A novel technique for detection of the toxic dinoflagellate, Karenia brevis, in the Gulf of Mexico from remotely sensed ocean color data

Karenia brevis, a toxic dinoflagellate that blooms regularly in the Gulf of Mexico, frequently causes widespread ecological and economic damage and can pose a serious threat to human health. A means for detecting blooms early and monitoring existing blooms that offers high spatial and temporal resolution is desired. Between 1999 and 2001, a large bio-optical data set consisting of spectral measurements of remote-sensing reflectance (R_rs(λ)), absorption (a(λ)), and backscattering (b_b(λ)) along with chlorophyll a concentrations and K. brevis cell counts was collected on the central west Florida shelf (WFS) as part of the Ecology and Oceanography of Harmful Algal Blooms (ECOHAB) and Hyperspectral Coastal Ocean Dynamics Experiment (HyCODE) programs. Reflectance model simulations indicate that absorption due to cellular pigmentation is not responsible for the factor of ∼3–4 decrease observed in R_rs(λ) for waters containing greater than 10⁴ cells l⁻¹ of K. brevis. Instead, particulate backscattering is responsible for this decreased reflectivity. Measured particulate backscattering coefficients were significantly lower when K. brevis concentrations exceeded 10⁴ cells l⁻¹ compared to values measured in high-chlorophyll (>1.5 mg m⁻³), diatom-dominated waters containing fewer than 10⁴ cells l⁻¹ of K. brevis. A classification technique for detecting high-chlorophyll, low-backscattering K. brevis blooms is developed. In addition, a method for quantifying chlorophyll concentrations in positively flagged pixels using fluorescence line height (FLH) data obtained from the Moderate Resolution Imaging Spectroradiometer (MODIS) is introduced. Both techniques are successfully applied to Sea-viewing Wide Field-of-view Sensor (SeaWiFS) and MODIS data acquired in late August 2001 and validated using in situ K. brevis cell concentrations.     

The growth dynamics of Karenia brevis within discrete blooms on the West Florida Shelf

As part of the ECOHAB: Florida Program, we studied three large blooms of the harmful bloom forming dinoflagellate Karenia brevis. These blooms formed on the West Florida Shelf during Fall of 2000 off Panama City, and during Fall 2001 and Fall 2002 off the coastline between Tampa Bay and Charlotte Harbor. We suggest that these blooms represent two different stages of development, with the 2000 and 2001 blooms in an active growth or maintenance phase and the 2002 bloom in the early bloom initiation phase. Each bloom was highly productive with vertically integrated primary production values of 0.47–0.61, 0.39–1.33 and 0.65 g C m⁻² d⁻¹ for the 2000, 2001 and 2002 K. brevis blooms, respectively. Carbon specific growth rates were low during each of these blooms with values remaining fairly uniform with depth corresponding to generation times of 3–5 days. Nitrogen assimilation by K. brevis was highest during 2001 with values ranging from 0.15 to 2.14 μmol N L⁻¹ d⁻¹ and lower generally for 2000 and 2002 (0.01–0.64 and 0.66–0.76 μmol N L⁻¹ d⁻¹ for 2000 and 2002, respectively). The highest K. brevis cell densities occurred during the 2001 bloom and ranged from 400 to 800 cells mL⁻¹. Cell densities were lower for each of the 2000 and 2002 blooms relative to those for 2001 with densities ranging from 100 to 500 cells mL⁻¹. The 2000 and 2001 blooms were dominated by K. brevis in terms of its contribution to the total chlorophyll a (chl a) pool with K. brevis accounting generally for >70% of the observed chl a. For those populations that were dominated by K. brevis (e.g. 2000 and 2001), phytoplankton C biomass (C_p,0) constituted <30% of the total particulate organic carbon (POC). However, in 2002 when diatoms and K. brevis each contributed about the same to the total chl a, C_p,0 was >72% of the POC. The fraction of the total chl a that could be attributed to K. brevis was most highly correlated with POC, chl a and salinity. Nitrogen assimilation rate and primary production were highly correlated with a greater correlation coefficient than all other comparisons.     

Nutrient availability in support of Karenia brevis blooms on the central West Florida Shelf: What keeps Karenia blooming?

