Seasonal occurrence of the ctenophore Pleurobrachia pileus in the western Dutch Wadden Sea

Due to the short residence time of the water in the western Dutch Wadden Sea the ctenophore Pleurobrachia pileus has to be considered as an allochthonous species, which enters the estuary passively from the North Sea. The seasonal occurrence of this species in the Dutch Wadden Sea is a reflection of its abundance in the coatal zone of the North Sea and is characterized by a pronounced spring peak of up to 17000 individuals per 1000 m³ and a lower autumn peak with numbers up to 400 individuals per 1000 m³. Similar to other areas, predation by Beroë gracilis, another ctenophore species, may be a factor in controlling the P. pileus population in this region, although B. gracilis abundance is low. P. pileus is only important as a predator for a short time, i.e. the month of May, when it reaches maximum biomass values of up to 12 00 mgC per 1000 m³. Both the estimated potential consumption rates in comparison with the available food as well as the observed suboptimal growth rate of the population suggest that in the coastal zone of the North Sea area food conditions may not be optimal for P. pileus.     

An evaluation of the ABC-method (abundance/biomass comparison) as applied to macrozoobenthic communities living on tidal flats in the Dutch Wadden Sea

The ABC-method proposed by Warwick (1986) for detecting pollution effects on marine zoobenthic communities, was tested on distribution patterns of numbers and biomass among species in macrozoobenthos samples taken annually for 13 yr in a uniform way at 15 tidal-flat stations in the western part of the Dutch Wadden Sea. Along the margins of the tidal-flat area studied, where exposition to either drainage or water movements is extreme, the k-dominance plots for numbers were generally situated above those for biomass. In these areas values for biomass and species richness were relatively low and a few small-sized species such asCorophium volutator andHydrobia ulvae were very numerous. In the central part of the area, where environmental conditions are less severe, values for biomass and species richness were higher. Biomass was dominated by large adults ofMya arenaria, Mytilus edulis and/orArenicola marina. However, in only a restricted part of this area were plots for numbers consistently below biomass plots. In most of the central area high numbers ofH. ulvae occasionally occurred, causing plots for numbers to be situated above those for biomass during such periods. There appears to be no reason to interprete a high abundance ofH. ulvae as a sign of pollution or any other kind of disturbance or stress. It is concluded that the ABC method cannot be applied to tidal flat communities without reference to long-term and spatial series of control samples. In areas where such smallsized and sometimes numerous species asH. ulvae occur in strongly fluctuating numbers, the method appears useless for assessing the pollution status of a benthic community.     

Effect of periphyton grazing by Hydrobia ulvae on the growth of Zostera noltii on a tidal flat in the Dutch Wadden Sea

The decrease of the intertidal seagrass Zostera noltii in the Dutch Wadden Sea may have been the result of enhanced periphyton load due to eutrophication. Decrease of this seagrass species coincided with an increase in the mudsnail Hydrobia ulvae. Feeding of this mudsnail on periphyton may have partly counteracted an increase of periphyton biomass. Exclosure experiments on seagrass stands in the Dutch Wadden Sea in 1987 showed that density of periphyton on leaves of Z. noltii decreased significantly with increasing density of grazers. An increased density of mudsnails significantly enhanced the density and biomass of seagrass, in particular of the below ground parts. Since this seagrass species survives winter in temperate climate zones mainly by means of rhizomes, grazing may also influence the seagrass dynamics in the long term. Results of the experiment were in agreement with field observations on coinciding low densities of mudsnails and high densities of fouling of seagrass stands on the tidal flats of western Wadden Sea in the late 1970s.     

