Environmental characteristics and distribution of macrobenthos in a mudflat of the west coast of Korea (Yellow Sea)

The spatio-temporal changes of environmental factors and the distribution patterns of macrobenthos were investigated on an intertidal mudflat near Panweol on the west coast of Korea. Temperatures of the surface sediments and salinities of the intertitial water were highest in summer and lowest in winter, while water content of the sediment was lowest in summer and highest in winter. Hourly variations of sediment temperature, water content and interstitial salinity during the daytime depended largely upon both the timing of ebb tide and daylight intensity.Temperature, water content and interstitial salinity showed a marked variation with sediment depth. Temperatures decreased sharply from the surface to 10 or 15 cm depth, and below this depth they were nearly constant, except in summer. Water content and interstitial salinity underwent a rapid decline in this upper layer and began to rise from this depth. From the high tide mark toward the main tidal channel, water content increased but salinity decreased. The sand fraction increased with the decrease of clay content toward the main tidal channel. This tidal flat could be divided into two zones: the “upper intertidal zone” and the “lower intertidal zone”, based on the spatial differences of the water content, interstitial salinity and grain-size composition in the surface sediment.There were three distinct zones of macrobenthos. The upper intertidal zone was dominated by burrowing deposit feeders, such as Ilyoplax dentimerosa, Helice tridens sheni, Cleistostoma dilatatum and Perinereis vancaurica tetradentata. The middle intertidal zone was characterized by other burrowing deposit feeders, such as Ilyoplax pingi, Macrophthalmus japonicus and Periserrula leucophryna. And the lower intertidal zone was dominated by suspension feeders represented by Laonome tridentata and Potamocorbula amurensis. Each zone was named after these characteristic species.Although the species composition of the upper intertidal community was different from that of the middle intertidal, the way of living and the feeding behaviour of the species occurring in these two zones were very similar. But the lower intertidal zone showed a characteristic community structure differing not only in species composition but also in ways of living and feeding of the species. Some sets of environmental factors are considered to explain the zonation. Water content of the sediment and interstitial salinity will be the main factors causing the observed zonation.     

The effects of bioturbation on sediment transport on an intertidal mudflat

Laboratory flume studies were conducted to determine the relative importance of various biological and physical factors controlling sediment erosion threshold and transport rate on an intertidal mudflat in the Bay of Fundy. Studies were conducted in a July period of maximum solar exposure. The upper and mid-intertidal stations of the flat were dominated by silt-clay sediments, while the lower intertidal was dominated by very fine sand. The tube-dwelling amphipod Corophium volutator was the most abundant infaunal species with densities exceeding ∼1300 ind·m⁻² based on counts of burrow openings. Sediment-penetrometry and water-content measurements indicated no change in unconsolidated shear strength and porosity, respectively, along the intertidal transect. Despite the apparently cohesive nature of the sediment, erosion occurred as small ripples. Critical shear velocities (u_crit) for erosion determined with intact cores in a laboratory flume were relatively consistent between stations and sampling dates (mean = 2.1 cm·s⁻¹ ± 0.2 SD), with no relationship to Corophium density, sediment chlorophyll a, or physical variables. Field-treatment of sediment with formalin did not cause an obvious change in u_crit as determined by flume experiments. Corophium seemed to have little effect on erosion thresholds because incipient motion could be observed between tube burrows, beyond the local influence of the amphipod. In contrast to erosion thresholds, sediment-erosion rates measured with bedload traps were negatively correlated with density of small Corophium, probably due to binding of sediment into burrows and the ambient sediment microfabric, all of which reduce the availability of sediment for transport. Adult amphipods, which occurred at low density probably due to territorial/competitive interactions, had no obvious effect on erosion rate since only a small proportion of the sediment surface was impacted by their bioturbation. Although a portion of the amphipod population exerts a stabilizing influence on sediment-erosion rates, concurrent studies at the site indicate that Corophium seasonally reduces the sediment-erosion threshold by grazing on microflora which would otherwise inhibit the initiation of grain motion. Due to this decoupling of erosion rate and threshold, it is necessary to measure both processes in assessing the effects of benthic biota on sediment transport.     