Identifying nutrient sources, primarily nitrogen (N) and phosphorus (P), sufficient to support high biomass blooms of the red tide dinoflagellate, Karenia brevis, has remained problematic. The West Florida Shelf is oligotrophic, yet populations >10⁶ cells L⁻¹ frequently occur and blooms can persist for months. Here we examine the magnitude and variety of sources for N and P that are available to support blooms. Annual average in situ or background concentrations of inorganic N in the region where blooms occur range 0.02–0.2 μM while inorganic P ranges 0.025–0.24 μM. Such concentrations would be sufficient to support the growth of populations up to ∼3×10⁴ cells L⁻¹ with at least a 1 d turnover rate. Organic N concentrations average 1–2 orders of magnitude greater than inorganic N, 8–14 μM while organic P concentrations average 0.2–0.5 μM. Concentrations of organic N are sufficient to support blooms >10⁵ cells L⁻¹ but the extent to which this complex mixture of N species is utilizable is unknown. Other sources of nutrients included in our analysis are aerial deposition, estuarine flux, benthic flux, zooplankton excretion, N₂-fixation, and subsequent release of organic and inorganic N by Trichodesmium spp., and release of N and P from dead and decaying fish killed by the blooms. Inputs based on atmospheric deposition, benthic flux, and N₂-fixation, were minor contributors to the flux required to support growth of populations >2.6×10⁴ cells L⁻¹. N and P from decaying fish could theoretically maintain populations at moderate concentrations but insufficient data on the flux and subsequent mixing rates does not allow us to calculate average values. Zooplankton excretion rates, based on measured zooplankton population estimates and excretion rates could also supply all of the N and P required to support populations of 10⁵ and 10⁶ cells L⁻¹, respectively, but excretion is considered as “regenerated” nutrient input and can only maintain biomass rather than contribute to “new” biomass. The combined estuarine flux from Tampa Bay, Charlotte Harbor, and the Caloosahatchee River can supply a varying, but at times significant level of N and P to meet growth and photosynthesis requirements for populations of approximately 10⁵ cells L⁻¹ or below. Estimates of remineralization of dead fish could supply a significant proportion of bloom maintenance requirements but the rate of supply must still be determined. Overall, a combination of sources is required to maintain populations >10⁶ cells L⁻¹.     

Cell cycle behavior of laboratory and field populations of the Florida red tide dinoflagellate, Karenia brevis

As a component of the ECOHAB Florida Regional Field Program, this study addresses cell cycle behavior and its importance to bloom formation of the Florida red tide dinoflagellate, Karenia brevis. The cell cycle of K. brevis was first studied by flow cytometry in laboratory batch cultures, and a laboratory mesocosm column, followed by field populations over the 5-year course of the ECOHAB program. Under all conditions studied, K. brevis displayed diel phased cell division with S-phase beginning a minimum of 6 h after the onset of light and continuing for 12–14 h. Mitosis occurred during the dark, and was generally completed by the start of the next day. The timing of cell cycle phases relative to the diel cycle did not differ substantially in bloom populations displaying radically different growth rates (μ_min 0.17–0.55) under different day lengths and temperature conditions. The rhythm of cell cycle progression is independent from the rhythm controlling vertical migration, as similar cell cycle distributions are found at all depths of the water column in field samples. The implications of these findings are discussed in light of our current understanding of the dinoflagellate cell cycle and the development of improved models for K. brevis bloom growth.     

Relationships among water column toxins,cell abundance and chlorophyll concentrations during Karenia brevis blooms

The assumptions that Karenia brevis cell abundance and brevetoxin concentrations are proportional and that cell abundance and chlorophyll are related were tested in a 3-year field study off the west coast of Florida. The relationship between K. brevis cell abundance and brevetoxins (PbTx-2+PbTx-3) in whole water samples was strong (R²=0.92). There was no significant difference between the brevetoxin concentrations in whole water and the >0.7 μm particulate fraction. Only 7% of the total brevetoxin concentration was measured in the <0.7 μm (cell free) filtrate. The relationship of K. brevis cell abundance >5000 cells L⁻¹ with chlorophyll for all cruises and at all depths was robust (R²=0.78). These data substantiate the use of chlorophyll as a proxy for K. brevis cell abundance and K. brevis cell abundance as a proxy for brevetoxins during blooms. The ratios of the brevetoxins, PbTx-2:PbTx-3, was significantly higher in surface water than in bottom water. This information in conjunction with K. brevis growth rates may provide a useful indicator for determining the physiological state of the bloom over time.     

