Evidence for effects of Spartina anglica invasion on benthic macrofauna in Little Swanport estuary,Tasmania

Spartina anglica is an exotic perennial grass that can rapidly colonise the intertidal zone of temperate estuaries and lagoons. Consequently, there is considerable concern about its impact on estuarine flora and fauna. This study provides the first investigation of ecological impacts by S. anglica in Australia. The objective was to investigate the impacts of S. anglica on benthic macroinvertebrate communities inhabiting mudflat and native saltmarsh habitats at Little Swanport estuary, Tasmania. The null hypothesis that species richness and species abundance of benthic macroinvertebrates in exotic S. anglica marsh does not differ from adjacent native saltmarsh and mudflat habitats was tested. Eighteen species and 3716 macroinvertebrates were collected from 60 intertidal core samples in three habitats. Species richness, total abundance of invertebrates, crustacean abundance and mollusc abundance of mudflat communities were significantly (P < 0.05) lower when compared to those inhabiting adjacent S. anglica marsh and native saltmarsh. However, species richness and total abundance of invertebrates of native saltmarsh and S. anglica marsh did not differ significantly. Ordination of macroinvertebrate data clearly separated mudflat sites from vegetated sites but showed remarkable similarity between exotic and native vegetated sites.     

Invasive alien plants in marine protected areas: the <Emphasis Type="Italic">Spartina anglica</Emphasis> affair in the European Wadden Sea

The common cord-grass Spartina anglica, a fertile hybrid of S. maritima and S. alterniflora, was planted in the European Wadden Sea extensively during the late 1920s and 1930s to promote sediment accretion. After establishment, it colonised as a pioneer plant in the upper tidal zone, where it occurs frequently in coherent swards at the seaward front of saltmarshes and in patches on the tidal flats. Often, a conspicuous, almost monotypic, belt of S. anglica is formed. Over the last two decades, an increase in abundance and accelerated spread of S. anglica was observed, possibly promoted by warmer spring temperatures. This alien species may benefit from global warming, and there is considerable concern about its harmful impacts on the native biocoenoses and native biodiversity of the unique Wadden Sea ecosystem, encompassing effects on hydromorphodynamics and coastal protection. For a definitive assessment, however, an adequate quantification and comparison of documented and potential effects of S. anglica is important, but currently unavailable. Consequently, no management strategy exists for the prevention or restoration of the Wadden Sea ecosystem. Thus, the development of an alien species plan on the level of the Trilateral Cooperation on the Protection of the Wadden Sea is essential.     

Remedial habitat creation: does Nereis diversicolor play a confounding role in the colonisation and establishment of the pioneering saltmarsh plant, Spartina anglica?

Increasing concerns over global warming and expected sea level rises have led to the adoption of new coastal management strategies around the south-east coast of England. This paper explores the role played by the estuarine invertebrate Nereis diversicolor in limiting the colonisation and establishment of the invasive pioneering salt marsh plant, Spartina anglica. The biology of N. diversicolor is briefly reviewed and data from field experiments are presented demonstrating significant negative effects of worm abundance on transplanted S. anglica biomass. Laboratory-based experiments demonstrated significant negative effects of N. diversicolor abundance on the survival of S. anglica seeds transplanted to sediment cores. The importance of estuarine invertebrates in engineering the mudflat habitat may confound the foreseen ecosystem services and function provided by saltmarsh management schemes. Received: 15 February 1999 / Received in revised form: 4 June 1999 / Accepted: 11 June 1999     

Impacts of an alien species (Spartina alterniflora) on the macrobenthos community of Jiangsu coastal inter-tidal ecosystem

