Effects of macroalgal mats on intertidal sandflats: an experimental study

The growth of green macroalgal mats is becoming increasingly common in many marine intertidal habitats. While the ecological effects of such growth has previously been experimentally investigated on mudflats, such experiments have rarely been performed on intertidal sandflats. This study investigated the ecological effects of macroalgal cover on a moderately exposed intertidal sandflat, Drum Sands, Firth of Forth, Scotland. Artificially implanted Enteromorpha prolifera (Müller) caused marked changes in the macrobenthos, together with significant changes in all the measured sediment variables. After 6 weeks, the weed significantly increased the macrofaunal diversity. The numbers of Pygospio elegans (Claparède) were significantly reduced under weed mats, while those of Capitella capitata (Fabricius), oligochaetes and gammarids increased. Percent water, organics and silt/clay contents, medium phi and sorting coefficient significantly increased in the sediments under weed mats which also became significantly more reduced between 1 and 8 cm depth. After 20 weeks, a macrofaunal community numerically dominated by C. capitata, with a significantly reduced diversity, was present under weed mats, while sediment variables were no longer significantly different from controls. The negative effect of E. prolifera on P. elegans was mainly due to larval filtering, suggesting that weed is likely to have detrimental effects on population maintenance of most species which rely on planktonic larval recruitment. These results are broadly similar to those obtained from algal manipulation experiments performed in much more sheltered, muddier environments. We suggest that a predictable deterioration in environmental quality results from the growth of macroalgal mats in soft-bottom habitats. However, the longer term effects of such algal growth are less predictable and depend upon the spatial distributions of the most abundant infaunal species and the spatial heterogeneity of weed mat establishment.     

Osmotic changes in an estuarine bivalve,Modiolus fluviatilis

• 1.1.Responses to different salinities monitored by opening and closing of the shell valves were observed in Modiolus fluviatilis.
• 2.2.The osmotic pressure, sodium and chloride ion concentrations were measured in the haemocoelic fluid of Modiolus fluviatilis under similar conditions.
• 3.3.Free amino acids (measured as ninhydrin-positive substances) were determined in the muscle tissue of Modiolus.
• 4.4.It appears that these free amino acids are involved in the ability of the estuarine bivalve Modiolus fluviatilis to osmoregulate in a wide range of salinities.

     

Macroinvertebrate response to different species of macroalgal mats and the role of disturbance history

Over the last 20 years, the Mondego estuary, Portugal has experienced excessive growth of macroalgae especially in the inner parts of the system, with several algal species implicated. In this study, we compare the effects of morphologically different species, the red alga Gracilaria verrucosa and the green macroalga Enteromorpha intestinalis on macrobenthic assemblages, by a field experiment whereby the biomass of algae was manipulated and the resultant changes in macrofauna abundance evaluated. The experiments were carried out in different areas (a relatively undisturbed sea grass bed and an upstream eutrophic area) experiencing different degrees of overall enrichment.Measurements of sediment redox potential revealed a rapid anoxia with a significant increase in algal biomass after 4 weeks. The effects of macroalgae were different at the two sites, being more marked in the eutrophic area. In addition, the effects of Gracilaria and Enteromorpha were significantly different, with Enteromorpha having a greater detrimental effect for most of the macrofauna, in particular Cyathura carinata, Scrobicularia plana, Cerastoderma edule and Alkmaria romijni. However, three of the most abundant invertebrates (Hydrobia ulvae, Hediste diversicolor and Capitella capitata) showed significant increases in abundance in weed affected compared to weed-free plots. Gracilaria had less of an impact on macrobenthic assemblages leading to a more enriched community. Between-site differences in overall impact were related to their previous disturbance history.     

