Feeding effects of the keystone deposit feeder Ilyanassa obsoleta (Neogastropoda,Gastropoda) on sedimentary diatoms

Microphytobenthos (MPB), typically comprised mainly of diatoms, is a key contributor to nearshore energy flow and nutrient cycles. Deposit-feeding invertebrates are known to alter the structure and activity of MPB. The eastern mud snail Ilyanassa obsoleta can reach extremely high densities in estuaries of the northwestern Atlantic, and their deposit-feeding and locomotion strongly influence other invertebrates and microbes. Our objective was to explore quantitative and qualitative effects of this keystone deposit-feeder on diatoms of intertidal sediments. We gathered snails from mudflat and sandflat habitats and collected their fresh fecal pellets in the laboratory. DNA metabarcoding allowed us to characterize diatom assemblages of ingested sediments and feces. We noted selective feeding such that reduction in MPB biomass with gut passage was difficult to quantify. Diatom α-diversity was reduced with gut passage in snails from both sedimentary regimes. Mudflat and sandflat diatom assemblages were distinct and differed markedly between feces and sediment in mud-feeding snails, whereas the difference in sand-feeding snails was minor. The sandy habitat was dominated by a mix of epipelic and epipsammic diatoms. In contrast, mudflat samples were dominated by epipelic and planktonic diatoms. Compositional differences between sediment and feces reflected preferential removal of planktonic taxa. Our results suggest the importance of phytodetritus to the mud snail diet, particularly in hydrodynamically quiescent environments. Due to the natural spatial patchiness of the snails and the capacity for rapid microbial recolonization, field experiments are recommended to determine whether MPB community changes attributed to gut passage are manifested at the landscape scale.     

Differential emigration causes aggregation of the amphipod Corophium volutator (Pallas) in tide pools on mudflats of the upper Bay of Fundy, Canada

We investigated mechanisms responsible for the formation of spatial patterns in a dominant macro-invertebrate, the burrowing amphipod Corophium volutator, in relation to tide pools on intertidal mudflats of the upper Bay of Fundy, Canada. A field survey of 3 different mudflats showed that density of C. volutator at low tide was consistently higher inside than outside tide pools, and a manipulative field experiment indicated that C. volutator selected tide pools over the adjacent emerged areas. Differential survival in immersed and emerged areas at low tide did not explain the observed pattern, because short-term survival of amphipods was not affected by immersion at low tide. As well, immigration rates of C. volutator into immersed and emerged areas at low tide were similar after one day, so preferential settlement did not explain aggregation of amphipods. However, marked C. volutator departed from experimental areas that were immersed at low tide at a slower rate compared to emerged experimental areas. Thus, formation of the spatial pattern observed in the field (higher densities inside tide pools) is likely the result of an effect of immersion at low tide on proportion of individuals emigrating from an area every day.     

Spatial patterns of benthic diatoms,carbohydrates and mud on a tidal flat in the Ems-Dollard estuary

The chlorophyll a content and two operational fractions of carbohydrate (water extractable and EDTA extractable) were measured every three months during one year along transects on a tidal flat in the Ems-Dollard estuary (The Netherlands). Chlorophyll a was used as an indicator of microphytobenthos biomass, which was composed predominantly of epipelic diatoms. Both carbohydrate fractions correlated significantly with chlorophyll a. EDTA extractable carbohydrates were more resistant towards degradation than the water extractable fraction. During most of the year, concentrations of chlorophyll a and carbohydrates were low, but in June, high concentrations of up to 90 g chlorophyll a/g sediment were found in a narrow zone running parallel to the channel. Maximum concentrations of water extractable carbohydrates and EDTA extractable carbohydrates ranged between 800–1200 and 600–800 g/g sediment, respectively. The mud content was high (± 90%) at the margin of the tidal flat. This was not limited to the growth season of the diatoms, but was observed throughout the year. This indicated that the high mud content at the mudflat margin was mainly caused by hydrodynamic factors, and not by biostabilization. In June, exceptionally high diatom densities were found in sediment with a high mud content. There was only minor evidence that biostabilization by epipelic diatoms lead to a further increase in the mud content of the sediment.     