Spartina alterniflora, a species vegetating on inter-tidal flats that was introduced from the eastern coast of United States, has become a hot topic, focusing on its invasion within local species in the coastal zone of China. Impacts of S. alterniflora on the inter-tidal macrobenthos community in the Jiangsu coastland are addressed by comparing the macrobenthos characteristics in a mudflat and in a four-year-old Spartina salt marsh that had earlier been a mudflat. During the period October 2002–July 2003, we studied the distribution pattern and diversity of macrobenthos, and discussed their correlation with environmental factors caused by Spartina vegetation. The results showed that a total of 43 macrobenthos species were found, mainly consisting of Mollusca, Crustacea, and Annelida. Ten macrobenthos species were found in the Spartina salt marsh, and 36 species were found in the mudflat. Life forms and functional groups of macrobenthos in the Spartina salt marsh were obviously distrinct from that of the mudflat. The study showed that macrobenthos diversity in the Spartina salt marsh decreased, and the community structure altered obviously, whereas the biomass showed no differences in different seasons. Statistical analysis demonstrated that seasonal change of macrobenthos diversity in the Spartina salt marsh negatively related to content of sediment organic matter, total N, bulk density, height and biomass of Spartina vegetation, and positively related to the density of Spartina. All these differences suggested the obvious effects of the Spartina vegetation on the Jiangsu inter-tidal benthic macroinvertebrate ecology. Furthermore, the investigation also showed that the niche of the native macrobenthos living in the mudflat has been transferred down, seaward, due to the invasion of Spartina in our study site.     

Rapid colonization of new habitats in the Wadden Sea by the ovoviviparousLittorina saxatilis (Olivi)

The intertidal periwinkleLittorina saxatilis completely lacks larval dispersal and adult vagility is low. Although this suggests a low dispersal rate,L. saxatilis is frequently found in recently established habitats “exotic” to the Wadden Sea. Populations occur on man-made structures like dikes, breakwater and groynes, some of which are not older than several years. Furthermore,L. saxatilis is found on marsh grassSpartina anglica, introduced to the Wadden Sea in the 1920s and 1930s, as well as on mats of green macroalgae, which have become an abundant feature on the tidal flats since the late 1970s. Seagrass beds are likely to be the original habitat ofL. saxatilis in the Wadden Sea. Since seagrass populations have dramatically declined over the last decades, colonization of new habitat types enabledL. saxatilis to maintain its Wadden Sea populations despite a changing environment. Colonizers can reach new habitats by means of passive transport, especially by rafting on macrophytes and by aerial dispersal attached to birds. In thew Wadden Sea, the ovoviviparously reproducingL. saxatilis has demonstrated its ability to successfully found new populations with only a few individuals. No reduction of genetic variablility (founder effect) was observed in recently established populations.     

Recently established <Emphasis Type="Italic">Crassostrea</Emphasis>-reefs versus native <Emphasis Type="Italic">Mytilus</Emphasis>-beds: differences in ecosystem engineering affects the macrofaunal communities (Wadden Sea of Lower Saxony,southern German Bight)

Since 1998 the non-indigenous Pacific oyster Crassostrea gigas (Thunberg 1793) has been invading the Wadden Sea of Lower Saxony, southern German Bight. C. gigas settles predominantly on intertidal Mytilus-beds (M. edulis L.) and subsequently create rigid reef-like structures. Both bivalve species are ecosystem engineers in sedimentary tidal flats. They provide hard substrate for sessile species, mobile organisms find refuge within the habitat matrix of dense suspension feeders, and biodeposits enrich the sediments with organic matter. The transformation of Mytilus-beds into Crassostrea-reefs gives rise to the question whether the invader may affect the native community. We investigated two parts of a changing bivalve bed in the backbarrier area of the island of Juist in March 2005. One part was still dominated by M. edulis whereas the other part was already densely colonized by C. gigas. Crassostrea-reefs compensate for the conceivable loss of Mytilus-beds in the intertidal of the Wadden Sea by replacing the ecological function of M. edulis. There was no indication of a suppression of indigenous species. This even applied to M. edulis, which persisted at the site invaded by C. gigas. The associated macrofaunal community showed increased species richness, abundance, biomass, and diversity in the Crassostrea-reef. The latter particularly favored sessile species like anthozoans, hydrozoans, and barnacles. Higher abundance and biomass for vagile epizoic species like the shore crab Carcinus maenas and the periwinkle Littorina littorea also occurred among oysters. Abundance of deposit feeding oligochaetes was enhanced by oysters as well. More opportunistic, facultative filter-feeding polychaetes occurred in the Crassostrea-reef.     

