Werner Armonies

Passive settlement of Macoma balthica spat on tidal flats of the Wadden Sea and subsequent migration of juveniles

Abstract Spatfall of the tellinid bivalve Macoma balthica may occur in fine-grained sediments at different tidal elevations in the Wadden Sea. but it is not clear which mechanism (active habitat selection or passive sinking of larvae, or both) can explain the observed distribution pattern. Spatfall and the subsequent development of juvenile abundances and size distribution were studied along a transect in the northern Wadden Sea near the Island of Sylt. Initial spatfall was highest in the lower intertidal and correlated significantly with hydrographic parameters. The enhanced larval settlement observed in dense aggregates of the tube-building polychaete Lanice conchilega confirmed the prominent role of hydrographic factors in initial spatfall. Subsequently, abundance strongly decreased in the lower intertidal but increased in the upper intertidal due to immigration of newly settled spat from lower sites. By the end of October, abundance was higher in the upper intertidal. Growth of the juveniles was higher in the upper intertidal. It is suggested that (1) initial spatfall is mainly ruled by hydrographic factors without active sediment selection; (2) young Macoma remain highly mobile during the first weeks of their benthic life and by byssus drift may achieve a net transport into areas with weak initial spatfall; (3) at the end of their first winter a second period of high floating activity occurs, bringing them back towards the lower intertidal and shallow subtidal. Thus, by successive postlarval migrations Macoma may several times change the intertidal site occupied during their first year of life. The comparatively low interannual variability of recruitment success noted in this species in the Wadden Sea may be a result of these migrations.

ON THE POPULATION DEVELOPMENT OF THE INTRODUCED RAZOR CLAM ENSIS AMERICANUS NEAR THE ISLAND OF SYLT (NORTH SEA)

The American razor clamEnsis americanus (=E. directus) was introduced into the eastern North Sea in the late 1970s. By larval and poslarval drifting the species rapidly extended its distribution, now ranging from the English Channel to the Kattegat. Near the island of Sylt in the eastern North Sea it has been recorded since 1979. Recruitment was rather irregular, with about six strong year-classes within two decades. Growth seems comparable with populations in its native range (Atlantic North America). Although present in the lower intertidal zone, maximum densities occurred in shallow subtidal sand with a biomass similar to that of dense beds of native cockles and mussels in the adjacent intertidal zone.Ensis americanus established in otherwise sparsely faunated sand (channels exposed to strong currents) as well as in dense infaunal assemblages (lower intertidal and subtidally). There were no significant interactions with resident species. In dense beds of razor clams, however, fine sediment particles accumulated which may have altered abundances of polychaetes. In spite of high annual variability,E. americanus has become a prominent component of the macrobenthos in shallow subtidal sands of the North Sea.

Meiofaunal emergence from intertidal sediment measured in the field: significant contribution to nocturnal planktonic biomass in shallow waters

Field studies on the occurrence of meiobenthos in the water column above intertidal sandflats have been performed near the Island of Sylt in the northern Wadden Sea. Swimming meiobenthos was strongly dominated by harpacticoid copepods. Many of them have a semiplanktonic life-style. They rest in superficial sediment layers at low tide and swin in the water column at high tide. Swimming activity correlated negatively with light. The abundance in the water column was one order of magnitude higher during the night. Strong currents caused by storm tides significantly decreased meiobenthic abundance in the water column. Light and flow being constant, no significant changes of meiobenthic abundance per unit area occurred over a tidal cycle. Since holoplankton and meroplankton abundances correlated positively with the height of the water column, semiplanktonic meiobenthos may dominate the mesozooplankton in shallow waters. On an average, emergence of meiobenthos increased the mesozooplanktonic biomass by about 2% during diurnal high tides over the entire tidal cycle, and by about 50% during nocturnal high tides. Because of seasonal cycles of the dominant harpacticoids, this high contribution to planktonic biomass may be a summer phenomenon.