Frequency distribution studies of epipelic diatoms along an intertidal shore

Live and dead cell counts of epipelic diatoms were analysed at 16 sites along a transect crossing a saltmarsh, sandflat, and mudflat at Berrow Flats, Somerset, U.K. Replicate monthly samples were taken from January 1982 to June 1984. The ratio of dead to live cells varied according to position along the transect. Temporal changes of the ratios did not display a cyclic seasonal pattern. The clustered mosaic of live cells and empty frustules on surface sediments was quantitatively assessed. Graphing variance against seasonal mean cell counts showed that the variance was proportional to the square of the mean. The Neyman Type A equation is suggested as a possible model to explain this relationship.     

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.     

Abundance and distribution of small infauna in mangroves of Missionary Bay, north Queensland, Australia

To assess the occurrence, spatial distribution and species composition of small infauna on a mangrove shore, core samples were taken along a transect in Missionary Bay at Hinchinbrook Island, north-east Australia. Three sites were arranged within the mangrove forest and one site was located in an adjacent mudflat. The sites were surveyed four times between November 1988 and October 1989. Based on the records from all samples and sites, 39 taxa were identified. Diversity (H') ranged from 1.18 to 2.38. Overall, total abundances of small infauna (retained on a 0.25 mm sieve) reached a mean value of 5,477 ind.m-2, with little variation throughout the transect or over time. Species numbers and diversity were higher in the mudflat than at the mangrove sites. The taxonomic composition changed between the mangrove forest and the mudflat: Oligochaeta were more abundant in mangrove sediments, whereas Polychaeta dominated in the mudflat. Of the polychaetes, Capitellidae were almost restricted to the mangrove sites, Sabellidae were recorded frequently at all sites, and Sigambra parva and Myriochele sp. were confined to the mudflat and the mangrove fringe. These species accounted also for dissimilarities between sites. Multivariate analyses showed a distinct assemblage at the mudflat compared to the mangrove sites. This survey showed that small infauna is an abundant component of mangrove sediments, which has been previously underestimated.     

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.     

Shorebirds, snails, and the amphipod (Corophium volutator) in the upper Bay of Fundy: top–down vs. bottom–up factors, and the influence of compensatory interactions on mudflat ecology

During their annual mid- to late-summer southward migration, Semipalmated Sandpipers (Calidris pusilla) feed intensively on the amphipod Corophium volutator on intertidal mudflats in the Bay of Fundy. Corophium, in turn, feed on diatoms and bacteria. Using a series of bird exclosures and fertilizer addition, we examined top–down and bottom–up effects, and investigated the presence of a trophic cascade in the mudflat community during the period when birds are abundant. Although both top–down and bottom–up forces were present in this system, neither transmitted beyond a single trophic link. Predation by shorebirds, which may be less size-selective than previously thought, reduced Corophium abundance in control plots by approximately 80% relative to exclosures, but most other species were unaffected. Shorebird predation did not result in an increase in diatom abundance, as predicted under the trophic cascade hypothesis. Fertilizer increased diatom abundance, but had no effect on Corophium abundance or bird predation, and little effect on other mudflat invertebrates. The only indirect effect observed was on mud snails (Ilyanassa obsoleta), which, by rapidly responding to changes in diatom abundance, compensated for both bird exclusion and fertilizer addition, and prevented the trophic cascade. This population response by snails, possibly stemming from competition with Corophium, probably contributed to the stability of the community. Our results provide an example of short-term compensation in a simple intertidal community, and highlight the importance of considering direct and indirect effects in community ecological studies. We conclude that while compensatory interactions that block trophic cascades may be more common in more complex ecosystems, they are not restricted to them.     