Food preferences of Shorelarks Eremophila alpestris,Snow Buntings Plectrophenax nivalis and Twites Carduelis flavirostris wintering in the Wadden Sea

Capsule Seeds of plants from lower salt marsh communities are preferred, with insects less important.

Methods Droppings of Shorelarks Eremophila alpestris, Snow Buntings Plectrophenax nivalis and Twites Carduelis flavirostris sampled in the German Wadden Sea were analysed and compared with food abundance to assess preferences.

Results Shorelarks prefer seeds of Salicornia sp., Suaeda maritima, Atriplex sp., Halimione portulacoides and unidentified small grass seeds. Insects are eaten mainly in periods of seed shortage, but are consumed in smaller amounts during the winter. The food composition of Snow Buntings is very similar, but additionally Triglochin maritimum is commonly eaten. Twites are specialized on seeds of Salicornia sp. and Suaeda maritima and rarely ingest other seeds and insects. All seeds consumed were of plants from lower salt marsh communities. Many halophyte seeds, and especially those that birds feed on, are rich in energy. Large seeds and those which need a long handling time are avoided.

Conclusion Changes in the lower salt marshes of the Wadden Sea by embankments and intensified grazing might have been responsible for the rapid population decline from the 1960s to 1980s.     

Macrobenthos of a tidal impoundment at the Mai Po Marshes Nature Reserve,Hong Kong

The composition and spatial distribution patterns of the macrobenthic faunal assemblages of an 8-ha tidal impoundment operated as a traditional shrimp pond at the Mai Po Marshes Nature Reserve, Hong Kong, were studied in relation to temporal changes in local environmental conditions. Species richness, abundance and biomass of macrobenthos across 5 different sub-habitats (seaward, middle, and landward parts of open water unvegetated areas, and Phragmites- and Kandelia-dominated, vegetated areas) within the pond were examined bimonthly between January 1997 and January 1998. Grab samples were collected randomly within the sub-habitats. Key physical environmental parameters of the sampling sites were also measured. A total of 46 species of macrobenthos was recorded: 11 polychaetes, 11 molluscs, 13 crustaceans and 11 insects. Mean species density in the five sub-habitats ranged from 0 to 3907 indm^–2, with mean biomass ranging from 0 to 96.9 gm^–2. The macrobenthos showed spatial and temporal differences among the sub-habitats and across sampling times. Species abundances of Polychaeta, Mollusca and Crustacea were significantly higher in the three open water areas than in the two vegetated (Phragmites- and Kandelia-dominated) areas, with an inverse pattern for Insecta. There were no clear temporal patterns although abundance and biomass generally increased in the cooler months. Results of a canonical correspondence analysis suggest that macrobenthic species richness, abundance and biomass in the open areas were positively correlated with salinity, while water depth, dissolved oxygen and sediment organic matter content had little relationship with the macrobenthic assemblage parameters. Ordination by multi-dimensional scaling suggests that different habitats supported distinct macrobenthic assemblages. The macrobenthic assemblage in the tidal pond was less species rich but denser than those in the neighboring tidal mangrove and mudflat, suggesting that conversion of these areas into extensively managed tidal aquaculture ponds results in reduced species richness in tropical mangrove habitats.     

