^?2) were low compared to other intertidal habitats in the Wadden Sea. Biomass was dominated by largesized individuals of the lugwormArenicola marina. The U-shaped burrows of these polychaetes were inhabited by high numbers ofUrothoe poseidonis. Maximum densities of these amphipods occurred in the deepest parts of the burrows. Sampling at approximately montly intervals revealed no apparent seasonality ofU. poseidonis abundance. Together with smallCapitella capitata, these amphipods constitute a deep-dwelling component of the macrofauna associated with lugworms, which is separated from all other macrofauna living at the sediment surface. As a response to rising sea level and increasing tidal ranges, we expect the unstable sandy shoals, inhabited by numerousSpio martinensis andUrothoe poseidonis, to expand within the Wadden Sea at the cost of stable sandy flats with abundant macrofauna.     

Temporal and spatial variations of biodeposits—A preliminary investigation of the role of fecal pellets in the Sylt-Rømø tidal area

The fecal pellets ofHeteromastus filiformis andHydrobia ulvae are abundant on the sediment surface of the Sylt-Rømø tidal flats. AlthoughCerastoderma edule is present in most parts of the study area, only small amounts of its pellets could be found. Fecal pellets are non-randomly distributed in the study area. Large amounts ofH. ulvae pellets are found in the mud flats and smaller amounts in the mised flats and sand flats, whereH. filiformis pellets predominate as biogenic component. This distribution pattern varies with time under the influence of environmental factors. Marked temperature fluctuations provide seasonal signals reflecting distinct patterns. The results show a net balance of biodeposit components in the sediment. This could be a result of the sheltered location of the area and this fits in well with the mostly stable sedimentological picture.     

Abundance and distribution of hydroids in a mangrove ecosystem at Twin Cays,Belize, Central America

Qualitative and quantitative collecting was undertaken in 1987 to determine the species composition, abundance, and distribution of hydroids in a mangrove system at Twin Cays, Belize. Of 49 species identified, the 5 most frequent were Ventromma halecioides, Nemalecium sp., Clytia hemisphaerica, Dynamena crisioides and Halopteris diaphana. Line-transect census data and qualitative observations showed that the hydroid fauna was sparse in sheltered, still-water areas of the mangal, but relatively abundant and diverse in areas exposed to waves and/or tidal currents. Species composition and relative abundance varied with depth at stations in exposed locations and in tidal creeks and channels. Although Turritopsoides brehmeri is known only from Twin Cays at present, it seems improbable that any of the species is restricted to mangrove ecosystems.     

A functional and morphological approach to evaluate the vertical migration of estuarine intertidal nematodes during a tidal cycle

We tested herein the hypothesis that exposure time significantly contributes to the vertical distribution profile of nematodes during a tidal cycle as a function of distinct feeding and locomotion behaviors, conditioned by body morphology. We showed that the vertical distribution profile of the slender with filiform tail, numerically dominant Terschellingia longicaudata is in fact significantly correlated with sediment changes induced by tidal variation. Conversely, none of the other nematode species showed unequivocal evidence of vertical migration. Horizontal spatial heterogeneity also influenced the vertical distribution of nematode associations, probably as a response to varying temperature and desiccation levels at the sediment surface. The resulting vertical profiles for individual or species groups are a trade-off among locomotory and feeding strategies and concordant morphological adaptations.     

Historical changes in the benthos of the Wadden Sea around the island of Sylt in the North Sea

The situation regarding the distribution and abundance of seagrass, macroalgae and benthic fauna near the island of Sylt in the south-eastern North Sea during the period 1923 to 1940 is compared with that of the 1980s. Evidence of organic enrichment in recent times is provided by (1) massive growth of green algal mats on sheltered tidal flats, (2) a decline of red algae in the subtidal zone, (3) an expansion of mussel beds along low water line and down to 20 m depth, (4) increased abundance of polychaetes inhabiting intertidal and subtidal sandy bottoms. Seagrass beds have undergone complex changes which remain unexplained. Intensified erosion has contributed to the loss of habitats in the intertidal zone, and probably affected sessile epifauna in the deep channels. Here, direct removal and disturbance by the bottom-trawling fishery may also have contributed to the observed species impoverishment. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

The distribution of living benthic foraminifera on the continental slope and rise off southwest Norway