Microphytobenthos,their Extracellular Polymeric Substances,and the Morphogenesis of Intertidal Sediments

Intertidal sediments are important areas that separate the land from the sea and form natural coastal defenses. They are known as highly productive ecosystems, fueling the coastal food web. It is also conceived that microphytobenthos contribute to the stability of intertidal sediments by increasing the erosion threshold and that they are major players in coastal morphodynamics. Depending on the sedimentary composition of intertidal flats, different types of microphytobenthos colonize the sediment surface. Fine sand sediment is often colonized by cyanobacteria, prokaryotic algae, which form dense and rigid microbial mats. Mudflats on the other hand are characterized by the development of thin biofilms of epipelic diatoms. Both groups of phototrophic microorganisms excrete extracellular polymeric substances (EPS), but they do so in different ways and for different reasons. Two operationally defined fractions, water- and EDTA-extractable EPS, have been obtained from intertidal diatom biofilms and from cultures. They differ in composition and their production seems to be under different metabolic control. Water-extractable EPS are considered to be closely associated with the diatoms and are rich in neutral sugars, notably glucose. These EPS show a dynamic relationship with the microphytobenthic biomass. EDTA-extractable EPS are tightly bound to the sediment, probably through bridging by divalent ions. This material is rich in uronic acids and other acid sugars and is weakly related to chlorophyll. These EPS have been conceived to be a major factor in the structuring and diagenesis of coastal sediments and essential for increasing the sediment erosion threshold. However, this relationship is now questioned.     

The meiobenthos of five mangrove vegetation types in Gazi Bay,Kenya

The vertical distribution of meiofauna in the sediments ofAvicennia marina,Bruguiera gymnorrhiza,Ceriops tagal,Rhizophora mucronata andSonneratia alba at Gazi Bay (Kenya), is described. Seventeen taxa were observed, with highest densities in the sediments ofBruguiera (6707 ind. 10 cm^–2), followed byRhizophora (3998 ind. 10 cm^–2),Avicennia (3442 ind. 10 cm^–2),Sonneratia (2889 ind. 10 cm^–2) andCeriops (1976 ind. 10 cm^–2). Nematodes accounted for up to 95% of total densities; other common taxa were copepods, turbellarians, oligochaetes, polychaetes, ostracods and rotifers. High densities occurred to about 20 cm depth in the sediment. EspeciallyCeriops sediments show still high densities of nematodes (342 ind. 10 cm^–2) and copepods (11 ind. 10 cm^–2) in the deepest layer (15–22 cm). Particle size and oxygen conditions were major factors influencing meiobenthic distribution;Uca burrows had a major impact on distribution and abundance of meiofauna.     

Prey Distribution,Physical Habitat Features,and Guild Traits Interact to Produce Contrasting Shorebird Assemblages among Foraging Patches

Worldwide declines in shorebird populations, driven largely by habitat loss and degradation, motivate environmental managers to preserve and restore the critical coastal habitats on which these birds depend. Effective habitat management requires an understanding of the factors that determine habitat use and value to shorebirds, extending from individuals to the entire community. While investigating the factors that influenced shorebird foraging distributions among neighboring intertidal sand flats, we built upon species-level understandings of individual-based, small-scale foraging decisions to develop more comprehensive guild- and community-level insights. We found that densities and community composition of foraging shorebirds varied substantially among elevations within some tidal flats and among five flats despite their proximity (all located within a 400-m stretch of natural, unmodified inlet shoreline). Non-dimensional multivariate analyses revealed that the changing composition of the shorebird community among flats and tidal elevations correlated significantly (ρ_s = 0.56) with the spatial structure of the benthic invertebrate prey community. Sediment grain-sizes affected shorebird community spatial patterns indirectly by influencing benthic macroinvertebrate community compositions. Furthermore, combining sediment and macroinvertebrate information produced a 27% increase in correlation (ρ_s = 0.71) with shorebird assemblage patterns over the correlation of the bird community with the macroinvertebrate community alone. Beyond its indirect effects acting through prey distributions, granulometry of the flats influenced shorebird foraging directly by modifying prey availability. Our study highlights the importance of habitat heterogeneity, showing that no single patch type was ideal for the entire shorebird community. Generally, shorebird density and diversity were greatest at lower elevations on flats when they became exposed; these areas are at risk from human intervention by inlet sand mining, construction of groins and jetties that divert sediments from flats, and installation of seawalls on inlet shorelines that induce erosion of flats.     