Restoration of Shorebird‐Roosting Mudflats by Partial Removal of Estuarine Mangroves in Northern Taiwan

Expansion of the monospecific mangrove, Kandelia obovata, has converted intertidal mudflats and other habitats into mangrove forests, thus reducing estuarine biodiversity in the Danshuei River estuary, northern Taiwan. Dense mangrove vegetation was removed to create a small patchwork of mudflats and a tidal creek in February 2007. Subsequent changes in sediment properties and biodiversity of the macrobenthos and avian communities were examined. The results showed that the creation of different habitats led to changes in sediment properties and biodiversity. The water content and sorting degree of the sediments differed significantly among the restored mudflat, the tidal creek, and the mangrove control site. Silt/clay, organic carbon content, and chlorophyll a concentrations varied seasonally, but not among sites. The abundance of polychaetes in the creek was greater than that in the mudflat or the mangrove (12.5 vs. 5.3 and 2.2 individuals/m², respectively), suggesting preferential colonization of infaunal polychaetes in habitats with prolonged submersion. Crabs showed seasonal changes in density, with higher densities in summer than in autumn and winter. The species richness of wintering shorebirds on the created mudflat increased dramatically from 2002 to 2007. The transformation of a vegetated area into an open mudflat appeared to benefit shorebirds by providing roosting habitat. Our study demonstrated that controlling the spread of estuarine mangrove forests could increase biodiversity, and could particularly benefit the migratory shorebird community.     

Restoration of Aegiceras corniculatum mangroves in Jiulongjiang Estuary changed macro-benthic faunal community

Macro-benthic faunal communities were compared between non-vegetation mudflat and Aegiceras corniculatum mangroves with different ages in Jiulongjiang Estuary, China. Faunal species number was highest in the mature mangrove and was higher in mangroves than in the mudflat, as snails and some crustaceans species were only collected in mangroves. The 5-year-old mangrove had the highest infaunal abundance and crustacean biomass. Snails had more abundance in the young mangroves. Uca arcuata was the dominant crab species in the non-vegetation mudflat and 5-year-old mangrove. Mangrove vegetation and sediment characteristics analyses indicated different habitats due to A. corniculatum mangrove restoration. However, overall poor correlations between faunal assemblage and sediment properties indicated that sediment properties were not the major factors influencing faunal distribution.     

Mass mortality in two common soft-bottom invertebrates,Hydrobia ulvae andCorophium volutator-the possible role of trematodes

Two littoral macrofaunal invertebrates,Hydrobia ulvae (Prosobranchia) andCorophium volutator (Amphipoda) suffered mass mortality on an intertidal mudflat in the Danish Wadden Sea in May–June 1990. Dissection of collectedH. ulvae individuals revealed a considerable increase from March to May in numbers of infected individuals by microphallid trematodes that useH. ulvae andC. volutator as first and second intermediate host, respectively. The numbers of infested snails were hereafter reduced by an amount equal to the observed mortality rate of snails. At the same time, theC. volutator population became extinct. Since other conceivable mortality factors could be ruled out, parasites are suspected to be the causative agent. Apart from the expected effects on potential predators by the decline in the two invertebrate populations, the benthic community changed and destabilization of the substratum occurred probably because of the die-off inC. volutator. Meteorological data suggest high temperatures as a triggering factor of the massdevelopment of the studied trematodes.     

The introduced ribbed mussel (<Emphasis Type="Italic">Geukensia demissa</Emphasis>) in Estero de Punta Banda,Mexico: interactions with the native cord grass, <Emphasis Type="Italic">Spartina foliosa</Emphasis>

Introduced populations of Guekensia demissa occur on the west coast of North America. They have been reported in San Francisco Bay, four southern California wetlands, and in Estero de Punta Banda (EPB), Baja California Norte, Mexico. We randomly sampled benthic invertebrates in four habitat types within EPB: marsh, channel, mudflat and pan. Geukensia demissa was the most abundant bivalve in the wetland at EPB. It was significantly associated with the native cordgrass, Spartina foliosa, and occurred at higher average densities in vegetated marsh sites (24/m²) and Spartina-dominated tidal channels (35/m²), compared to mudflat (0/m²), and pan (0/m²) sites. We estimated that the total biomass of this invader was over four times that of the next most abundant bivalve, Tagelus spp., in EPB. We examined G. demissa for parasites and found that only a few native parasites colonized this introduced host at very low prevalences and intensities. We performed bird surveys to determine the habitat overlap and potential impact of this mussel on the EPB population of light-footed clapper rails (Rallus longirostrus levipes), an endangered species in the United States. The high abundance of G. demissa in EPB, its presence in clapper rail habitat, and its known effects on salt marsh habitat in it’s native range, warrant further investigations of the impact of this invader in EPB and elsewhere.     