Hermaphroditism in brown shrimp: lessons from field data and modelling

This study was undertaken to further clarify whether the brown shrimp, Crangon crangon (Linnaeus 1758), is a gonochorist, a facultative or an obligate hermaphrodite. Juvenile shrimps were sampled from intertidal habitats along the German Wadden Sea coast with a push net and from a power plant water inlet to quantify the share of primary females. Length-based sex ratios were determined for about 27,000 individuals using external characteristics. Observed sex ratios were mainly female-biased, and also large males occurred regularly in the catch. This indicates that sex at hatch is not male as would be characteristic for an obligate protandric hermaphrodite and that not all male shrimps change sex. A cohort-based computer simulation, including sex-specific growth rates, mortality and seasonally varying recruitment, generated sex ratios comparable to the field. The observed decline in the proportion of males with increasing size can be explained solely by faster growth of females without involving hermaphroditism. Based on temperature-dependent growth and moult rates as well as length-specific numbers of eggs per female, the potential egg production of primary and secondary females was modelled, yielding contributions of secondary females of <1%. Sex change in C. crangon has previously been observed and may be interpreted as an evolutionary relict of this species having evolved in a habitat characterized by lower population densities, lower predation levels and increased longevity compared to today’s living conditions in North Sea coastal waters.     

The contribution of seagrass beds (<Emphasis Type="Italic">Zostera noltii</Emphasis>) to the function of tidal flats as a juvenile habitat for dominant,mobile epibenthos in the Wadden Sea

Since the substantial loss of subtidal eelgrass (Zostera marina L.) in the 1930s, seagrass beds in the Wadden Sea are limited to the intertidal zone and dominated by Z. noltii Hornem. This study deals with the effect of vegetated tidal flats on quantities of mobile epifauna and proves empirically the function of seagrass canopies as a refuge for marine animals remaining in the intertidal zone at ebb tide. Drop-trap samples were taken in the Sylt-Rømø Bight, a shallow tidal basin in the northern Wadden Sea, on vegetated and unvegetated tidal flats during July and August 2002, and during the entire growth period of Z. noltii from May to September in 2003. The species composition in Z. noltii and bare sand flats showed minor differences since only two isopod species (Idotea baltica and I. chelipes) occurred on Z. noltii flats exclusively. Juvenile shore crabs (Carcinus maenas L.), brown shrimps (Crangon crangon L.) and common gobies (Pomatoschistus microps Krøyer) were also found abundantly on bare sand flats. However, the results showed significantly higher abundances and production of these dominant species on vegetated tidal flats. Additionally, the analyses of faunal size classes indicated higher percentages of small individuals in the seagrass bed during the entire sampling period. Despite drastic diurnal fluctuations of dissolved oxygen at low tide, faunal density in the residual water layer remaining in seagrass canopies at ebb tide was found to be consistently higher than that found in artificially created tide-pool units. Although species composition of mobile epifauna did not basically differ between vegetated and unvegetated tidal flats, Z. noltii beds are considered to contribute quantitatively to the function of tidal flats, as an extended juvenile habitat for some of the most important species of the Wadden Sea food web.Communicated by O. Kinne, Oldendorf/Luhe     

Habitat related growth and reproductive investment in estuarine waters,illustrated for the tellinid bivalve <Emphasis Type="Italic">Macoma balthica</Emphasis> (L.) in the western Dutch Wadden Sea

In estuarine areas, bivalve species can be found in a variety of environments, where they experience large differences in environmental conditions. In the present paper, the importance of different habitats (intertidal, subtidal and adjacent coastal waters) for the persistence of the population was evaluated for the bivalve Macoma balthica (L.) in the western Dutch Wadden Sea estuary. Intra-specific variation in growth and reproductive output were followed during the year and related to local abiotic conditions. Significant differences in growth and reproductive investment were found between locations. Young individuals were mostly found in the intertidal area, where growth in terms of somatic mass was good. These areas were not favourable for adult individuals, since growth in shell length was low and many individuals did not reproduce. In the subtidal, where the highest densities were found, somatic and gonadal mass indices were low. Coastal areas had the lowest densities and showed high growth in terms of shell length and body mass. The habitat with the highest reproductive effort per individual was not the most important habitat in terms of reproductive output due to differences in density and in size of the habitat type. For M. balthica, the subtidal habitat contributed the most to the reproductive output of the western Dutch Wadden Sea population although the highest reproductive output per individual was in the coastal area.     

A meiofaunal “thiobios” limited to the anaerobic sulfide system of marine sand does not exist

The spatial relationship of interstitial metazoans to sources of oxygen has been studied on a tidal flat in the Wadden Sea near Sylt (eastern part of the North Sea). Consistently, meiofauna remains in close proximity to oxygenated layers or pocket areas within the sediment. This is exemplified by the pattern of meiofaunal abundance and species composition alongside oxic burrows of the lugworm Arenicola marina L. A specific meiofauna confined to oxygen-deficient horizons of the sediment does not exist.     