Impact of intercropping aphid-resistant wheat cultivars withoilseed rape on wheat aphid (Sitobion avenae) and its natural enemies

The effects of intercropping of wheat cultivars and oilseed rape on the densities of wheat aphid, Sitobion avenae, and their arthropod natural enemies were evaluated. Three winter wheat cultivars with different resistant levels to S. avenae were used: ‘KOK’ (high resistance), ‘Xiaobaidongmai’ (low resistance) and ‘Hongmanghong’ (susceptible). The results showed that the densities of S. avenae were significantly higher on the monoculture pattern than on either the 8-2 intercropping pattern (eight rows of wheat with two rows of oilseed rape) or the 8-4 intercropping pattern (eight rows of wheat with four rows of oilseed rape). The mean number of predators and the mummy rates of S. avenae were significantly higher in two intercropping patterns than those in the monoculture pattern. The densities of S. avenae, ladybeetles, and mummy rate of S. avenae were significantly different among different wheat cultivars. The highest densities of S. avenae and ladybeetles were found on wheat cultivar Hongmanghong. The lowest densities of S. avenae associated with high mummy rate of S. avenae were found on wheat cultivar Xiaobaidongmai. The results showed that wheat-oilseed rape intercropping conserved more predators and parasitoids than in wheat monoculture fields, and partial resistance of wheat cultivar Xiaobaidongmai had complementary or even synergistic effects on parasitoid of S. avenae.     

Diatoms and dinoflagellates in Terra Nova Bay (Ross Sea-Antarctica) during austral summer 1990

During January/February 1990 the floristic composition and biomass of diatoms and dinoflagellates in Terra Nova Bay (Ross Sea) were investigated. The study area included eight stations, seven of which were inshore and one offshore. Of the 94 taxa identified, 58 were diatoms and 34 were dinoflagellates. Cell densities were higher in surface layers and particularly at two stations, one located at the centre of the bay, and the other to the south of the Terra Nova station. While density data showed that the diatoms predominated over the dinoflagellates and over Phaeocystis sp., the biomass values were largely made up of the dinoflagellates contribution.     

Effect of the snail Ilyanassa obsoleta (Say) on dynamics of the amphipod Corophium volutator (Pallas) on an intertidal mudflat

We investigated interactions between two dominant invertebrate species of intertidal soft-sediment environments of the northwest Atlantic, the mud snail Ilyanassa obsoleta and the burrowing amphipod Corophium volutator, on a mudflat of the upper Bay of Fundy, Canada. Distribution of I. obsoleta on the mudflat was highly patchy and negatively correlated with density of C. volutator. Manipulation of snail density in cages showed that I. obsoleta influences C. volutator; specifically, increasing density of snails reduced density, increased patchiness in distribution, decreased recapture rates and decreased immigration of C. volutator. Ilyanassa obsoleta seems to be affecting C. volutator through an influence on survival rate and emigration rate, although temporal variation in these effects was observed. Given that both I. obsoleta and C. volutator show a preference for tide pools, an important microhabitat on mudflats, snails might have a profound impact on C. volutator population dynamics.     

Diatoms from surface sediments of Amurskiy Bay,Sea of Japan

The results of a study of diatoms from surface sediments (0–1 cm) of Amurskiy Bay are presented for the first time. The specific composition (221 species and intraspecific taxa) and ecological structure of the diatom flora were determined. The diatom species composition of phytoplankton, periphyton, and surface sediments is compared.     