Surface sediment samples taken by ? corer from 45 stations on the Norwegian continental margin and in the Norway Basin have been investigated for their benthic foraminiferal content. Unlike previous studies, the living benthic foraminiferal fauna was differentiated from empty tests comprising the foraminiferal death assemblage. Factor analysis of both the living and dead faunal data reveals six living species assemblages and five corresponding dead assemblages. The additional living assemblage is characterized by the arenaceous speciesCribrostomoides subglobosum that dominates between 1400 and 2000 m water depth, but is rare in the dead faunal data.Trifarina angulosa and, to a lesser extent,Cibicides lobatulus characterize the shallowest foraminiferal assemblage from 200 to 600 m water depth, where it is associated with strong bottom currents and warm, saline Atlantic water of the North Atlantic Drift. On the slope between 600 and 1200 m water depth, theMelonis zaandami Species Assemblage dominates, particularly in areas characterized today by rapid sedimentation of terrigeneous material. Between 1000 and 1400 m depth, where the slope is covered by fine grained, organic-rich, terrigeneous mud, the living foraminiferal assemblage is characterized byCassidulina teretis andPullenia bulloides. Below 1400 m, three foraminiferal assemblages are found:C. subglobosum is found from 1400 to 2000 m,Cibicidoides wuellerstorfi andEpistominella exigua predominantly live from 2000 to 3000 m water depth, and below 3000 m,Oridorsalis umbonatus andTriloculina frigida dominate the fauna.All of theElphidium excavatum tests found in this study and theCassidulina reniforme tests found above 500 m water depth were found to be reworked.Analysis of the sediment grain-size distribution and the organic carbon content in surface samples from the deepest stations suggest that the abundance ofC. wuellerstorfi andE. exigua is positively correlated to relatively coarse (caused by planktic foraminifera) and organic-rich sediments, whereas high frequencies ofO. umbonatus andT. frigida coincide with low organic carbon content. We suggest thatC. wuellerstorfi is adapted to deep-sea environments with relatively high food supply, tolerating relatively low interstitial water oxygen content, whereasO. umbonatus may tolerate lower food supply prefering well-oxygenated interstitial waters.     

The effect of the dung pad fauna on the emergence ofMusca tempestiva [Dipt.: Muscidae] from dung pads in southern France

The emergence ofMusca tempestiva Fallèn, from cow pads in southern France was reduced by a mean of 98% and 96% respectively in nine experiments concluded in 1979 and 1981, by the action of the complete dung fauna. Fauna emerging from dung pads and captured in simultaneous trapping using dung baited pitfall traps consisted of 13 Dipterous species (9 Muscidae, 2 Sepsidae and 2 Sphaeroceridae), 25 Coleopterous species (7 Scarabaeidae, 1 Geotrupidae, 2 Aphodiidae, 3 Hydrophilidae, 12 Staphylinidae), 3 Hymenopterous parasites and 1 species of macrochelid and 2 species of parasitoid mites carried phoretically by the dung beetles. The total number of insects and mites per trap was usually small. The Staphylinidae which included 4Aleochara species of whichAleochara tristits Gravenhorst andA. bipustulata (L.) are known parasitoids of Diptera pupae, and 6 predatory species, were the most abundant insects present. Whilst the overall abundance of insects and mites trapped was relatively low it is probable that the most important factor in the control ofM. tempestiva was the combined influence of the different elements making up the dung pad fauna.     

Sewage,green algal mats anchored by lugworms,and the effects on Turbellaria and small Polychaeta

On sandy tidal flats at the Island of Sylt (North Sea) ephemeral mats of green algae covered wide areas in the vicinity of sewage outflows. Algae became anchored in the feeding funnels of lugworms (Arenicola marina) and thus were able to resist displacement by tidal currents. Below the algal mats anoxic conditions extend to the sediment surface. After about one month a rough sea removed all algae. Polychaetes endured this short-term environmental deterioration, while the more sensitive Turbellaria decreased in abundance and species richness. Diatom-feeders were affected most, predators to a medium extent, and bacteria-feeders the least affected. Rare and very abundant species were more affected than moderately abundant ones. None of the turbellarian species increased in abundance and none colonized the algal mats above the sediment. In a semicontrolled experiment with daily hand-removal of drift algae from a 100-m² plot within an extensive field of algal mats, this cleaned "island" served as a refuge to Turbellaria escaping from their algal covered habitat. Here abundance doubled relative to initial conditions and was 5-times higher than below algal mats.     

