Short-term and long-term facilitation of goose grazing by livestock in the Dutch Wadden Sea area

We studied the impact of livestock grazing on the distribution ofBranta bernicla bernicla (Dark-bellied Brent goose) in the Dutch Wadden Sea during spring. It was hypothesized that livestock facilitate short-term (within-season) grazing for geese as well as long-term (over years). Therefore we measured grazing pressure by geese in salt marsh and polder areas that were either grazed (spring-grazed) or ungrazed during spring (summer-grazed). Additionally, we carried out a preference experiment with captive geese to test the preference between spring-grazed and summer-grazed polder swards. We furthermore compared patterns of use by geese between long-term ungrazed and grazed salt marshes. In May, there is a difference in grazing pressure by geese between polder pastures that are grazed or ungrazed during spring. In this month, the ungrazed polder pastures are abandoned and the geese shift to either the grazed polder pastures or to the salt marsh. Vegetation in the polder that had been spring-grazed had a lower canopy height and a higher tiller density than summer-grazed vegetation. The captive geese in the preference experiment showed a clear preference for vegetation that had been spring-grazed by sheep over ungrazed vegetation. Goose grazing pressure was negatively correlated to canopy height, both on the polder and on the salt marsh. Within the plant communities dominated byFestuca rubra andPuccinellia maritima, marshes that were intensively grazed by livestock generally had higher grazing pressure by geese than long-term ungrazed or lightly grazed salt marshes.     

Reintroduction of eelgrass (Zostera marina) in the Dutch Wadden Sea; review of research and suggestions for management measures

Eelgrass (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal flats. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.     

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.     

Stepwise acclimation—a method for estimating the potential euryhalinity of the gastropod Hydrobia ulvae

The White Sea gastropod Hydrobia ulvae (Pennant) was exposed to step-wise lowering or increase of the habitat salinity. The time allowed for acclimatization to the successive salinity levels was sufficient to complete non-genetic adaptation. In this way, the lower and upper salinity limits were extended. The tolerance limits obtained are assumed to be indicative of the capacity for non-genetic adaptation and to serve as a genotypical characteristic. The tolerance of specimens colleced from in situ conditions (mid littoral, 20 S) ranged between 14 and 34 S. After non-genetic adaptation, the lower tolerance value shifted to 6 S (adaptation limit), and the upper value to 76 S (final limit not reached). There is no reason for considering White Sea H. ulvae to represent a special physiological race of specimens from those on the coast of Great Britain.     

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.     

附着生物对沉水植物伊乐藻生长的研究

以沉水植物伊乐藻(Elodea nuttalii(Plant.)ST.John.)为研究对象,室内利用原位湖水进行培养,通过测定伊乐藻的生物量、叶绿素质量分数、光合作用速率等指标,研究了附着生物量的增加对沉水植物伊乐藻的影响。结果表明,随着附着生物处理量的增加,伊乐藻的生物量、叶绿素质量分数以及光合作用速率随之下降,50d后,各处理的生物量与对照相比分别减少了约5%~15%,叶绿素质量分数以及光合作用速率下降了20%~43%、10%~36%。同时也分析了丝状藻对沉水植物伊乐藻的遮阴作用,初步推测附着生物对沉水植物伊乐藻的影响可能是由于附着生物的遮荫作用。研究的结果为揭示富营养化湖泊沉水植被衰退的机理提供了理论依据。     

New evidence for habitat-specific selection in Wadden Sea Zostera marina populations revealed by genome scanning using SNP and microsatellite markers

Eelgrass Zostera marina is an ecosystem-engineering species of outstanding importance for coastal soft sediment habitats that lives in widely diverging habitats. Our first goal was to detect divergent selection and habitat adaptation at the molecular genetic level; hence, we compared three pairs of permanently submerged versus intertidal populations using genome scans, a genetic marker-based approach. Three different statistical approaches for outlier identification revealed divergent selection at 6 loci among 46 markers (6 SNPs, 29 EST microsatellites and 11 anonymous microsatellites). These outlier loci were repeatedly detected in parallel habitat comparisons, suggesting the influence of habitat-specific selection. A second goal was to test the consistency of the general genome scan approach by doubling the number of gene-linked microsatellites and adding single nucleotide polymorphism (SNP) loci, a novel marker type for seagrasses, compared to a previous study. Reassuringly, results with respect to selection were consistent among most marker loci. Functionally interesting marker loci were linked to genes involved in osmoregulation and water balance, suggesting different osmotic stress, and reproductive processes (seed maturation), pointing to different life history strategies. The identified outlier loci are valuable candidates for further investigation into the genetic basis of natural selection.     