Siphon size and burying depth in deposit- and suspension-feeding benthic bivalves

This paper analyses the significance of siphon investment in the life strategy of benthic bivalves. It describes the relationships between siphon weight, burying depth and shell size in Mya arenaria, Cerastoderma edule. Scrobicularia plana and Macoma balthica. All data were collected on an intertidal flat in the Dutch Wadden Sea during seven successive winter and summer periods. The four species have in common that (1) the increase of depth in relation to size can be described with an S-curve; (2) there is a linear relationship between log siphon weight and log shell size; (3) siphon investment is maximal for the size classes with the greatest increase in their depth; (4) siphon weight, in proportion to total body weight, decreases gradually for the larger size classes whose depth does not increase; (5) burying depth increases with siphon weight if individuals within a same size class are compared, but burying depth levels off above a certain siphon weight. Macoma balthica and Scrobicularia plana live twice as deep in winter as in summer, although siphon weight for both seasons is about the same. In summer both species use a part of the siphon to graze the surface around the burrow, whereas deposit feeding does not occur in winter. This might explain the seasonal variation in burying depth. On the other hand Cerastoderma edule and Mya arenaria, which are both suspension feeders, show hardly any increase of depth in winter as compared to summer. For benthic bivalves the risk of being taken by a predator decreases with depth. The burying depth levels off where individuals reach the depth refuge (though in winter Scrobicularia plana live at greater depth). The conclusion is that siphon size is one of the main factors determining the burying depth of benthic bivalves and thus plays a critical role in their survival.     

Local changes in the composition and community structure of suprabenthic peracarid crustaceans on the bathyal Mediterranean: influence of environmental factors

Deep-water assemblages of suprabenthic peracarids were analyzed in the SW Balearic Islands (Algerian Basin, southwestern Mediterranean) between 249 and 1,622 m depth; the patterns of species composition, possible zonation, and trophic structure found in this area were compared with those exhibited by peracarids in the mainland side of the Catalan Sea slope (northwestern Mediterranean). One hundred and four peracarid species (plus one leptostracan) were identified on the Balearic Islands slope, amphipods being the most diversified taxon (45 species). On the Balearic slope, two distinct depth assemblages were distinguished: one at the upper slope (US), between 249 and 402 m depth and the second at the deep slope, between 543 and 1,620 m depth. A remarkable species substitution occurred at depths between 402 and 638 m. In the Catalan Sea, in addition to the US assemblage occupying depths between 208 and 408 m, a second boundary of faunal change was found around 1,250 m. Suprabenthos biomass increased from 242 to approximately 500 m. Suprabenthos attained the highest biomass values (100 g wet weight/10,000 m²) at intermediate depths between 504 and 1,211 m, as also occurred with the associated zooplankton collected with suprabenthos (peak biomass between 502 m and 898 m). Suprabenthos biomass did not show any significant correlation with any environmental water-column variable. In contrast, zooplankton (especially small fish and decapod crustaceans) showed a significant positive correlation with fluorometry and turbidity at different levels of the water column. The feeding guilds of species showed important differences between the two areas only on the US, with a higher abundance of deposit feeders in the Catalan Sea (20.4%) than in the Balearic Islands (4.2%). The low contribution of deposit feeders in the SW Balearic Islands may ultimately be a consequence of the lack of river discharges in this area.Communicated by S.A. Poulet, Roscoff     

Are Wadden Sea tidal systems with a higher tidal range more resilient against sea level rise?

Accelerated sea level rise may have serious implications for the Wadden Sea ecosystem in its present state. If sediment accumulation rates on the extensive intertidal flats stay behind sea level rise, the flats will eventually submerge. Drowning of the flats has negative consequences for nature conservation and for coastal risk management. Based upon an evaluation of steady state relations for Wadden Sea tidal basins, Hofstede (Zeitschrift für Geomorphologie 59(3): 377-391, 2015) postulated that the capacity of these basins to balance sea level rise by accumulation on intertidal flats seems positively related to mean tidal range. In the present study, morphodynamical simulations with a numerical model were performed for two tidal basins in the German Wadden Sea to verify the empirically established hypothesis. The following conclusions are established. Larger mean tidal range improves the capacity of Wadden Sea tidal basins to balance sea level rise. Wadden Sea intertidal flats are effective sediment sinks and seem quite resilient against (higher rates of) sea level rise. Finally, subtidal gullies may constitute a significant sediment source for accumulation on intertidal flats in response to sea level rise. With respect to the limited comparability of the two investigated tidal systems, morphodynamical modelling of all Wadden Sea tidal systems should be conducted.     