Bacterioplankton dynamics along the gradient from highly eutrophic Pearl River Estuary to oligotrophic northern South China Sea in wet season: implication for anthropogenic inputs

Bacterioplankton abundance (BA) and biomass (BB) from the eutrophic Pearl River Estuary (PRE) to the oligotrophic northern South China Sea (NSCS) were studied in the wet season. BA was significantly higher (p < 0.05) in PRE (12.51 ± 3.52 × 10⁸ cells L⁻¹), than in the continental shelf neritic province (CSNP, 4.95 ± 2.21 × 10⁸ cells L⁻¹) and in the deep oceanic province (OP, 3.16 ± 1.56 × 10⁸ cells L⁻¹). Nutrient-replete PRE waters (DIN > 100 μM and PO₄ > 1 μM) resulted in high chl a and BB, whereas nutrient-depleted offshore waters (DIN <5 μM and PO₄ < 0.5 μM) had low biomass. Temperature (>26 °C) was not the controlling factor of BA. BB was significantly correlated with chl a biomass both in PRE and NSCS. The bacteria to phytoplankton biomass (BB/PB) ratio increased clearly along the gradient from near-shore PRE (0.15) to offshore CSNP (0.93) and deep OP (2.75), indicating the important role of small cells in the open ocean compared to estuarine and coastal zones.     

Ostreopsis cf. ovata bloom in the northern Adriatic Sea during summer 2009: ecology, molecular characterization and toxin profile

Intense blooms of the benthic dinoflagellate Ostreopsis cf. ovata have occurred in the northern Adriatic Sea since 2006. These blooms are associated with noxious effects on human health and with the mortality of benthic organisms because of the production of palytoxin-like compounds. The O. cf. ovata bloom and its relationships with nutrient concentrations at two stations on the Conero Riviera (northern Adriatic Sea) were investigated in the summer of 2009. O. cf. ovata developed from August to November, with the highest abundances in September (1.3 × 10⁶ cells g⁻¹ fw corresponding to 63.8 × 10³ cells cm⁻²). The presence of the single O. cf. ovata genotype was confirmed by a PCR assay. Bloom developed when the seawater temperature was decreasing. Nutrient concentrations did not seem to affect bloom dynamics. Toxin analysis performed by high resolution liquid chromatography-mass spectrometry revealed a high total toxin content (up to 75 pg cell⁻¹), including putative palytoxin and all the ovatoxins known so far.     

On the remote monitoring of Karenia brevis blooms of the west Florida shelf

In situ surveys (1997–2002) of Karenia brevis distribution on the west Florida shelf were used to explain spectral remote sensing reflectance, chlorophyll-a concentration, and backscattering coefficient estimates derived using SeaWiFS satellite data. Two existing approaches were tested in an attempt to differentiate K. brevis blooms from other blooms or plumes. A chlorophyll-anomaly method used operationally by the National Oceanic and Atmospheric Administration (NOAA) sometimes correctly identified K. brevis blooms but also generated false positives and false negatives. The method identified approximately 1000 km² of high chlorophyll-anomalies (>1 mg m⁻³) off southwest Florida between the 10 and 50-m isobaths nearly every day from summer to late fall. Whether these patches were K. brevis blooms or not is unknown. A second method used a backscattering:chlorophyll-a ratio to identify K. brevis patches. This method separated K. brevis from other blooms using in situ optical data, but it yielded less satisfactory results with SeaWiFS data. Spectral reflectance (R_rs) estimates for K. brevis blooms, diatom blooms, and coastal river plumes are statistically similar for many cases. Large pixel size, shallow water, and imperfect algorithms distort satellite retrievals of bio-optical parameters in patchy blooms. At present, a combination of chlorophyll-a, chlorophyll-anomaly, backscattering:chlorophyll-a ratio, RGB composites, MODIS fluorescence data, as well as time-series analysis and ancillary data such as winds, currents, and sea surface temperature can improve K. brevis bloom assessments. Progress in atmospheric correction and bio-optical inversion algorithms is required to help improve capabilities to monitor K. brevis blooms from space. Further, satellite sensors with improved radiometric capabilities and temporal/spatial resolutions are also required.     