The possible origin of Zostera noltii in the Canary Islands and guidelines for restoration

Seagrasses and their habitat are declining worldwide. Zostera noltii in the Canary Islands has been drastically reduced, mainly by anthropogenic disturbance, to three small surviving patches in a single harbor in Lanzarote. A previous genetic study, using neutral microsatellite markers, revealed that these patches consist of a single clonal individual. Here, an assignment test, using microsatellite data, was used to locate the most likely population of origin from a set of possible donor populations. Our results show that the Moulay Bousselham population in northern Morocco is assigned as the most likely population of origin (88%), although the probability of being at one generation time distance is low (2.7%). This result, however, allows locating the most closely related stands that may be the most successful donor populations for future restoration based on shoot or seed transplantation.     

Food sources used by sediment meiofauna in an intertidal Zostera noltii seagrass bed: a seasonal stable isotope study

In an intertidal Zostera noltii Hornem seagrass bed, food sources used by sediment meiofauna were determined seasonally by comparing stable isotope signatures (δ¹³C, δ¹⁵N) of sources with those of nematodes and copepods. Proportions of different carbon sources used by consumers were estimated using the SIAR mixing model on δ¹³C values. Contrary to δ¹⁵N values, food source mean δ¹³C values encompassed a large range, from −22.1 ‰ (suspended particulate organic matter) to −10.0 ‰ (Z. noltii roots). δ¹³C values of copepods (from −22.3 to −12.3 ‰) showed that they use many food sources (benthic and phytoplanktonic microalgae, Z. noltii matter). Nematode δ¹³C values ranged from −14.6 to −11.4 ‰, indicating a strong role of microphytobenthos and/or Z. noltii matter as carbon sources. The difference of food source uses between copepods and nematodes is discussed in light of source accessibility and availability.     

Control of microbial growth and of amphipod grazing by water-soluble compounds from leaves of Zostera marina

The role of minor components of the leaves of Zostera marina L. in altering the activity of micro-organisms directly (and indirectly by affecting amphipod grazers) was investigated in laboratory experiments, using plants collected at Roberts Bank (49°2N; 123°8W) on the west coast of Canada. Water-soluble extracts of eelgrass leaves inhibited the growth of a micro-alga (Platymonas sp.) and many marine bacteria at concentrations equivalent to as little as 10 mg dry leaf l^-1. The potency of leaf extracts was higher (1) in young, actively metabolizing tissue than in older leaves, and (2) in leaves collected during rapid growth in summer 1980 than during the following winter. Water-soluble inhibitors (especially phenolic acids) may explain the low biomass of epiphytes on actively growing leaves. Three phenolic acids inhibited the test micro-organisms at concentrations as low as 0.3 mg l^-1; caffeic acid was more potent than either protocatechuic or gentisic acid. Extracts of young leaves also inhibited grazing by amphipods [Eogammarus confervicolus (Stimpson)] on dead leaves. The time required for leaching of soluble inhibitors may account for the delay between the loss of leaves from the plants and the onset of grazing. Thus, interactions among the biotic components of the detritus ecosystem may be significantly modified by minor compounds in the leaves of Z. marina     

Analysis of the small-scale spatial patterns of free-living marine nematodes from tidal flats in the East Frisian Wadden Sea

Serial samples with a multiple corer were taken in order to understand the small-scale aggregation patterns in the nematode community. A total of 77 species was found in the sample. Fifteen dominant nematode species were examined by means of a dispersion analysis. Only one species (Metachromadora scotlandica) may be considered randomly distributed. A variogram analysis gave no proof of aggregation in that species, but all the other (aggregated) species displayed different spatial patterns: (1) a maximum/aggregation at the sample's margin, (2) two maxima/aggregations in the centre of the sample, (3) aggregation of individuals distributed at random over the entire sample (Metalinhomoeus typicus) and (4) aggregation of individuals distributed at random only in the centre of the sample (Paralinhomoeus ilenensis). These results allow the prognosis that distribution patterns are repetitive in five species (Microlaimus conothelis, M. marinus, Odontophora rectangula, Oncholaimellus calvadosicus, Viscosia rustica). Finally, by means of MDS-ordination (based on cosine similarity) two nematode associations were distinguished: one group represented by diatom-feeders (sensu lato), the other one comprised of DOM-users (sensu lato). Received: 13 May 1998 / Accepted: 15 January 1999     

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.     

Food sources of macrobenthos in an estuarine seagrass habitat (Zostera noltii) as revealed by dual stable isotope signatures

Stable carbon and nitrogen isotope analysis was used to examine the resources and position of macrobenthos in an estuarine seagrass food web in two sampling moments, during summer and winter. The contribution of each food source to the carbon requirements of consumers was estimated by a mixing model. The used carbon sources were largely seagrass associated, although seagrass tissues were utilized by only few species, and equally contributed to microphytobenthos and suspended particulate organic matter. Based on isotopic data, Lucinidae bivalves have an alternative trophic pathway via symbiosis with chemoautotrophic bacteria. Resource utilization inside and adjacent to seagrass beds did not differ significantly, implying that seagrass-associated inputs extend well beyond the borders of the vegetation patches.     