Continuous plankton records: Geographical variations in numerical abundance,biomass and production of euphausiids in the North Atlantic Ocean and the North Sea

Geographical variations in the numbers, biomass and production of euphausiids and the contribution of common species to the total are described from samples taken during 1966 and 1967 in the North Atlantic Ocean and the North Sea by the Continuous Plankton Recorder at 10 m depth. Euphausiids were most abundant in the central and western North Atlantic Ocean and the Norwegian Sea. Thysanoessa longicaudata (Krøyer) was numerically dominant. Biomass was greatest in the Norwegian Sea and the north-eastern North Sea where Meganyctiphanes norvegica (M. Sars) accounted for 81 and 59%, respectively, of the total biomass. Production was highest off Nova Scotia and in Iberian coastal waters; the dominant species were T. raschi (M. Sars) in the former area and Nyctiphanes couchi (Bell) in the latter. The mean P:B ratios were correlated with temperature.     

Spatial and temporal patterns of habitat use by Eurasian oystercatchers (<Emphasis Type="Italic">Haematopus ostralegus</Emphasis>) in the eastern Wadden Sea revealed using GPS data loggers

Detailed knowledge on species–habitat relationships is of crucial importance for the understanding of processes in marine ecosystems. Being top-predators, birds are important bio-indicators for marine systems. The aim of this study was to elucidate precise information on foraging habitat use and foraging times of oystercatchers (Haematopus ostralegus) on wide tidal flats using global positioning system (GPS) data loggers. The study was conducted to collect hints for the negative population trends in oystercatchers in the Wadden Sea. It is the first time that GPS technique has been used in a shorebird species. Although oystercatchers are known to exhibit foraging site fidelity, a number of individuals visited multiple sites. Foraging trips at night were longer, and the targeted sites were further away than those used during the day. These patterns were likely to be caused by higher risks of clutch predation by avian predators during the day that led adults to reduce their absence to defend their clutches. Our methodological approach enabled the subtle spatio-temporal patterns of habitat use to be determined on a very fine spatio-temporal scale. We suggest further potential studies using GPS data loggers that may help to reveal the reasons for the current declines in oystercatcher populations in the German Wadden Sea.     

Food habits and intestinal parasites of deep demersal fishes from the upper continental slope east of Newfoundland,northwest Atlantic Ocean

Stomach contents and intestinal parasite faunas of 471 individuals of demersal fishes in 14 species were examined from the Carson Canyon region (Lat. 45°30N; Long. 48°40W) of the upper continental slope of the Grand Banks off Newfoundland, Canada. Individual species tended to feed either on benthic or on pelagic/benthopelagic organisms, but pelagic prey assumed the greatest importance overall. Data from stomach contents were supported by the parasite information. Prevalence of parasites was higher in benthic feeders (53.1%) than in pelagic feeders (28.9%), and relative abundance by major group was: Digenetic Trematoda 5.8% benthic vs 27.8% pelagic, Nematoda 53.1% vs 72.2%, and Acanthocephala 40.9% vs 0%. Of the dominant fishes, there were more species of benthic feeders (5) than pelagic feeders (3), but pelagic feeders were numerically more abundant (pelagic 70.9%, benthic 20.5%). Benthic feeders were on average larger (=270.6g) than pelagic ones (=130.6g), but pelagic feeders represented a larger proportion of the biomass (pelagic 43.3%, benthic 25.9%). The results of this study combined with those from other areas suggest that feeding from the pelagial by demersal fishes at upper continental slope depths is probably the general rule.     