Effects of an invasive reef-building polychaete on the biomass and composition of estuarine macroalgal assemblages

Invasive species can transform ecological communities. Their profound effects may alter the sources and pathways of primary production. We investigated the effects of the reef forming polychaete Ficopomatus enigmaticus invasion on the biomass and distribution of estuarine macroalgae in a SW Atlantic coastal lagoon (Mar Chiquita, 37° 40′S, 57° 23′W, Argentina). Reefs built by this species serve as substrates for macroalgal development and furnish structures that modify physical and biological conditions for the surrounding benthos. We showed that (1) the red macroalga Polysiphonia subtilissima settles and grows almost exclusively on the surface of the reef, (2) the green macroalgae Cladophora sp. and Enteromorpha intestinalis are found almost exclusively in areas without reefs attached to mollusk shells and, (3) no macroalgae occur in the sediment between reefs. Manipulative experiments show that reefs provide a complex substrate for settlement and survival and therefore benefit red macroalga. These experiments also show that the invasive reef builder has negative indirect effects on green macroalgae by increasing grazing and probably by increased sedimentation between reefs. Via these direct and indirect effects, reefs change the relative biomass contribution of each macroalgal species to the overall production in the lagoon. Knowledge of these processes is important not only for predicting net effects on primary production but also because changes in macroalgal species composition may produce effects that cascade through the food web.     

The role of bivalve molluscs as tools in estuarine sediment toxicity testing: a review

Estuarine sediments frequently are repositories and therefore potential sources of anthropogenic contaminants. Many organic and metallic chemical compounds released into aquatic systems bind to particulates and so accumulate in the sediments, thus, sediments become repositories of contaminants in estuaries. These may also cause contamination through diffusion of porewater, resuspension of particulates and dispersal of benthic fauna. There is a need to assess the biological affects of these anthropogenic contaminants because they may be toxic to infauna and bottomfish. Sediment toxicity bioassays are a means for carrying out such an assessment and primarily provide data on toxicity by measuring the effects on the test organism. Existing sediment toxicity bioassays rely on a battery of aquatic toxicity tests, which are based on the extraction of pore water, and elutriate from sediments and then subjecting these sediment phases to toxicity testing regimes. Two estuarine bivalve molluscs, Scrobicularia plana and Tapes semidecussatus were used to assess the ecotoxicity of field-collected sediments from estuarine and coastal areas around the Irish and English Coast over a 3-year study period. A variety of endpoints were measured during the study including survival in air, behaviour, animal condition, biochemistry, soft tissue metal concentrations, lysosomal membrane integrity and histopathology. Of these endpoints, the most sensitive were survival, survival in air, lysosomal membrane integrity, behaviour and histopathology.     

Effects of a macroalgal mat (Ulva lactuca) on estuarine sand flat copepods: an experimental study

The effects of a macroalgal mat (Ulva lactuca) on intertidal sand flat copepods were studied in Jamaica Bay, NY. In 1999, Ulva was removed from paired quadrats in the algal mat. Density and species richness were estimated from sediment cores over seven biweekly periods and compared with Ulva-removal quadrats and with an unvegetated reference site. In a separate analysis at the same site, species composition and density of all copepods within the algal mat were determined from replicated quadrat samples over seven dates. Emergence traps were deployed in all treatments in 1999 to quantify copepod movements from sediments into the water column during flooding tide. Dissolved oxygen (DO) was continuously monitored over several 96-h periods in and above the algal mat. In 2000, three biomass levels of Ulva (0, 100, 200 g DW per m²) were added to paired quadrats in the unvegetated zone. Density and species richness were estimated over eight time periods. Ulva removal resulted in significant increases in density and species richness, but levels remained lower than at the unvegetated site. Likewise, addition of Ulva biomass produced significant declines in copepod density. However, the differences between low- and high-Ulva-addition treatments were insignificant. Increases in phytal copepods after Ulva addition did not compensate for losses of epibenthic and infaunal species. The overall effect of the Ulva mat was a net loss of copepod density, including the loss of nearly all infaunal species. Anoxia in sediments and within the algal mat are correlated with declines in infaunal species and with escape by some infaunal species from the sediment into the water column.     