Plankton community as an indicator of water quality in tropical shrimp culture ponds

The plankton was examined as an indicator of water quality in 14 shrimp Litopenaeus vannamei farms in Brazil in 2003. The ponds were categorized by high stocking density (>30 PL m⁻²) of phytoplankton, consisting of 51 species with concentrations ranging from 365,218 ± 416,615 cells mL⁻¹ to 1,961,675 ± 3,160,172 cells mL⁻¹. Diatoms contributed to almost 70% of the species number and high densities resulted from Cyanophyta blooms, mainly Pseudanabaena cf limnetica. Forty zooplankton taxa were registered and were essentially composed of typical marine euryhaline species and suspension-feeders. Copepoda dominated (45%) the make-up, followed by Protozoa (18%), Rotifera (12%), and Mollusca (12%) larvae. Zooplankton varied from 972 ± 209 ind m⁻³ to 4235 ± 2877 ind m⁻³. Enhanced nutrient input affected plankton density and composition. Diatom and Copepoda dominance was replaced by cyanobacteria, protozoan, and rotifers as nutrient concentrations increased with the cultured period, indicating that plankton structure is affected by eutrophic conditions.     

Effects of lanthanum(III) and EDTA on the growth and competition of Microcystis aeruginosa and Scenedesmus quadricauda

Rare earth elements (REEs) are widely used to increase crop production in China. However, little attention has been paid to their impacts on aquatic ecology. Batch cultivation was used here to study the effects of lanthanum (La) and EDTA on the growth and competition of the cyanobacterium Microcystis aeruginosa and the green alga Scenedesmus quadricauda. When EDTA was present at a very low concentration (0.269 μmol L⁻¹), low lanthanum concentrations (?7.2 μmol L⁻¹) had little stimulative effect on the growth of M. aeruginosa and S. quadricauda, whereas a high lanthanum concentration (72 μmol L⁻¹) had significant inhibitory effect on both of them. The results of cultivation experiments suggested that the inhibitory effect on M. aeruginosa was higher than that on S. quadricauda and S. quadricauda could become dominant in mixed cultures. When lanthanum was not added to the culture medium, high EDTA concentrations (>13.4 μmol L⁻¹) had a great inhibitory effect on the growth of M. aeruginosa but little effect on the growth of S. quadricauda, which could become dominant in the mixed cultures.Lanthanum and EDTA had complex effects on the growth and competition of M. aeruginosa and S. quadricauda. EDTA did not change the stimulation of low lanthanum concentrations on both, but at intermediated concentrations (2.69-13.4 μmol L⁻¹) it could greatly alleviate lanthanum inhibition on M. aeruginosa; thus, M. aeruginosa would dominate S. quadricauda in these mixed cultures. Lanthanum at low concentration (7.2 μmol L⁻¹) could also alleviate the inhibition of high EDTA on M. aeruginosa, but did not alter the outcome of the competition.     

Bacterial distribution and nutrient limitation in relation to different water masses in the coastal and northwestern South China Sea in late summer

In order to study heterotrophic bacterial responses to upwelling in the northern South China Sea (SCS) and the influence of the Pearl River estuarine coastal plume, two cruises were conducted to investigate the distribution of bacterial abundance (BA) in September-October 2004 and 2005, along with measurements of inorganic nutrients, particulate and dissolved organic carbon (POC and DOC) in 2004. Surface BA was 10±2×10⁸ cells l⁻¹ near the Pearl River estuary and 6±1×10⁸ cells l⁻¹ in oligotrophic offshore waters of the SCS in both 2004 and 2005. In contrast, BA was 15±3×10⁸ cells l⁻¹ in western coastal waters during the upwelling period in 2004, and decreased to 10±2×10⁸ cells l⁻¹ in 2005 when upwelling was absent, indicating that upwelling exerted a significant influence on BA (p<0.05). Nutrient addition experiments were conducted and showed that phosphorus availability limited bacterial growth in coastal upwelled waters and near the Pearl River estuary, while bacteria in offshore waters were mainly C limited. The upwelled waters brought up considerable amounts of nutrients to the surface (e.g. DOC ∼70 μM, DIN ∼4 μM and PO₄ ∼0.1 μM). However, P addition increased BA and bacterial production (BP) by 20±5% and 30±5%, respectively, in the upwelled water, which was higher than those near the Pearl River estuary (2±1% and 20±3%, respectively) (p<0.05). In the upwelled waters, phosphorus was low relative to nitrogen, which resulted in a high N:P ratio of 40:1 at the surface and hence potential P deficiency in bacteria. Consequently, there was a higher increase in BP in response to a PO₄ addition.     