Microalgae biodiversity and biomass status in Qua Iboe Estuary mangrove swamp,Nigeria

Microalgae composition, abundance, diversity and biomass of the Qua Iboe Estuary mangrove swamp were studied. The results revealed the rich assemblage of the brackish ecosystem. Six major taxanomic classes were encountered. These were the Bacillariophyceae, Cyanophyceae, Chlorophyceae, Chrysophyceae, Euglenophyceae and Phaeophyceae. Their composition, abundance and diversity exhibits strong seasonal variation. Variations between pelagic and sedimentary habitats were also noticed. The diatoms (Bacillariophyceae) dominated the habitats. Actinoptychus undalatus, Navicula radiosa and Amphora ovalis co-dominated the pelagic water column; while the epipellic (intertidal) and benthic (subtidal) sediments were co-dominated by A. ovalis and Actinoptychus undulatus. Analyses of the Shannon’s index of general diversity (H¹), McArthur-Terborgh species equitability index (E) and Simpson’s index of dominance (D) of the microalgae communities revealed that A. ovalis with a mean H ¹ -value of 0.25, E-value of 0.08 and D-value of 0.05 was the most prevalent genus in the mangrove ecosystem despite its uneven distribution. The densities of the microalgae communities corresponded with their biomass statuses and were seriously impacted by oil spillage. This forms the basis of concern because the estuary is associated with a high probability of major oil pollution with serious consequences for ecological stability and fisheries.     

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.     

Associations between macrobenthos and invasive cordgrass, Spartina anglica, in the Danish Wadden Sea

We conducted a survey to provide knowledge on mechanisms controlling spatial and temporal variability of macrobenthos in an intertidal Wadden Sea area partly covered by invasive Spartina anglica. Benthic macrofauna was collected seasonally at seven stations along a transect covering non-vegetated mudflat and vegetated marsh areas. Shannon diversity index was consistently higher for macrobenthos in the open mudflat compared to the marsh area. Infaunal species, like Arenicola marina, Tubificoides benedeni and Macoma balthica, were more abundant in the mudflat than the marsh, while the opposite was evident for epifaunal species, like Hydrobia ulvae and Littorina littorea. The infaunal crustacean, Corophium volutator, on the other hand, appeared particularly attracted to the mudflat–marsh boundary. The biomass of below-ground plant materials and macrodetritus was positively correlated with the total macrofaunal abundance, while the biomass of dead below-ground plant materials alone showed a significant negative correlation with infaunal abundance. Total macrofaunal abundance was highest during summer in the mudflat, while no seasonal pattern was evident in the marsh. The negative influence of S. anglica invasions on infaunal abundance may have serious implications for higher trophic levels, such as waterbirds, that forage in Wadden Sea areas and thus for the overall biodiversity and ecosystem functioning.     

Genotype-Dependent morphogenetic potentiality of various explants of a food legume,the pigeon pea (Cajanus cajan L.)

Summary The morphogenetic response of various explants of seven different cultivars of a food legume, the pigeon pea (Cajanus cajan L.), has been studied. The stimulation and elongation of shoot buds into shoots derived from the mature embryo axis and intact seed on Murashige and Skoog&#x2019;s medium supplemented with 2.32&#x00B5;M kinetin and 22.2&#x00B5;M benzyladenine was found to be optimum in Murashige and Skoog&#x2019;s medium supplemented with 0.46&#x00B5;M kinetin, 0.53&#x00B5;M naphthalene acetic acid, and 0.29&#x00B5;M gibberellic acid. Even though the response of these two explants for formation of shoot buds in all the genotypes is 30&#x2013;100% depending on media composition, subsequent growth and elongation of these shoot buds into plants is genotype dependent and is restricted to two genotypes. Cotyledon and epicotyl explants of pigeon pea cultivars on the other hand differentiated directly into four to eight and two to four shoots, respectively, depending on the media composition and genotype. In vitro rhizogenesis of regenerated shoots was 80% and the survival of these plantlets in the field was 70&#x2013;80%. NCL Communication no.: 5667.