Is spread of the neophyte Spartina anglica recently enhanced by increasing temperatures?

The introduced cordgrass Spartina anglica, a fertile hybrid of S. maritima and S. alterniflora, grows as a pioneer plant in the upper intertidal zone and has invaded most sheltered shorelines of the Wadden Sea. After its introduction in 1927 S. anglica has spread vigorously along the mainland shore and on some of the more southern islands. In contrast, it has later established on Sylt and spread at a lower pace. On the island of Sylt it occurs near at its northern limit in Europe. Due to rising sea level and storm frequency a decrease or steady state of S. anglica was expected because its niche is narrowing rather than widening and plants are more frequently eroded. Contrary to that, many new sites were colonized and dense monotypic swards have formed after 1985. This new spread coincided with a shift in the local temperature regime around 1987. The monthly mean temperature from January to April has increased significantly after 1987. Furthermore, the important physiological thresholds of 4 °C for germination and 7 °C for photosynthesis were more often exceeded during spring after 1987 than before. We suggest that warmer spring seasons since 1988 could have promoted germination, growth and the recent accelerated spread of this neophyte.     

Limited seed retention during winter inhibits vegetation establishment in spring,affecting lateral marsh expansion capacity

Coastal systems worldwide deliver vital ecosystem services, such as biodiversity, carbon sequestration, and coastal protection. Effectivity of these ecosystem services increases when vegetation is present. Understanding the mechanisms behind vegetation establishment in bio‐geomorphic systems is necessary to understand their ability to recover after erosive events and potential adaptations to climate change. In this study, we examined how seed availability affects vegetation establishment in the salt marsh–intertidal flat transition zone: the area with capacity for lateral marsh expansion. This requires vegetation establishment; therefore, seed availability is essential. In a 6‐month field experiment, we simulated a before and after winter seed dispersal at two locations, the salt‐marsh vegetation edge and the intertidal flat, and studied seed retention, the seed bank, and the seed viability of three pioneer marsh species: Salicornia procumbens, Aster tripolium, and Spartina anglica. During winter storm conditions, all supplied seeds eroded away with the sediment surface layer. After winter, supplied seeds from all three species were retained, mostly at the surface while 9% was bioturbated downwards. In the natural seed bank, A. tripolium and S. anglica were practically absent while S. procumbens occurred more frequently. The viability of S. procumbens seeds was highest at the surface, between 80% and 90%. The viability quickly decreased with depth, although viable S. procumbens seeds occurred up to 15 cm depth. Only when seeds were supplied after winter, many S. procumbens and some S. anglica individuals did establish successfully in the transition zone. Viable seed availability formed a vegetation establishment threshold, even with a local seed source. Our results suggest that, although boundary conditions such as elevation, inundation, and weather conditions were appropriate for vegetation establishment in spring, the soil surface in winter can be so dynamic that it limits lateral marsh expansion. These insights can be used for designing effective nature‐based coastal protection.     

Microphytobenthic dynamics in a Wadden Sea intertidal flat – Part II: Seasonal and spatial variability of non-diatom community components in relation to abiotic parameters