Utilisation of mangroves and seagrasses by fishes in the Negombo Estuary,Sri Lanka

In two years of sampling, 10 287 fishes belonging to 62 species (39 families) were collected from mangroves and seagrasses of the Negombo Estuary. Of the species collected, 70% were marine, 27% estuarine and 3% freshwater. Aplocheilus melastigma, Ambassis dayi, Etroplus suratensis, E. maculatus, Bathygobius fuscus and Arothron immaculatus constituted 90% of the catch. Sixteen species were caught only from mangroves, 6 only from seagrasses, and 40 from both. The number of species, individuals, biomass and diversity were higher in seagrasses than in mangroves. Gobies and some siganids moved to the mangrove habitat as they grew, but cichlids, triacanthids, some eleotrids and tetraodontids preferred seagrasses. The number of species and individuals caught increased at the beginning of the north-east monsoon when the rainfall was low. The number of species and individual numbers were positively correlated with salinity and dissolved oxygen and negatively correlated with water temperature. Of the 33 species examined, seagrasses formed >25% of the gut content in juvenile Lates calcarifer, Apogon thermalis, E. suratensis, E. maculatus, Siganus javus, Butis butis and Bathygobius fuscus. Mangrove litter was the main gut content of the family Syngnathidae. E. suratensis, E. maculatus and S. javus from seagrasses contained more filamentous algae than seagrasses in their stomachs. S. javus collected from mangroves also contained more seagrasses in their stomachs than mangrove litter, indicating that the association of juvenile fishes with mangroves or seagrasses may not be obligate in estuaries subject to low tidal fluctuations.     

Sea anemones and brittle stars: unexpected predatory interactions during induced in situ oxygen crises

During oxygen crises, benthic faunas exhibit a series of behavioural patterns that reflect the duration and severity of the event. During artificially induced oxygen deficiencies at 24 m depth in the Northern Adriatic Sea, we photographically documented predation by the sea anemones Cereus pedunculatus (Pennant, 1777) and Calliactis parasitica (Couch, 1842) on the brittle star Ophiothrix quinquemaculata (DelleChiaje, 1828). Five predatory events were recorded with four anemones during nine deployments totalling 817 h of observation. Under near-anoxic conditions, individuals of both actinians made contact with, pulled in and consumed the brittle stars. The duration of each predatory event was 1.5–7.5 h. In three of the five events, brittle star remains were regurgitated after an additional 2.0–12.5 h of digestion by the anemones. Our time-lapse sequences demonstrate that oxygen deficiency, beyond eliciting a series of specific behaviours in members of each species, also promotes previously unobserved interspecific interactions. Our results show that sea anemones are not only highly resistant to anoxia, but may also benefit by taking advantage of prey that are more vulnerable to anoxic conditions.     

Observations on the biology of Donax trunculus and D. vittatus at Ile d'Oléron (French Atlantic coast)

Observations have been made on the biology of two species of the bivalve genus Donax, on two beaches, Vertbois and St. Trojan, on the Ile d'Oléron, French Atlantic coast. The two species displayed different vertical distribution: D. trunculus Linné in the intertidal; and D. vittatus (da Costa) between low-water spring tide (LWST) and 5 to 6m depth, with only the fringe of the population extending to the shore. D. trunculus also showed differential distribution by size or age, the youngest individuals being highest on the shore, and the oldest near LWST. The two species exhibited a similar pattern of seasonal recruitment and growth, but in both there were differences in growth characteristics between the two beaches, individuals at St. Trojan growing to a larger size than those at Vertbois. The mean pattern of mortality in the two areas was similar, and productivity per 100 recruits surviving to the first winter following settlement was greater at St. Trojan than at Vertbois. For D. trunculus the mean population density at Vertbois was higher and the production per unit area on the two beaches was similar. Production to biomass (B) and elimination to biomass (B) ratios were similar for the two beaches for both species, and while production was greater for D. trunculus at both beaches, the B and B ratios were higher for D. vittatus. Comparisons with other published data reveal that the rate of growth of D. trunculus is similar throughout its range from Brittany to the Mediterranean Sea, while that of D. vittatus shows more variation. In both species, there is a trend of increasing P/B and E/B ratios from north to south throughout the geographical range.     