Sedimentary environment influences the effect of an infaunal suspension feeding bivalve on estuarine ecosystem function

The suspension feeding bivalve Austrovenus stutchburyi is a key species on intertidal sandflats in New Zealand, affecting the appearance and functioning of these systems, but is susceptible to several environmental stressors including sedimentation. Previous studies into the effect of this species on ecosystem function have been restricted in space and time, limiting our ability to infer the effect of habitat change on functioning. We examined the effect of Austrovenus on benthic primary production and nutrient dynamics at two sites, one sandy, the other composed of muddy-sand to determine whether sedimentary environment alters this key species' role. At each site we established large (16 m(2)) plots of two types, Austrovenus addition and removal. In winter and summer we deployed light and dark benthic chambers to quantify oxygen and nutrient fluxes and measured sediment denitrification enzyme activity to assess denitrification potential. Rates of gross primary production (GPP) and ammonium uptake were significantly increased when Austrovenus was added, relative to removed, at the sandy site (GPP, 1.5 times greater in winter and summer; ammonium uptake, 8 times greater in summer; 3-factor analysis of variance (ANOVA), p<0.05). Denitrification potential was also elevated in Austrovenus addition plots at the sandy site in summer (by 1.6 times, p<0.1). In contrast, there was no effect of Austrovenus treatment on any of these variables at the muddy-sand site, and overall rates tended to be lower at the muddy-sand site, relative to the sandy site (e.g. GPP was 2.1 to 3.4 times lower in winter and summer, respectively, p<0.001). Our results suggest that the positive effects of Austrovenus on system productivity and denitrification potential is limited at a muddy-sand site compared to a sandy site, and reveal the importance of considering sedimentary environment when examining the effect of key species on ecosystem function.     

A short-term study of the effects of algal mats on the distribution and behavioural ecology of estuarine birds

The effects of algal mats on wintering wading bird distribution and behaviour were studied at two sites during 1998/99 to identify changes that occurred during the winter breakdown of the algal mat. Numbers of Black-tailed Godwit Limosa limosa did not increase as the algal mat reduced in extent but a significantly greater number of birds were observed in areas without algal cover than in areas with mats in most months. The use of clear spaces, created between the algal patches as the mat broke down, by Black-tailed Godwit increased across the winter. We suggest that algal mats may physically interfere with the deep probing action of this wader. In contrast, when algal mats were most dense in November 1998, significantly greater numbers of Redshank Tringa totanus were observed in algal areas at one site. Redshank appeared not to be deterred by algal cover and, at times, preferentially utilized the algal infauna as prey.     

Carbon-13 depletion in an estuarine bivalve: Detection of marine and terrestrial food sources

Summary Stable carbon isotope analysis was used to define the food sources of bivalve Chione (Austrovenus) stutchburyi in the Avon-Heathcote Estuary, Christchurch, New Zealand. ¹³C values of C. stutchburyi tissue were significantly different (from -16.7 to -23.5 relative to the PDB standard) at five locations separated by less than 4 km but subject to different hydrological regimes. This is related to differences in the isotopic composition of the suspended particulate matter of the inflowing water. C. stutchburyi is shown to utilise carbon of terrestrial and marine origin depending upon its position within the estuary and local hydrology.     

The distribution and temporal dynamics of the estuarine macroalgal community of San Francisco Bay

Long-term sampling of intertidal macroalgae along permanently marked transects within San Francisco Bay has shown a marked decline in overall species number along the estuarine gradient from the ocean to the river, presumably as a result of decreasing salinity and a progressive lack of hard substrata in the upstream direction. Green algae penetrated further landward than either brown or red species. Seasonally, macroalgal species diversity is lowest during the winter-spring months when salinity, temperature, and irradiance are at yearly minima. Macroalgal abundance as measured by percent cover was maximum during the late spring near the mouth of the estuary and during late summer towards the head. The seasonal increase in algal abundance is related to increasing salinity, temperature, and light availability to the bottom. The summer increase in irradiance is due to the longer photoperiod, increased frequency of day-time low tides, and reduced levels of suspended sediments. The aperiodic occurrence of algal blooms in San Pablo Bay may be caused by a combination of physical factors which are ultimately associated with the river inflow. A hypothesis based on interannual differences in river inflow and the contribution of phytoplankton to nutrient cycles in the benthos is presented to explain the occurrence of nuisance algal blooms.     