The microphytobenthos colonizing the intertidal flats forms an important component of the Wadden Sea. Ten sampling points along a 1-km transect were studied in a fringe area of the Solthörn tidal flat, southern North Sea, in order to determine seasonal differences in the microphytobenthos. An accompanying paper deals with the major component of the flora, the diatoms; here we, focus on the minor taxonomic groups. From May 2008 to May 2009 surface sediments were collected during low tide. Variation of environmental factors as well as microphytobenthic density (abundance and chlorophyll a) were monitored. The area investigated was a mixed-sediment mudflat, with a gradient from coarse to fine. Highest biomass was obtained in summer 2008 with 215.9?±?12.6?mg chlorophyll a m^–2. In late autumn the chlorophyll a concentration decreased continuously at all investigated stations. Lowest values were detected in December 2008. Species abundances varied considerably both along the transect and seasonally, depending on species-specific requirements as well as hydrodynamic conditions (tidal currents). Higher densities of benthic pro- and eukaryotic microalgae were observed in sites characterized by fine sediments. Apart from the diatoms, the most abundant microphytobenthic group was the cyanophytes. Coccoid cyanophytes, mainly Merismopedia sp., were most abundant during summer, with cell numbers up to 5.72?×?10⁶ cells cm^?2, while diatoms dominated in winter, spring and autumn. Filamentous cyanophytes, particularly Microcoleus chthonoplastes, were most abundant during autumn, while coccoid chlorophytes (spring: Chlorococcum submarinum, Crucigenia tetrapedia, Tetraselmis suecica), euglenophytes (summer: Euglena obtusa), dinophytes (autumn: Amphidinium operculatum, A. herdmanii) and cryptophytes (autumn: Hillea marina, Hemiselmis virescens) contributed to the microphytobenthos during warmer seasons. The statistical analysis confirmed that the composition of the microphytobenthos was related to sediment features and to characteristics of particular seasons.     

The American slipper limpet <Emphasis Type="Italic">Crepidula fornicata </Emphasis>(L.) in the northern Wadden Sea 70 years after its introduction

In 1934 the American slipper limpet Crepidula fornicata (L.) was first recorded in the northern Wadden Sea in the Sylt-R?m? basin, presumably imported with Dutch oysters in the preceding years. The present account is the first investigation of the Crepidula population since its early spread on the former oyster beds was studied in 1948. A field survey in 2000 revealed the greatest abundance of Crepidula in the intertidal/subtidal transition zone on mussel (Mytilus edulis) beds. Here, average abundance and biomass was 141 m^–2 and 30 g organic dry weight per square metre, respectively. On tidal flats with regular and extended periods of emersion as well as in the subtidal with swift currents in the gullies, Crepidula abundance was low. The main substrate of attachment was live mussels. Compared with the years following their initial introduction, Crepidula is more abundant today and has shifted from the now extinct oyster beds to the epifaunal community of the mussel beds. Their present abundance is considerably lower than at more southern European coasts where the species may dominate the epifauna. Low winter temperatures are suggested to have limited the population expansion in the northern Wadden Sea until now. Electronic Publication     

Foraging strategies involved in habitat use of shorebirds at the intertidal area of Chongming Dongtan, China

Dense flocks of migratory shorebirds from diverse species often concentrate in the intertidal areas for stopover. Trophic structure, food partition, prey availability and selectivity, predation risk, and abiotic factors are often used to explain the differences in habitat use of coexisting shorebirds. We sampled the macrobenthos and surveyed the distribution of shorebird populations to study the effects of foraging strategies on the habitat use of shorebirds at Chongming Dongtan, an important stopover site for shorebirds on the East Asian–Australasian Flyway. Results show that the relative abundance of epifaunal macrobenthos in salt marshes was much higher than that in the bare flats, whereas the relative abundance of infaunal macrobenthos in salt marshes was much lower than that in bare flats. The relative abundance of two life forms of macrobenthos was similar in the transitional zones between the salt marshes and the bare flats. Shorebirds with different foraging strategies exhibited different habitat uses. Pause-travel shorebirds mainly utilized the salt-marsh fringes, while tactile continuous shorebirds relied heavily on the bare flats. There was no significant difference in habitat use for visual continuous shorebirds. The density of tactile continuous shorebirds was positively correlated with bivalve density, and that of visual continuous shorebirds positively with crustacean density. Meanwhile, the relative abundance of pause-travel foraging shorebirds was positively correlated with the relative abundance of epifaunal, but negatively with infaunal macrobenthos. In contrast, the relative abundance of tactile foraging shorebirds had a positive correlation with infaunal but a negative one with epifaunal life form. Therefore, foraging strategies may play important roles in shorebirds’ habitat use in intertidal areas.     