Animal traces on a tidal flat in Hangzhou Bay,China

Despite prevailing low salinity (S ranging from 1 to 10) of the Hangzhou Bay, its extensive (up to 2 km wide) flats along the northern coast have a species-rich macrobenthic fauna of crustaceans, bivalves, gastropods and polychaetes. These produce a variety of crawling, feeding, dwelling and resting traces, some comparable to those observed on the flats of Korea and Taiwan. The characteristic starlike feeding traces of the bivalve Tellina (Moerella) iridescens are described here for the first time. Sediment reworking appears to be largely confined to the surface layer. The burrow-inhabiting crabs are also mainly surface sediment feeders. These Chinese flats lack organisms reworking deeper sediment layers such as lugworms or callianassid crustaceans. As compared to the Wadden Sea, sedimentation rate is higher and subsurface sediment reworking less intense, resulting in relatively few bioturbation structures in vertical sections of the Hangzhou tidal flat sediments.     

Adaptation to anaerobic metabolism in two mussel species,Mytilus edulis and Mytilus galloprovincialis,from the tidal zone at Arcachon Bay,France

Aspects of anaerobic metabolism were investigated in two sympatric mussel species, viz. Mytilus edulis and Mytilus galloprovincialis, living in the tidal zone in Arcachon Bay, France. Specific activities of pyruvate kinase (PK) and phosphoenolpyruvate kinase (PEP-CK) were remarkably similar in the two sympatric species and generally corresponded more closely to those observed in M. galloprovincialis in the Mediterranean than with M. edulis in the Dutch Wadden Sea. However, the values for the radio PK: PEP-CK for the two species in Arcachon Bay agreed with those of intertidal M. edulis from the Dutch Wadden Sea.Succinate accumulation during the first 24 h of anaerobicsis was about the same as in M. galloprovincialis in the Mediterranean, but decreased during the second 24 h, particularly in M. edulis, obviously due to propionate formation. Decrease in ATP concentrations in the tissues during anaerobiosis corresponded to that of intertidal M. edulis from the Dutch Wadden Sea. With the exception of specific activities of PK and PEP-CK, all properties investigated in both species were as expected in intertidal mussels.     

Community structure and biomass distribution of seagrasses and macrofauna in the flores sea,Indonesia

In several locations in the Flores Sea region the community structure and the biomass distribution of seagrasses were studied along transects perpendicular to the shoreline. The share of each species within a sample plot was estimated, divided in above- and below-ground biomass. Statistics regarding substrate coverage, shoot density and leaf-area index were sampled. A standard relation was calculated between seagrass dry weight, ash-free dry weight and organic carbon content.The biotic data were related to environmental factors: DOC and nutrients in the water, salinity, tidal amplitude, sediment composition. A relation was estimated between bottom coverage of seagrasses and standing stock. Further calculations of biomass-production ratios allow a quick and rough estimate of seagrass productivity. Maximum above-ground biomass values (500–700 g AFDW·m⁻²) together with qualitative data indicate resource (= space) partitioning among the component seagrasses within a community, and suggest a carrying capacity of the reefflat habitat for seagrass density and biomass.A tentative model was constructed, starting from a constant, non-distributed multispecies vegetation in the lower intertidal and subtidal zone on sand and coral rubble, and moving into several suboptimal situations. The upper shore carries an impoverished, constrained vegetation (irregular tides, desiccation, harvesting). Sediment reworking by animals and physical displacement of sand disturbs the vegetation and favours pioneer species. Muddy habitats bordering mangroves carry monospecific stands showing extremely high biomass (e.g. below-ground Enhalus acoroides 3500 g AFDW·m⁻²). Thalassia hemprichii and Enhalus acoroides are the most constant species in all habitats mentioned.Macrofauna biomass within the seagrass beds fluctuated widely (maximum values 50–70 g AFDW·m⁻² in mixed seagrass vegetations) and only a weak relation between benthic macrofauna biomass and seagrass community structure and biomass was found.     

Biotope and biology of Armadillidium album Dollfuss,a terrestrial isopod of sandy beaches,in the SW Netherlands

The supralittoral isopod Armadillidium album Dollfus is common, although patchily distributed, under driftwood on the foreshore of broad sandy beaches on the outer coast of the Delta area in the SW Netherlands. The isopods are very tolerant of immersion in seawater, but are nevertheless confined to a narrow zone just above normal spring tides.A. album is a sexually reproducing isopod, with a single well-defined reproductive period in summer and a lifespan of two years. In spite of its very specialized biotope, the life cycle and reproductive strategy of A. album do not deviate substantially from those of related ubiquitous terrestrial isopods.     