Empty habitat in coastal sediments for populations of macrozoobenthos

Abstract. Species with wide dispersal are expected to have little empty habitat. This was tested by analysing habitat use in the Wadden Sea near the island of Sylt (North Sea). Sampling covered 222 intertidal sites spread across the Königshafen tidal flats (4.5xa0km2, mapping approach conducted once) and 270 subtidal sites along 12xa0km of a tidal channel system (stratified random sampling in spring and autumn). For any single species, suitable habitat was extrapolated from the ranges of water depth and sediment composition present at the ten sites with the highest frequency or abundance of the species. On average, macrobenthic species actually used less than half of the suitable sites within the scale of local populations; this was far less than expected from a lognormal distribution. In the subtidal, abundance of most species changed between the two sampling seasons. A 50% increase in overall abundance was accompanied by a decrease in empty habitat of only about 25%. Thus, a doubling of abundances would not fill the empty space but just of half of it. The polychaete Scoloplos armiger was an exception in occupying almost all of its suitable habitat in the intertidal and subtidal sediments. A distinct patch of high species richness occurred where flood waters persistently form a large gyre which may enhance larval settlement. We suggest that limitations to larval settlement and/or juvenile survival primarily cause the extent of the observed habitat emptiness.

Synoptic patterns of meiofaunal and macrofaunal abundances and specific composition in littoral sediments

During recent years, many investigations on small zoobenthos have been performed at the island of Sylt. As these studies were carried out sporadically over many years and as different extraction methods were used, comparisons of the results have been hampered. Therefore, in August/September 1986, 24 sites were sampled and evaluated using one quantitative method throughout. Sites range from mud to exposed sand and from the sublittoral to the supralittoral. Macrofauna and the taxa Plathelminthes, Polychaeta, and Oligochaeta are determined to species level. Macrofaunal (>0.5 mm) abundance is highest in mud and continuously decreases with increasing exposure to wave action. Meiofaunal (<0.5 mm) abundance is less variable. Nematoda dominate in mud and muddy sand, Copepoda in sheltered and exposed sand, other taxa only intermittently. Related to surface area, no correlation between macro-and meiofaunal abundance is apparent. Plathelminthes and Copepoda reach highest abundance per surface area in sand but their per volume density is higher in mud and muddy sand. Related to sediment volume instead of surface area, the meiofaunal abundance pattern is very similar to the macrofaunal pattern. The faunal composition changes gradually along the tidal gradient without general faunal boundaries. On an averange, the faunal similarity of neighbouring sites is highest in Oligochaeta and lowest in Plathelminthes. Presumably, Oligochaeta tolerate wider ranges of environmental factors. This may explain the low number of oligochaete species. On the other hand, Plathelminthes seem to adapt to relatively narrow ranges of factors and their species richness is highest. Because of macrofaunameiofauna interaction it is suggested that the meiofaunal assemblage will be least stable in mud and muddy sand, and most stable in exposed sand.

On the spatial scale needed for benthos community monitoring in the coastal North Sea

Abstract In community monitoring an attempt is made to identify long-term trends by regular sampling of selected sites. Since the benthos is reputed to be fairly sedentary, the spatial resolution is often reduced to single sites. However, members of many benthic invertebrate species have been found drifting across sedimentary seabeds in shallow waters. Transportation by currents may result in changes of their spatial pattern in the sediment, thereby changing local community composition. The quantitative importance of drifting was tested by repeated sampling of a 2-km 2 shallow (10xa0m) offshore area west of the island of Sylt (North Sea). Within the fortnight period between two samplings the benthic community composition had changed dramatically. Despite fairly calm weather, translocation of organisms by currents exceeded 1xa0km. In about half of the species, the spatial changes in abundance within these two weeks roughly equalled the average variation between consecutive years. This example suggests that community monitoring needs a wide spatial scale to discriminate long-term temporal changes from short-term variability. Extending the sampling area from 2 to 180xa0km 2 strongly reduced the variability of abundance estimates. However, only in a few species was the spatial distribution over the sampling sites found on one sampling date a suitable estimator for the spatial pattern found one or two months later, at the same sites. Instead the spatial patterns of the fauna changed strongly during a single month with a spatial scale of re-distribution exceeding several km in some species. At the same time the granulometric sediment composition changed, indicating changes of habitat quality. Hence, sampling of a large area, with random selection of the sampling sites on each sampling date, is suggested to yield the most reliable estimates of population development in the coastal North Sea. However, in view of the expected spatial scale of re-distribution during storm tides and the spatial variability of recruitment, even a 180-km 2 sampling area may be too small.