The significance of spatial and temporal patterns of algal mat deposition in structuring salt marsh vegetation

Question: Are there hot spots of algal mat deposition in space and time at the marsh scale and, if so, how does this affect the coexistence of a dominant (Spartina anglica) and gap dependent (Salicornia europaea) species? Location: The Rattekaai salt marsh in the Scheldt estuary in the southwestern Netherlands (NW Europe). Methods: Mat cover and the abundance of the gap dependent species Salicornia europaea were monitored at the scale of a marsh. The effects of mat cover on the vegetation structure were studied by applying three mat removal treatments over three growing seasons. Results: The low marsh border was found to be a hot spot of algal mat deposition during the growing season, which had a correlated spatial pattern between two successive years at a 20 m X 20 m scale. The combination of duration, timing and repetition of mat cover determined growth inhibition of the competitive dominant Spartina anglica, and thereby the abundance of subordinates such as Salicornia europaea. Mat cover reduced the storage of carbon reserves in Spartina and our results imply that repetition of non‐lethal mat cover can lead to ‘gap creation’. Gaps gave only temporary habitat to less dominant species since Spartina quickly re‐invaded them. The gap dependent annual Salicornia was most abundant at intermediate levels of disturbance measured as a function of both space and time. Conclusions In addition to disturbance level, the spatial and temporal distribution of disturbance are important in creating and maintaining habitat for gap dependent species. Relatively small disturbances will have a large effect on diversity if the spatial and temporal distribution of the disturbances leads to ‘disturbance hot spots’.     

Long‐term vegetation changes in experimentally grazed and ungrazed back‐barrier marshes in the Wadden Sea

Abstract. Vegetation succession in three back‐barrier salt marshes in the Wadden Sea was studied using a data set comprising 25 years of vegetation development recorded at permanent quadrats. The effect of livestock grazing on succession was assessed by comparing quadrats where grazing was experimentally prevented or imposed. We studied changes at the species level as well as at the level of the plant community. Special attention is given to effects on plant species richness and community characteristics that are relevant for lagomorphs (hares and rabbits) and geese. Inundation frequency and grazing were most important in explaining the variation in species abundance data. The three marshes studied overlap in the occurrence of different plant communities and the observed patterns were consistent between them. Clear differences in frequency and abundance of plant species were observed related to grazing. Most plant species had a greater incidence in grazed treatments. Species richness increased with elevation, and was 1.5 to 2 × higher in the grazed salt marsh. Grazing negatively influenced Atriplex portulacoides and Elymus athericus, whereas Puccinellia maritima and Festuca rubra showed a positive response. The communities dominated by Elymus athericus, Artemisia maritima and Atriplex portulacoides were restricted to the ungrazed marsh. Communities dominated by Puccinellia maritima, Juncus gerardi and Festuca rubra predominantly occurred at grazed sites. As small vertebrate herbivores prefer these plants and communities for foraging, livestock grazing thus facilitates for them.     

Impact of the nemertean Lineus viridis on its polychaete prey on an intertidal sandflat

Interactions of the predator Lineus viridis (O. F. Muller 1774) with infaunal polychaetes were investigated in the German Wadden Sea. L. viridis is the dominant heteronemertean in this intertidal area with a mean abundance of 14 individuals per square meter. Predator enclosure experiments in the field revealed that L. viridis preyed on predatory polychaetes. The preferred prey species was Nereis diversicolor; others were Anaitides maculata. Nephtys hombergii and Harmothoe sarsi. These results were supported by a series of feeding experiments in the laboratory. The rate of consumption was estimated by laboratory and field experiments. The feeding rate of Lineus viridis was estimated by laboratory enclosure experiments to 5.6 g ashfree dry weight (AFDW) per square meter annually.