Transport,deposition and in situ decay of seagrasses in a tropical mudflat area (Banc D'Arguin,Mauritania)

Extensive seagrass beds cover large areas of the mudflats and subtidal parts of the Banc d'Arguin (Mauritania, West Africa). The fate of seagrass biomass (mainly Zostera noltii and Cymodocea nodosa) was investigated during a 1-month expedition to the area (May 1988). Only minor amounts of leaf material were found to be transported by water currents at that time. Subtidal detritus depositions were not found in the area. Though conspicuous accumulations of leaf material were present on some parts of the shore, the amounts washed ashore per flood tide formed only a minor proportion of the daily seagrass production. Thus, transport of leaf litter away from the seagrass beds of the Banc d'Arguin appears insignificant. The major part of senescent leaves probably remains trapped within the seagrass beds and will decompose in situ. The observation that a large part of the particulate carbon present in the surface sediment of seagrass beds was derived from above-ground biomass (55% and 37% in Zostera and Cymodocea beds, respectively), is in agreement with this conclusion.Using the litterbag technique, the rate of leaf decomposition in a seagrass bed was investigated. The time required for 50% weight loss of Zostera and Cymodocea leaf litter in the intertidal zone of the seagrass bed was 158 and 50 days, respectively. In the subtidal zone these figures were 49 and 37 days.     

Flocking and feeding in the fiddler crab (UCA tangeri): Prey availability as risk-taking behaviour

For a full understanding of prey availability, it is necessary to study risk-taking behaviour of the prey. Fiddler crabs are ideally suited for such a study, as they have to leave their safe burrow to feed on the surface of the intertidal flats during low tide, thereby exposing themselves to avian predators. A study in an intertidal area along the coast of Mauritania showed that small crabs always stayed in the vicinity of their burrow, but large crabs wandered in large flocks (also referred to as droves) to feed on sea-grass beds downshore. Transplanting downshore feeding substrate to the burrowing zone of the small crabs proved that they too preferred to feed on it. Since small crabs can be preyed upon by more species of birds, this suggests that the decision not to leave the burrowing zone might be related to the risk of being fed upon by birds. We calculated predation risk from measurements on the density and feeding activity of the crabs, as well as the feeding density, the intake rate and the size selection of the avian predators. Per hour on the surface, crabs in a flock were more at risk than crabs feeding near their burrow. Thus, though flocking crabs may have benefited from ‘swamping the predator’ by emerging in maximum numbers during some tides only, this did not reduce their risk of predation below that of non-flocking crabs. Furthermore we found that irrespective of activity, large crabs suffered a higher mortality per tide from avian predators than small crabs. This suggests that large crabs could not sufficiently reduce their foraging time to compensate for the increased risk while foraging in a flock, even though they probably experienced better feeding conditions than small crabs staying near their burrow. The greater energy demands of large crabs were reflected in a greater surface area grazed. Thus, with increasing size a fiddler crab has to feed further away from its burrow and so may derive less protection from staying near to it. It seems that growing big does not reduce the risk of predation for fiddler crabs, as it does in many other species with indeterminate growth. As in such species, the most probable advantage of growing big is increased mating success. Ultimately, therefore, prey availability must be understood from the life-history decisions of the prey species.     

Scale and intensity of intertidal habitat use by knots Calidris canutus in the Western Wadden Sea in relation to food,friends and foes

In August–October 1988–1992 we studied the distribution and abundance of knots Calidris canutus around Griend in the western Wadden Sea, and the extent to which these can be explained by benthic prey availability and presence of avian predators. Numbers in the nonbreeding season showed monthly averages of 10 000 to 25 000 birds. Over 100 000 knots were recorded on three occassions. Knots feed in large flocks, individual birds usually experiencing 4 000 to 15 000 flock-mates. The Siberian-breeding/west-African wintering canutus subspecies passed through in late July and early August. Otherwise the Greenlandic/Canadian breeding islandica subspecies was present. Over the period 1964–1992 there were no clear trends in the number of knots, but canutus-knots were particularly abundant in July–August 1991, whereas in 1992 both subspecies were absent.Macoma balthica was the preferred prey of both subspecies. Hydrobia ulvae, Mytilus edulis and Cerastoderma edule were eaten when Macoma was absent close to the surface of the sediment. As Macoma buried deeper from July onwards, canutus faced better average feeding conditions than islandica later in the year. The spatial distributon of knots feeding on the intertidal flats around Griend was best explained by the harvestable biomass of the prevalent prey species in a particular year and season, i.e. Macoma (main prey when their harvestable biomass densities were greater than ca 0.8 g AFDM per m²) and Cerastoderma, and by the avoidance of situations where they run the risk of attack by bird-eating birds. Flocks of knots covered most of the intertidal flats in the Western Dutch Wadden Sea in a couple of tidal cycles. This is about 800 km², much larger than the equivalent area used by knots on their wintering grounds in Mauritania (10–15 km²), a difference that is correlated with prey spectrum, prey availability and predictability.     