Model simulations of conditions suitable for the establishment of Enteromorpha sp. (Chlorophyta) macroalgal mats

Floating mats of algae regularly appear in shallow bays in many coastal areas around the world. This paper presents a model that simulates the growth of ephemeral macroalgae in shallow bays, and interprets the growth according to physical and biological prerequisites. Model runs emphasized the role of internal nutrient cycling to provide the enhanced nutrient availability needed to ensure abundant growth of macroalgae. Thus, the presence of macroalgae, as well as long water residence times, influenced the nutrient cycling in the bay to support further growth of macroalgal mats.     

The earliest known estuarine bivalve assemblage, Lower Ordovician of northwestern Argentina

The oldest known estuarine bivalve assemblage is documented from the Lower Ordovician (upper Arenig-lower Llanvirn) Alto del Cóndor Formation, which crops out in the Cordillera Oriental of northwestern Argentina. This unit displays most of the diagnostic sedimentary attributes of estuarine environments. Biotic components include low-diversity trace fossils and a peculiar bivalve fauna consisting of the new genera Konduria, Pseudoredonia, and Pucamya, and the new species Redonia condorensis. This constitutes the earliest known occurrence of bivalves in brackish waters, suggesting that the capability of this clade to colonize estuarine environments developed early in their radiation.     

Spatiotemporal changes in distributions of the two dominant bivalve species in an estuarine river connecting two brackish lakes,Japan

Spatiotemporal changes in the biomass and abundance of the two dominant bivalve species Corbicula japonica and Arcuatula senhousia were investigated at 14 stations positioned along the Ohashi River from November 2005 to August 2007. Corbicula japonica predominantly inhabited the upstream section of the primary branch and Kensaki stream of the Ohashi River, whereas A. senhousia predominantly inhabited the downstream section of the Ohashi River. However, the distributions were highly variable among stations and seasons. Intrusions of saline water from Lake Nakaumi varied with astronomical and climatological factors, and consequently, the halocline of the Ohashi River is subject to spatiotemporal changes. Changes in distributions of C. japonica and A. senhousia occurred in relation to a flood event, indicating that environmental disturbances affect the distributions of the two bivalve species. Responses of the two species to continual saline water intrusion were also observed during the period of normal flow conditions. We suggest that the distribution of the two species is closely related to the dynamics of the ambient saline water, which is the most important factor correlated to changes in the distributions of C. japonica and A. senhousia in the Ohashi River.     

High macroalgal cover and low coral recruitment undermines the potential resilience of the world's southernmost coral reef assemblages

Coral reefs are under increasing pressure from anthropogenic and climate-induced stressors. The ability of reefs to reassemble and regenerate after disturbances (i.e., resilience) is largely dependent on the capacity of herbivores to prevent macroalgal expansion, and the replenishment of coral populations through larval recruitment. Currently there is a paucity of this information for higher latitude, subtropical reefs. To assess the potential resilience of the benthic reef assemblages of Lord Howe Island (31°32'S, 159°04'E), the worlds' southernmost coral reef, we quantified the benthic composition, densities of juvenile corals (as a proxy for coral recruitment), and herbivorous fish communities. Despite some variation among habitats and sites, benthic communities were dominated by live scleractinian corals (mean cover 37.4%) and fleshy macroalgae (20.9%). Live coral cover was higher than in most other subtropical reefs and directly comparable to lower latitude tropical reefs. Juvenile coral densities (0.8 ind.m(-2)), however, were 5-200 times lower than those reported for tropical reefs. Overall, macroalgal cover was negatively related to the cover of live coral and the density of juvenile corals, but displayed no relationship with herbivorous fish biomass. The biomass of herbivorous fishes was relatively low (204 kg.ha(-1)), and in marked contrast to tropical reefs was dominated by macroalgal browsing species (84.1%) with relatively few grazing species. Despite their extremely low biomass, grazing fishes were positively related to both the density of juvenile corals and the cover of bare substrata, suggesting that they may enhance the recruitment of corals through the provision of suitable settlement sites. Although Lord Howe Islands' reefs are currently coral-dominated, the high macroalgal cover, coupled with limited coral recruitment and low coral growth rates suggest these reefs may be extremely susceptible to future disturbances.     