Faunenverarmung in einem Schlickgebiet in der Nähe Helgolands

Variations in abundance and species composition of macro-zoobenthos were studied at a sublittoral muddy-bottom station in the inner part of the German Bight from 1969 to 1976. The fauna of this area can be included in the classicalAbra alba community. A total of 98 species were found in 62 samples. Of these, only 4 species occurred in all samples:Nucula nitidosa, Diastylic rathkei, Ophiura texturata andNephtys hombergii. Species richness has declined since 1969. Populations of several species disappeared in the investigated area and its surroundings. Other populations are shown to exhibit remarkable fluctuations in abundance with sometimes heavy spatfalls or sudden breakdowns in population density, mainly during late summer. The long-term trend of impoverishment of the fauna is attributed to the development of anaerobic conditions in the muddy sediment. Such conditions are favoured by stable, thermohaline strafications of the water during summer, resulting in low oxygen levels and, probably, enrichment of hydrogen sulfide even in near-bottom water. Moreover, both continuou influx into the area of Elbe water, rich in suspended matter and nutrients and dumping of sewage sludge about 8 km east of the investigated area are considered to have increased the strees on the bottom fauna.     

Evidence for tidally-induced vertical migration of some gelatinous zooplankton in the Wadden Sea area near Sylt

Gelatinous zooplankton is more abundant in the Wadden Sea area of North Sylt than in the adjacent North Sea. The hypothesis is tested that medusae maintain their position in the North Sylt Wadden Sea by ascending to the surface at flood tides and descending to the bottom during ebb tides, thus avoiding the strong ebb currents which could carry them out of this food-rich area. Surface sampling at a main tidal channel revealed great differences between high tide and low tide abundances of five species of medusae in the surface layer.Rathkea octopunctata, Sarsia tubulosa, Eucheilota macultata andPleurobrachia pileus were all more abundant around high tide than during low tide. Bongo net sampling in different depth layers revealed thatPleurobrachia pileus, Bougainvillia ramosa andEucheilota maculata showed a preference for the surface layers around high tide, whereas most of the individuals were found in the deepest layer around low tide. The results suggest tidally-induced vertical migration of medusae in tidal channels. This may assist maintenance of the populations in the Wadden Sea area near Sylt.     

Circular pools in the seagrass beds of the Banc d’Arguin,Mauritania, and their possible origin

The tidal flats of the Banc d’Arguin, Mauritania, are covered by vast beds of Zostera noltii. At low tide these seagrass beds appear to be interspersed with partly vegetated, circular pools of 5–25 m diameter. Between February and May 2001 we described these pools and studied their possible origin. Several hypotheses regarding the origin have been developed. The first group of hypotheses assumes that the pools result from erosion activity. Since human disturbance of seagrass beds at the Banc d’Arguin is virtually non-existent, causes should be found in natural bed disturbances and/or tide or wave action. Therefore, small gaps, simulating holes dug by the crab Callinectes marginatus, were made to see if they would further erode by tidal currents or waves. The experiments showed no erosion. Neither we found support for other hypotheses assuming erosion to be the cause of circular pools. The alternative group of hypotheses stated that sedimentation on the flats would be responsible. We conclude that accretion of creek remnants is the most likely process behind the development of the pools; this conclusion is based on both mapping of the pattern of pools, the sediment profile in and around the pools and the distribution of seagrass biomass. Also the disturbance experiments showed bed accretion rather than bed erosion and support this hypothesis.     

Investigations on the ecophysiology of Geleia nigriceps Kahl (Ciliophora,Gymnostomata) inhabiting a sandy beach in Bermuda

Summary The dominant ciliate species Geleia nigriceps inhabiting a sheltered beach in Bermuda was characterized by its horizontal and vertical distribution on the beach in correlation with sediment composition and grain size, temperature, pH, oxygen, redox potential, and water coverage, its tolerance of high and low temperature under normoxic and oligoxic conditions, and its tolerance of extreme pH-values.From tolerance experiments it can be concluded that G. nigriceps is able to tolerate relatively high temperatures (up to 37.3°C) for only short time (e.g., when low tide occurs around noon on hot days).This species, inhabiting both the oxidized and the reduced milieu below the redox potential discontinuity layer of the beach, is only to a small degree dependent upon oxygen availability and can be regarded as a more or less habitual dweller of anoxic or oligoxic sediments. The functions relating LT₅₀ and exposure time were almost identical for normoxic and oligoxic conditions.Tolerance experiments revealed that pH values from 8.2 to 9.6 present no obstacle to G. nigriceps in its occupation of the upper millimeters of the sediment. In addition, it could be demonstrated that pO₂ has no effect on the tolerance of alkaline conditions.The vertical pattern of distribution of G. nigriceps correlated with water coverage during one period of outgoing tide suggests vertical migration. The upper sediment layer is more densely colonized than the lower ones at stations where water coverage exceeds 45%. A highly significant correlation exists between the population density of the upper centimeter and water coverage.Contribution No. 724 from the Bermuda Biological Station for Research     