Copper,zinc and cadmium in benthic organisms from the Java Sea and estuarine and coastal areas around East Java

A study was made of the concentrations of copper, zinc and cadmium in benthic organisms, representing the phyla Mollusca, Arthropoda, Echinodermata and Pisces, from the riverine and estuarine areas of the rivers Brantas and Solo (East Java) and the adjacent coastal area. Moreover, an assessment was made of the contamination of the benthic biota with these elements in the Java Sea and Bali Sea.Benthic organisms show a species-specific uptake pattern for each element. Compared to the same type of animals from estuaries and coastal areas in temperate regions of western Europe, the concentrations of cadmium are considerably higher, while copper and zinc concentrations are somewhat lower.There is no general trend in concentration levels of the metals in specimens from rivers, estuaries, coastal zone and open sea. In some groups of organisms (e.g. shrimp, starfish) the concentrations of copper and zinc are highest in specimens from rivers and estuaries. In contrast, cadmium concentration levels in e.g. crab, shrimp and squid are lowest in riverine and estuarine areas.Significant differences in metal concentrations in these organisms were found between the dry monsoon period (July, August) and the beginning of the wet monsoon (November, December).No relationship existed between the metal concentration of the organisms and the silt fraction of the sediment (grain size < 63 μm) or the bulk sediment.     

The macrozoobenthos of the subtidal western dutch wadden sea. I. Biomass and species richness

During a one-year period in 1981–1982, a survey was conducted on the macrozoobenthos of the subtidal areas of the western half of the Dutch Wadden Sea. In total 80 species were found, half of them polychaetes. In terms of biomass, Mytilus edulis dominated the macrozoobenthos, with Hydrobia ulvae, Heteromastus filiformis, Carcinus maenas and Macoma balthica as other important species. Numerically important were also the polychaetes Pygospio elegans and Scoloplos armiger. Average macrozoobenthic biomass amounted to 43.7 g·m⁻² ash-free dryweight. This value is in the same range as values from intertidal areas in the Wadden Sea. The relatively high value in comparison with data from similar subtidal areas is attributed to the important mussel culture in the area.     

An evaluation of intertidal feeding habitats from a shorebird perspective: Towards relevant comparisons between temperate and tropical mudflats

Macrozoobenthic communities of intertidal soft sediments are reviewed worldwide from the perspective of a mollusc-eating shorebird species. Based on 19 sites, total biomass figures varied between 5 and 80 g AFDM per m² (average 24 g AFDM per m²); no latitudinal trends are apparent. The contribution made by bivalves and gastropods varies between 1% and 99%, north-temperate intertidal flats having relatively more molluscs than tropical flats. Intertidal flats in the tropics contain a greater variety of taxa, with brachiopods in Indonesia and echinoderms in northwest Australia contributing significantly to biomass only there. Limits to the occurrence of avian predators of intertidal benthos are set by the harvestable fraction of the biomass on offer and the costs of living at a particular site. No systematic differences in the harvestable fraction of the total mollusc-biomass for a worldwide occurring shorebird species specializing on molluscs (knots Calidris canutus) were apparent between temperate and tropical intertidal areas, in spite of large differences in maintenance metabolism incurred by these birds. The harvestable fractions of bivalves in the two West African areas (Banc d'Arguin, Mauritania and Guinea-Bissau) tended to be high (23–84% of total biomass in six species), they were relatively low (2–52% in five species) in the temperate Wadden Sea and the tropical northwest Australian site. Harvestable biomass determines the intake rate of shorebirds, as illustrated by functional-response curves of knots feeding on two bivalves species. We argue that the collection of information on size-depth relationships along with faunal and biomass surveys at a range of sites is bound to greatly increase our understanding of both the biology of tidal-flat invertebrates and the resource base underpinning the spectacular seasonal migrations of shorebirds.     