The good,the bad and the Ulva: the density dependent role of macroalgal subsidies in influencing diversity and trophic structure of an estuarine community

Worldwide, ecological subsidies enhance ecosystem productivity and therefore trophic support for greater biodiversity of taxa. While studies in terrestrial and aquatic ecosystems demonstrate that the magnitude of subsidies into ecosystems differs widely, the thresholds where subsidies may switch from exerting positive to negative effects are poorly understood. In estuaries, eutrophication promotes drift macroalgae that deposit on the benthos, cover intertidal flats for months and serve as pressed resource subsidies for benthic consumers. We hypothesized there would be a critical threshold of macroalgal biomass where ecosystem‐level effects would turn from positive to negative. We used manipulative field experiments varying macroalgal mat thickness (0.5, 1.5 and 4 cm) over eight weeks and quantified effects on macrofauna on a lagoon mudflat in California. We documented that plots with mat depths of 0.5 and 1.5 cm had higher diversity by supporting both surface feeding and burrowing detritivores. Non‐metric multidimensional scaling showed that the benthic community diverged with mat depth over the course of the experiment. After eight weeks, surface deposit feeders were associated mainly with 0.5 cm macroalgal subsidies, whereas subsurface deposit feeding capitellids were closely linked with 4 cm mats. Depth profiles of pore water sulfide concentration collected from 4 cm mats were 7622 ± 5294 μM, mean ± SE, (mean of means across depth profiles), whereas 0.5 cm treatments resulted in sulfide concentrations that were 0.25% of the 4 cm treatments. This suggests that the mechanism of negative effects for elevated macroalgal subsidies was development of anoxic conditions promoting sulfide accumulation. Thus, our study was the first to find a critical threshold, or ecological tipping point, beyond which the effects of anthropogenically enhanced subsidies to estuarine mudflat communities switched from positive to negative and to describe the mechanism by which elevated subsides altered the abiotic environment and likely reduced ecosystem functioning.     

Hemolymph cells of the bivalve mollusc Mercenaria mercenaria: an electron microscopical study.

The hemolymph cells of Mercenaria mercenaria were studied with the transmission electron microscope. Three morphological types of cells, granulocytes, hyalinocytes, and fibrocytes, are distinguishable and their fine structural characteristics are described. However, as a result of analyzing the fine structural features of the so-called fibrocytes of M. mercenaria, i.e., the inclusion of large aggregates of glycogen granules in their cytoplasm and the occurrence of primary phagosomes enclosing partially degraded exogenous material and digestive lamellae, it is suggested that fibrocytes are actually granulocytes which are at the terminal phase of their physiologic cycle relative to the degradation of phagocytized nonself materials. The cytoplasmic granules of M. mercenaria granulocytes are structurally different from those of Crassostrea virginica in that they are delimited by a unit membrane, rather than by a complex wall, and include a homogenously electron-dense material. Lipidlike droplets are reported from both granulocytes and hyalinocytes of M. mercenaria for the first time.     

A few is enough: a low cover of a non-native seaweed reduces the resilience of Mediterranean macroalgal stands to disturbances of varying extent

There is increasing evidence that the severity of the ecological impact of non-native species does not necessarily scale linearly with their abundance in the introduced range. Nonetheless, the potential of low abundance invaders to alter the resilience of native communities to disturbance has been poorly explored. On Mediterranean rocky reefs, we tested the hypothesis that (1) a pulse disturbance opening gaps within canopy stands formed by the fucoid seaweed Cystoseira brachycarpa facilitates the establishment of the non-native seaweed Caulerpa cylindracea and that (2), once established, the seaweed can reduce the recovery of macroalgal canopies. In July 2011, C. cylindracea was experimentally transplanted in small and large plots that were either cleared of the canopy or left untouched. After 45 months, the cover of C. cylindracea was greatest in small canopy-removal plots, without, however, achieving values exceeding ~10%. Nonetheless, such a low abundance of C. cylindracea caused a threefold reduction in canopy recovery. The establishment of C. cylindracea in canopy-removal plots did not alter the structure of the understory assemblages or the cover of turf-forming, erect and encrusting algae and sessile invertebrates. Our results suggest that some non-native species may be stronger competitors than natives, despite their low abundance. This property has important implications for control programs since not achieving the total eradication of the targeted invader would make little progress towards the mitigation of its impacts. Finally, our results show that non-native species acting as passengers of change can ultimately promote the persistence of alternative degraded states.     