In situ study of short-term variations of redox species chemistry in intertidal permeable sediments of the Arcachon lagoon

We investigated the composition of porewaters in intertidal sediments in response to the diurnal rise and fall of tides. For this reason, we deployed an in situ voltammetric system to measure vertical distribution and time-series at defined depths of O₂, Mn(II), Fe(II), and S(?II) in the porewater of permeable sediments from a protected beach in the Arcachon Bay. We also report microprofiles of O₂ and pH together with sediment properties (organic carbon, particulate reactive manganese and iron, porosity and permeability). Results shows that the oxygen dynamics in the upper sediment at low tide appeared to be mainly controlled by microphytobenthos activity, which may migrate downward just before immersion. The tidal forcing seemed to influence the oxygen dynamic in a minor way through flushing of the uppermost sediment porewater layer at the beginning and end of immersion. Vertical profiles and time-series measurements showed that the distributions of reduced species varied with tides. Although this work reveals that the upper sediment layer was subject to redox changes, the response of vertical distributions of redox species to tidal and night?Cday cycles did not have a cyclic pattern.     

Macrobenthic turnover in the subtidal Wadden Sea: The Norderaue revisited after 60 years

The benthic macrofauna of a tidal inlet in the northern Wadden Sea was sampled with grab and dredge in 1924–1926 (Hagmeier & Kändler, 1927), and again in 1985 and 1986. The comparison of surveys from consecutive years, as well as observations from an adjacent area, are employed to separate spurious from real long-term changes. Several epibenthic species of the 1920s became rare or absent in the 1980s. Oyster beds and reefs of the colonial polychaeteSabellaria spinulosa have disappeared completely. On the other hand, mussel beds have extended their range, and the abundance of mobile infauna has increased. The total number of species has remained approximately the same. Compared to surveys from consecutive years, the 60-year interval has doubled the species turnover rate, and has decreased the similarity in relative abundances by one third. The observed losses are best explained by the impact of dredging and trawling on the benthic fauna, while gains seem to indicate coastal eutrophication.     

Habitat preferences as patch selection of larval and emerging chironomids (Diptera) in a gravel brook

Four common species of larval and emerging chironomids were investigated during 1981 and between 1984/1985 in an alpine gravel brook. The emergence patterns ofN. dubius, O. rivulorum andH. ornaticollis were significantly affected by fluctuations of the water level, while rises in water temperature tended to trigger the peaks in emergence of the same species and ofO. frigidus. N. dubius exhibited its main emergence activity over areas of bed sediment which had a lower mean water velocity or were discontinuously flooded. On the other hand, the imagines ofO. frigidus, O. rivulorum andH. ornaticollis reached peak densities in zones of the main current channel in the Seebach. The larvae of these species inhabited horizontally and vertically different areas of bed sediment depending on the developmental stage. Early instars ofN. dubius andH. ornaticollis preferred zones near the banks and reached their maximum densities in sediment depths of 10 to 20 cm and 20 to 30 cm, respectively. In contrast, the larvae ofO. frigidus andO. rivulorum reached their highest densities in the first 10 cm of the sediments with significant preferences for the zones of stronger water flow.     