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     

Seasonal variations in nitrogen-fixation (acetylene reduction) and sulphate-reduction rates in the rhizosphere of Zostera noltii: nitrogen fixation by sulphate-reducing bacteria

Nitrogen-fixation (acetylene reduction) rates were measured over an annual cycle in meadows of the seagrass Zostera noltii Hornem in the Bassin d'Arcachon, south-west France, between March 1994 and February 1995, using both slurry and whole-core techniques. Measured rates using the slurry technique consistently overestimated those determined on whole cores, probably due to the release of labile organic carbon sources as a result of root damage during preparation of the slurries. Thus, the whole-core technique may provide a more accurate estimate of in situ activity, since disturbance of physicochemical gradients of oxygen, sulphide, nutrients and the relationship between the plant roots and the rhizosphere microflora is minimised. Rates measured by the whole-core method were 1.8- to 4-fold greater (dependent upon season) in the light than those measured during dark incubations, indicating that organic carbon diffusing from the plant roots during photosynthesis was an important factor in regulating nitrogen fixation in the rhizosphere. Additions of sodium molybdate, a specific inhibitor of sulphate-reducing bacteria (SRB) inhibited acetylene-reduction activity by >80% as measured by both the slurry and whole-core techniques throughout the year, inferring that SRB were the dominant component of the nitrogen-fixing microflora. A mutualistic relationship between Z. noltii and nitrogen-fixing SRB in the rhizosphere, based on the exchange of organic carbon and fixed nitrogen is proposed. Acetylene- and sulphate-reduction rates showed distinct summer peaks which correlated with a reduced availability of ammonium in the sediment and the annual growth cycle of Z. noltii in the basin. Overall, these data indicate that acetylene reduction (nitrogen fixation) activity in the rhizosphere of Z. noltii was regulated both by the availability of organic carbon from the plant roots and maintenance of a low NH ₄ ⁺ concentration in the vicinity of the plant roots due to efficient assimilation of NH ₄ ⁺ by Z. noltii during the growth season. Nitrogenfixation rates determined from acetylene-reduction rates measured using the whole-core technique ranged from 0.1 to 7.3 mg N m^-2d^-1, depending on season, and were calculated to contribute between 0.4 and 1.1 g N m^-2yr^-1, or 6.3 to 12% of the annual fixed nitrogen requirement of Z. noltii.     

Productivity of bacteria and microalgae and the effect of grazing by holothurians in sediments on a coral reef flat

Bacterial productivity in sandy sediments on reef flats at Lizard Island, Great Barrier Reef was determined from the rate of incorporation of tritiated thymidine into DNA. The study was conducted during January 1982 and July 1983. A small diurnal increase occurred in sediments having a dense population of microalgae. Bacterial production was 120 to 370 mg C m^-2 d^-1 in summer on reef flats, which was equivalent to 30–40% of primary production by benthic microalgae. In winter, rates of primary production by benthic microalgae and secondary production by bacteria were about one-half to one-fifth of those in summer. There was much variation in production, due to patchiness in the distribution of benthic microbes, especially microalgae. Doubling times for the bacteria in surface sediment were 1 to 2 d in summer and 4 to 16 d in winter on the reef flats. These high productivity values for bacteria indicated that a net input of organic matter to the sediment was needed to support the growth of bacteria. Sediment bacteria thus have a very important role in transforming organic matter on the reef flats. Grazing by Holothuria atra depressed both primary production and bacterial production. It was estimated that these holothurians ate about 10 to 40% of bacterial carbon produced each day in summer, and thus have an important role in the carbon cycle. Harpacticoid copepods were numerically important components of the benthic meiofaunal community and probably had a significant impact on bacterial density as grazers.     

Reintroduction of eelgrass ( Zostera marina ) in the Dutch Wadden Sea; Review of research and suggestions for management measures

Eelgras (Zostera marina L.) in the Dutch Wadden Sea historically covered an area varying from 65–150 km² in the eulittoral as well as the sublittoral zones. At present, this area comprises less than 1 km² eulittoral eelgrass stands, with an associated decrease in habitat diversity. The causes for this decline are presumably connected with the ‘wasting disease’ and the closure of the former Zuiderzee in the early 1930s resulting in increased tidal range and increased currents. After a slight recovery of the eelgrass populations on the intertidal flats a definite decline started in the early 1970s, possibly connected to increased turbidity. The present water quality and turbidity do not negatively influence eelgrass growth up to a depth of at least 0.6m below Mean Sea Level. Based on mesocosm experiments and field experiments it is concluded that re-establishment of eelgrass should be possible in sheltered bays and on unexposed tidal falts. The most suitable depths for a reintroduction are those between 0 and 20–40 cm below mean sea level.