De novo assembly of the complete organelle genome sequences of azuki bean (Vigna angularis) using next-generation sequencers

Since chloroplasts and mitochondria are maternally inherited and have unique features in evolution, DNA sequences of those organelle genomes have been broadly used in phylogenetic studies. Thanks to recent progress in next-generation sequencer (NGS) technology, whole-genome sequencing can be easily performed. Here, using NGS data generated by Roche GS Titanium and Illumina Hiseq 2000, we performed a hybrid assembly of organelle genome sequences of Vigna angularis (azuki bean). Both the mitochondrial genome (mtDNA) and the chloroplast genome (cpDNA) of V. angularis have very similar size and gene content to those of V. radiata (mungbean). However, in structure, mtDNA sequences have undergone many recombination events after divergence from the common ancestor of V. angularis and V. radiata, whereas cpDNAs are almost identical between the two. The stability of cpDNAs and the variability of mtDNAs was further confirmed by comparative analysis of Vigna organelles with model plants Lotus japonicus and Arabidopsis thaliana.     

Rj (rj) genes involved in nitrogen-fixing root nodule formation in soybean

It has long been known that formation of symbiotic root nodules in soybean (Glycine max (L.) Merr.) is controlled by several host genes referred to as Rj (rj) genes, but molecular cloning of these genes has been hampered by soybean’s complicated genome structure and large genome size. Progress in molecular identification of legume genes involved in root nodule symbiosis have been mostly achieved by using two model legumes, Lotus japonicus and Medicago truncatula, that have relatively simple and small genomes and are capable of molecular transfection. However, recent development of resources for soybean molecular genetic research, such as genome sequencing, large EST databases, and high-density linkage maps, have enabled us to isolate several Rj genes. This progress has been achieved in connection with systematic utilization of the information obtained from molecular genetics of the model legumes. In this review, we summarize the current status of knowledge of host-controlled nodulation in soybean based on information from recent studies on Rj genes, and discuss the future research prospects.     

Environmental effects of the growth rate of intertidal invertebrates and some implications for foraging waders

The paper describes effects of intertidal height and sediment type on growth rate of the bivalves Cerastoderma edule, Macoma balthica, Mya arenaria, Mytilus edulis and Scrobicularia plana, and of the worms Arenicola marina, Nephtys hombergii and Nereis diversicolor in the eastern part of the Dutch Wadden Sea. In most species, exposure time was negatively correlated with length growth, although interfering effects of sediment type could not be ruled out. When controlled for the effects of exposure time, clay content of the sediment appeared to affect the growth of all species, but in different ways. The variation was related to the foraging methods of the invertebrates. Foraging waders may use the spatial variation in growth rate of the invertebrates to optimize the exploitation of individual cohorts.     

The National BioResource Project (NBRP) Lotus and Glycine in Japan

The objective of the National BioResource Project (NBRP) in Japan is to collect, conserve and distribute biological materials for life sciences research. The project consists of twenty-eight bioresources, including animal, plant, microorganism and DNA resources. NBRP Lotus and Glycine aims to support the development of legume research through the collection, conservation, and distribution of these bioresources. Lotus japonicus is a perennial legume that grows naturally throughout Japan and is widely used as a model plant for legumes because of such advantages as its small genome size and short life cycle. Soybean (Glycine max) has been cultivated as an important crop since ancient times, and numerous research programs have generated a large amount of basic research information and valuable bioresources for this crop. We have also developed a “LegumeBase” a specialized database for the genera Lotus and Glycine, and are maintaining this database as a part of the NBRP. In this paper we will provide an overview of the resources available from the NBRP Lotus and Glycine database site, called “LegumeBase”.     