The benthic macrofauna along the estuarine gradient of the Schelde estuary

The intertidal benthic macrofauna of the Schelde estuary (The Netherlands and Belgium) was sampled in late autumn of 1990 at 50 stations along the whole salinity gradient (between Vlissingen and Dendermonde), including the freshwater tidal part. All stations were situated in sheltered areas with a relatively muddy sediment. Species richness, diversity and total biomass of the benthic macrofauna decreased along the salinity gradient from Vlissingen to Dendermonde, while total density showed no clear trend. Especially the oligohaline and freshwater tidal part of the Schelde estuary was characterized by a very impoverished benthic community, composed only of Oligochaeta. No other species (freshwater, marine or brackish) was observed in this part of the estuary. The marine part had a more diverse macrozoobenthos structure than that of the brackish part. Species found only in the marine zone areCerastoderma edule, Tharyx marioni, Eteone longa, Nephtys hombergii andCapitella capitata. In the brackish part of the estuary,Corophium volutator was a typical, dominant species. However, a lot of the dominant species were common in both the marine and brackish part of the Schelde estuary (e.g. Heteromastus filiformis, Pygospio elegans, Nereis diversicolor, Macoma balthica). The observed gradient in species composition and dominance is compared with some other European estuaries. The marine and brackish part of the Schelde estuary is quite similar to other european estuaries. The freshwater tidal part, however, was more impoverished.     

Planktonic foraminifera of the northern Indian Ocean: Distribution and preservation in surface sediments

In the northern Indian Ocean, planktonic foraminiferal tests accumulate in a wide variety of surface-water environments and depositional settings. This variability enables us to isolate the effects that surface-water ecology and differential dissolution have on the distribution of planktonic foraminifera from 251 geographically widespread surface sediment samples.Foraminiferal abundance varies from 0 to > 10⁴ whole foraminifera in the greater than 150 μm fraction per gram dry sediment. Values < 10 characterize the three deep basins of the equatorial Indian Ocean and the western Bay of Bengal. Foraminiferal tests are most abundant on carbonate covered Ninety-East and Carlsberg Ridges. Absolute abundance patterns are mainly controlled by non-ecological processes. Variations in dissolution resistant species (RSP) with water depth reveal that the foraminiferal lysocline (FL) varies regionally. The FL is deepest (3,800 m) in the equatorial region, rises abruptly to 3,300 m in the Arabian Sea, and varies from 2,600 m to near 2,000 m moving northward in the Bay of Bengal. Deep samples with anomalously low RSP (< 30%) suggest redeposition.Systematic geographic and depth-related variation is observed for the 17 most abundant foraminiferal species. Dissolution resistant species (G. menardii, G. tumida, G. dutertrei, P. obliquiloculata) generally exhibit a rapid and continuous increase in relative abundance at and below the FL. Susceptible species (G. ruber, G. bulloides, G. glutinata, for example) exhibit a rapid and continuous decrease in relative abundance at and below the FL. Moderately susceptible species (G. conglobatus, G. aequilateralis, G. conglomerata, for example) rapidly increase in abundance at the FL and systematically decrease with depth below the FL.Principal components analysis (PCA) of faunal data from minimally dissolved (< 30% RSP) samples reveals important ecologically related species intercorrelations. The major biogeographic gradient is the negative covarying relationship between aG. bulloides-G. glutinata species pair and a grouping ofG. sacculifer, G. conglobatus, G. aequilateralis, andG. ruber. PCA of all samples demonstrates how differential dissolution alters this and other species relationships. Species groupings that incorporateG. ruber, G. menardii, andG. dutertrei are particularly affected by dissolution.Comparison of average faunal data from minimally dissolved samples in the northern Indian Ocean with similar samples from other tropical regions suggests varying environmental factors produce distinct faunas within the tropical ocean. For example,G. bulloides, G. falconensis, andG. hexagona are significantly more abundant in northern Indian Ocean surface sediments while such species asG. ruber, G. sacculifer, G. dutertrei, andP. obliquiloculata dominate in other tropical regions.     

Impact of hydrodynamic changes on the zonation of helophytes

Water movement has a strong impact on the development and persistence of helophyte stands along large water bodies. The decline ofScirpus andPhragmites stands in the Rhine-Meuse estuary (The Netherlands), due to changed hydrodynamic conditions (reduced tidal action, increased shoreline erosion), illustrates the effects of large-scale hydrological regulation. Hydrodynamic factors partly explain the distribution and composition of helophyte vegetation bordering the open water. A vertical zonation emerges during the stages of dispersal, germination, seedling establishment and adult growth of the plants. Differences between species may result in the occupation of different water-depth zones. Additionally, gradients of wave exposure result in a horizontal zonation along water bodies. Restrictions to helophyte development due to the hydrological regime and wave attack should be taken into account in wetland restoration studies. Water-level management, instead of a strict control of water levels, should be one of the key elements of water management in the future.