Effects of the severe winter 1995/96 on the benthic macrofauna of the Wadden Sea and the coastal North Sea near the island of Sylt

Abstract. The development of benthic macrofauna in the Wadden Sea and in the coastal North Sea after the severe winter of 1995/96 is compared with the preceding years with mild to moderate winters. In the intertidal of the Wadden Sea, ice-drift and low temperature caused the expected changes in species composition by increasing winter mortality in sensitive species, and by exceptionally high recruitment of some species during the succeeding summer. In the shallow subtidal (10–20xa0m depth), similar winter effects were observed. However, recovery of many subtidal populations was still incomplete until the summer of 1997. It is suggested that this was due to hydrographic conditions that carried many larvae or drifting juveniles into more distant offshore areas. This may have limited larval supply and may have delayed recovery at the onshore sites. Since in the eastern North Sea severe winters are accompanied by frequent easterly winds, it is not clear whether decreasing winter abundances in some species were due to increased mortality, or to a seaward dislocation of organisms.

Short-term changes of meiofaunal abundance in intertidal sediments

Some species of various meiofaunal taxa may actively emerge from the sediment and swim in the water column, preferably at night. In the water column they get dispersed by tital currents. The hypothesis that this drift may cause significant short-term changes in the abundance of these species was tested. Such changes were verified for harpacticoid copepods and plathelminths. Depending on the hydrographic conditions, abundance changes may be unpredictable or show regularly alternating patterns. In the light of these results, small-scale spatial estimates of abundance become generally very difficult to make in the taxa containing emergent species. It is suggested that reliable averages can only be obtained by repeated or large-scale spatial sampling.

Hydrodynamic factors affecting behaviour of intertidal meiobenthos

Abstract Intertidal meiobenthos may actively emerge from the sediment and enter the water column. Changing hydrostatic pressure and the dimensions of the water column in laboratory experiments did not affect the meiobenthic emergence pattern. Varying current speed in laboratory flumes significantly changed the abundance of Copepoda, Plathelminthes, and Nematoda in the water column while Ostracoda were not affected. Nematode emergence correlates significantly with current speed and is predominantly a result of passive erosion, possibly enhanced by a vertical migration of the specimens to the sediment surface. A current speed > 1 and ≥ 10 cm · s-1 significantly reduced the numbers ofCopepoda and Plathclminthes actively entering the water column. Observation on the swimming behaviour of members of the latter two groups shows that the horizontal component of their active movement is negligible. Presumably, most of these specimens actively enter the water column in order to disperse by currents.

The diet of Enoplus brevis (Nematoda) in a supralittoral salt marsh of the North Sea

The gut content of nearly 2000 specimens ofEnoplus brevis was quantitatively analysed.E. brevis is an opportunistic omnivore. Main food categories found were cyanobacteria, diatoms, oligochaetes, nematodes, and rotifers. With a few exceptions, the diet of females and males was identical. Predominantly, juveniles fed on cyanobacteria and adults on animal prey. Experiments revealed that living oligochaetes are attacked and preyed upon. The quantities of the food categories eaten byEnoplus varied according to their ambient abundance, both temporarily and spatially on a small scale. At least the nematode prey was strongly selected: some species were significantly preferred and others avoided.