Kazusa Marker DataBase: a database for genomics,genetics, and molecular breeding in plants

In order to provide useful genomic information for agronomical plants, we have established a database, the Kazusa Marker DataBase (http://marker.kazusa.or.jp). This database includes information on DNA markers, e.g., SSR and SNP markers, genetic linkage maps, and physical maps, that were developed at the Kazusa DNA Research Institute. Keyword searches for the markers, sequence data used for marker development, and experimental conditions are also available through this database. Currently, 10 plant species have been targeted: tomato (Solanum lycopersicum), pepper (Capsicum annuum), strawberry (Fragaria × ananassa), radish (Raphanus sativus), Lotus japonicus, soybean (Glycine max), peanut (Arachis hypogaea), red clover (Trifolium pratense), white clover (Trifolium repens), and eucalyptus (Eucalyptus camaldulensis). In addition, the number of plant species registered in this database will be increased as our research progresses. The Kazusa Marker DataBase will be a useful tool for both basic and applied sciences, such as genomics, genetics, and molecular breeding in crops.     

What determines the densities of feeding birds on tidal flats? A case study on dunlin,Calidris alpina,in the Wadden Sea

The tidal and seasonal pattern of habitat utilization by dunlin was studied in a tidal-flat area in a shallow bay called Königshafen, island of Sylt, by counting the number of dunlins on marked plots at 10-min intervals over whole tidal cycles. Sediment type, tidal elevation, and seasonal changes in food choice were found to influence the feeding densities of dunlin irrespective of total numbers present in the area. Densities of dunlin were generally highest on low muddy substrates. A preference for high sandy substrates was found in late summer. The tidal pattern of habitat utilization differed between areas. In preferred areas dunlin stayed during the whole emersion period. Other areas were only used by following the moving tide line. Seasonal changes in habitat utilization were apparently triggered by changes in food choice. In spring dunlin fed almost exclusively on polychaetes. In late summer a relatively high proportion of shrimps, Crangon crangon, was found in the diet of dunlins. The preference for shrimps may force the birds to stay away from the tide line, where the shrimps can escape into deeper waters. It is recommended that studies on habitat utilization on tidal flats should consider the tidal behaviour of the target species, as single low-tide counts may not give valid results.     

Cone index and root growth in surface and subsurface microirrigated hardpan soil

Restricted root growth caused by subsurface hardpans and low water holding capacity reduces crop yields in many United States southeastern Coastal Plain soils. With intensive irrigation it is possible to obtain suitable yields without deep tillage. The objective of this study was to find differences of root growth and cone indices between surface and subsurface applied sources of irrigation water. We measured root growth and cone index in intensively managed irrigation plots of green beans (Phaseolus vulgaris L.) in 1988 and 1989. Microirrigation tubes were placed on the surface or in the subsurface — buried at a depth of approximately 0.25 m — and irrigated either continuously or with intermittent pulses of water. Mean profile cone indices for the surface tube placement were significantly lower than for the subsurface tube placement. Mean profile cone indices for the continuous irrigation treatment were significant lower than for the intermittent irrigation treatment. However, no one treatment significantly reduced the cone indices within the hardpan. Total root count was significantly greater for the subsurface tube placement with increased root growth below the 0.1-m depth. There was a high density of roots next to the subsurface tube that would have been effective in uptake of water from the microirrigation tube. However, yield was significantly greater for the surface tube placement. For all treatments, the largest concentration of root growth occurred in the top 0.2 m of the soil within the row. We expected this for the surface irrigation treatment. However, we also measured this for the subsurface treatment, probably because of high rainfall and upwelling of water from the subsurface micro-irrigation tube.     

The effect of organic pollution on local distribution of Nereis diversicolor and Corophium volutator

Distribution patterns of Nereis diversicolor and Corophium volutator in the SE-Dollard, a 92 km² tidal flat area in the NE Netherlands are described. Body condition as well as the maximal size of juvenile worms correlated positively with elevation of the mudflat, probably caused by a better food supply on the higher mudflats. No spatial variation in size was found for C. volutator.N. diversicolor disappeared from ∼1 km² of the intertidal mudflats in the autumn of 1985 as a consequence of the seasonal waste discharges, whereas C. volutator disappeared from an area of ∼4 km². Though the discharges decreased by almost 50% from 1982 to 1985, N. diversicolor disappeared from about the same area in the 2 years.N. diversicolor is the main prey species of the avocet Recurvirostra avosetta, large flocks of which stay in the Dollard during autumn migration. Though the avocets lost ∼30% of their potential feeding area in the Dollard in the 1970s because of the high waste discharges, against only 5 to 10% in 1982 and thereafter (at significant lower discharges), the numbers of the avocets did not respond to the decrease of the discharges.