Population dynamics and production of the planktonic copepods in a eutrophic inlet of the Inland Sea of Japan. I. Centropages abdominalis

Population dynamics and production of the calanoid copepod Centropages abdominalis were studied from November 1986 to November 1987 in Fukuyama Harbor, in the central part of the Inland Sea of Japan. This species was present in the plankton during a cold-water period from November to June (temperature range: 8.9 to 21.1 °C), with a peak abundance (23 600 ind m^–3) in February. During this period, six generations could be detected, and each generation time agreed well with that predicted from food-satiated laboratory experiments, indicating that the natural population was not food-limited. The population suffered extremely high mortality during the period from egg to naupliar stage (N) II: only 0.02 to 4% of the eggs survived to NII. However, the mortality in stages older than NII was almost negligible. The growth rate of C. abdominalis increased exponentially with increasing temperature. Its biomass and production rate showed marked seasonal variations largely in parallel with numerical abundance. The estimated production between 7 November 1986 and 29 May 1987 was 355 mg C m^–3 or 2.66 g C m^–2, 95% of which occurred during February and March. The daily production rate to biomass ratio increased exponentially with temperature from 0.18 at 8.9°C to 0.37 at 19°C.     

Macrozoobenthos diversity in an oxygen minimum zone off northern Namibia

A benthological survey in the Benguela upwelling area off northern Namibia (located at 17.3°S and water depth ranging between 26 and 117 m) showed the concentration of dissolved oxygen and the accumulation of organic-rich sediments to control macrozoobenthic community patterns. In contrast to highly biodiverse nearshore areas with well-structured shell deposits of the brachiopod Discinisca tenuis (Sowerby 1847), the benthic community in the oxygen minimum zone (OMZ) decreased strongly in species numbers. Nevertheless, a well-established community ranging from 13 to 31 species persisted. Species densities (300–3,350 ind m⁻²) and biomass (4–109 g afdw/m²) were surprisingly high for areas with near bottom oxygen concentrations from 0.06 to 0.88 ml l⁻¹. In contrast to OMZ’s of other upwelling areas, where the benthic macrofauna is generally dominated by small-bodied polychaetes, off Namibia larger key organisms like the bivalve Nuculana bicuspidata (Gould 1845) and the snail Nassarius vinctus (Marrett 1877) accounted for a large proportion of the macrozoobenthos >1 mm. This is supposed to have a distinct effect on the functional properties of the sediments.

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Recurrent and changing seasonal patterns in phytoplankton of the westernmost inlet of the Dutch Wadden Sea from 1969 to 1985

Data for phytoplankton composition and abundance in the Marsdiep are presented for the period from 1969 to 1985 inclusive. Only a few species dominated the phytoplankton. A recurrent pattern was observed in the seasonal succession: in winter, total cell numbers were invariably low, but freshwater algae, sluiced into the Wadden Sea from IJssel Lake, showed highest densities in winter. A diatom spring peak was observed around mid-April, followed by a Phaeocystis pouchetii peak about three weeks later. Later in summer usually two more diatom peaks followed by non-diatom peaks were present. The exact timing of the spring peak varied from year to year, with the extremes being late March and early May. A relatively late spring peak usually coincided with a relatively high turbidity in the preceding winter. An increase in total cell numbers was found over the 17-year observation period. Diatoms decreased from 1969 to 1974 but have increased since then, reaching values above those of 1969 during recent years. Flagellates showed a consistent increase over the entire observation period.     

Influence of anoxia on adaptation of euryhaline polychaetes to hyposmotic conditions

The metabolic reactions ofArenicola marina andNereis diversicolor (both euryoxic and euryhaline polychaetes from the intertidal) to hyposmotic shock under anaerobic conditions were investigated in 1987–1988 using specimens from the East Frisian Wadden Sea. Although reductions in salinity were within the limits of tolerance under normoxic conditions, during anoxia they resulted in increased stress and a significant reduction in survival time. Both species were unable to sufficiently intensify anaerobic energy production to match the increased requirement for energy during adaptation to hyposmotic conditions. Moreover, as an adaptation to anoxic conditions, worms reduced their metabolic activity after the first 6 h. Thus, specimens remained more swollen than at the same reduction in salinity under normoxic conditions. InA. marina only the extracellular volume was reduced all be it to a limited extent.N. diversicolor probably reacted in the same or similar way. In both species, cells remained swollen. The concentrations of cellular free amino acids were not reduced; that means the main mechanism of cellular volume regulation was not activated under these conditions.