Biotic resistance to invasion along an estuarine gradient

Biotic resistance is the ability of native communities to repel the establishment of invasive species. Predation by native species may confer biotic resistance to communities, but the environmental context under which this form of biotic resistance occurs is not well understood. We evaluated several factors that influence the distribution of invasive Asian mussels (Musculista senhousia) in Mission Bay, a southern California estuary containing an extensive eelgrass (Zostera marina) habitat. Asian mussels exhibit a distinct spatial pattern of invasion, with extremely high densities towards the back of Mission Bay (up to 4,000 m⁻²) in contrast with near-complete absence at sites towards the front of the bay. We established that recruits arrived at sites where adult mussels were absent and found that dense eelgrass does not appear to preclude Asian mussel growth and survival. Mussel survival and growth were high in predator-exclusion plots throughout the bay, but mussel survival was low in the front of the bay when plots were open to predators. Additional experiments revealed that consumption by spiny lobsters (Panulirus interruptus) and a gastropod (Pteropurpura festiva) likely are the primary factors responsible for resistance to Asian mussel invasion. However, biotic resistance was dependent on location within the estuary (for both species) and also on the availability of a hard substratum (for P. festiva). Our findings indicate that biotic resistance in the form of predation may be conferred by higher order predators, but that the strength of resistance may strongly vary across estuarine gradients and depend on the nature of the locally available habitat.     

13种双壳类贝壳的扫描电镜观察

采用扫描电子显微镜(SEM)对湛江地区的丽文蛤(Meretrix lusoria)、琴文蛤(M.lyrata)、文蛤(M.meretrix)、波纹巴非蛤(Paphia undulata)、菲律宾蛤仔(Ruditapes philippinarum)、杂色蛤仔(R.variegata)、伊萨伯雪蛤(Clausinella isobellina)、格粗饰蚶(Anadara clathrata)、泥蚶(Tegillarca granosa)、栉孔扇贝(Chlamys farreri)、尖紫蛤(Sanguinolaria aauta)、锈色朽叶蛤(Coecella turgida)和栉江珧(Atrina pectinata)这6科13种双壳类的贝壳形态微观结构特征进行观察。结果表明,不同种的贝壳表面和横切面微观结构有一定差异。这些差异主要表现在晶体的组成和排列方式两个方面。贝壳角质层根据表面形态特征分为5种类型:光滑平整、颗粒状、不规则多边形、蜂窝状和沟壑状。贝壳棱柱层的晶体形状有棱柱状、短柱状、片状和不规则形状。不同贝壳的晶体有两种排列方向,垂直于横切面和平行于横切面。13种贝壳珍珠层的晶体有颗粒状、砖块状、圆形、块状和不规则的多边形。不同种的贝壳角质层、棱柱层和珍珠层的厚度也不同。研究贝壳微观结构之间的差异,可以为分类提供基本资料。     

Short-term testing of antifouling surfaces: the importance of colour

Data from short-term biofouling assays are frequently used to evaluate the performance of antifouling (AF) coatings. There are a large number of factors, however, that may influence community development. One such factor is colour. The hypothesis was that differences in colour may impact the short-term development of a biofouling community and therefore bias the results. An experiment was designed to investigate the effect of black and white substrata on settlement of fouling organisms in the field. Both Ulva sp. and Spirorbis sp. had significantly higher settlement on black surfaces. This result emphasises the importance of considering colour and other factors when undertaking short-term testing of AF coatings.