First record of Prorocentrum lima (Dinophyceae) inside harbor areas and along the Abruzzo region coast, W Adriatic

Prorocentrum lima (Ehrenberg) Dodge has been found for the first time during the summer of 2007 inside Ortona harbor and along the coast of the Abruzzo region, a slightly eutrophic area influenced by runoff from a nearby river. The investigations were conducted in two harbors and at six coastal sampling stations. Samplings were conducted using a phytoplankton net and with a pump. Average P. lima cellular concentrations were 3.2 × 10⁴ cells L⁻¹. Other well-known toxic and potentially toxic phytoplankton species have been considered. The number of toxic cells from net samples were higher than the numbers of toxic cell samples collected without the net. Occurrences of P. lima with abiotic factors revealed that temperature was positively correlated with P. lima abundance (p = 0.01), while salinity was highly negatively correlated with P. lima presence (p = 0.001). The total phytoplankton community was studied.     

Influences of temperature and nutrients on Synechococcus abundance and biomass in the southern Mid-Atlantic Bight

Synechococci are small (<1 μm) coccoid prokaryotes that play a significant ecological role in microbial food webs and are important contributors to carbon and nitrogen biogeochemical cycles. Under funding from NOAA and NASA, we developed a time series observatory to understand the seasonal variability of Synechococcus and other phytoplankton. Our goal is to understand the distribution and relative contribution of Synechococcus to the carbon cycle and how they relate to nutrients and temperature. Synechococcus in the southern Mid-Atlantic Bight exhibited a clear seasonal abundance pattern in both inshore and offshore waters—peaking in abundance (11×10⁴ cells ml⁻¹) during warm periods of summer. Synechococci were numerically important during periods of stratification when waters were warm and macronutrients were low. Using a simple algorithm to convert cellular volume to cellular carbon using image analysis, we estimated that Synechococcus cellular carbon ranged from 0.1 to 1.5 pg C per cell and was most significant compared to total particulate carbon in the summer peaking at ∼25% of the total carbon available. No direct correlations were found between Synechococcus abundance and nitrate, nitrite, ammonium, phosphate, and silicate. However, inshore Synechococcus abundance peaked at 10⁴ cells ml⁻¹ when nitrogen concentrations were lowest. Our results suggest that Synechococcus is adapted to warm temperatures and are capable of demonstrating rapid growth during summer when macronutrients are limiting. The ability of Synechococcus to take advantage of high summer temperatures, low nutrient concentrations and low light levels allows them to maintain a picoplankton community during periods of low detritus and nanophytoplankton is nutrient limited. Temperature-dependence is important in altering the size spectrum of the phytoplankton community and affects the carbon cycle on the Mid Atlantic Bight.     

Harmful algal bloom species and phosphate-processing effluent: field and laboratory studies

In 2002, the Florida Department of Environmental Protection began discharging phosphate-processing effluent into Bishop Harbor, an estuary within Tampa Bay. Because of concerns that the effluent would serve as a nutrient source for blooms of the toxic dinoflagellate Karenia brevis, a field monitoring program was established and laboratory bioassays were conducted. Several harmful algal bloom (HAB) species, including Prorocentrum minimum and Heterosigma akashiwo, were observed in bloom concentrations adjacent to the effluent discharge site. Blooms of diatoms were widespread throughout Bishop Harbor. K. brevis was observed with cell concentrations decreasing with increasing proximity to the effluent discharge site. Bioassays using effluent as a nutrient source for K. brevis resulted in decreased cell yields, increased growth rates, and increased time to log-phase growth. The responses of HAB species within Bishop Harbor and of K. brevis to effluent in bioassays suggested that HAB species differ in their response to phosphate-processing effluent.     

Optimization of nutrient component for diesel oil degradation by Rhodococcus erythropolis

A novel bacterium T7-2 was isolated from the oil-polluted sea-bed mud of Bohai Sea, northern China, which can degrade diesel oil at 15 °C. This bacterium was identified as a strain of Rhodococcus erythropolis according to its 16S rDNA gene. In order to enhance degradation efficiency, a five-level, three-factor central composite design was employed to optimize the nutrition supplied to artificial seawater. The results indicate that a supplement of 2.53 g (NH₄)₂SO₄ L⁻¹, 2.75 g Na₂HPO₄ L⁻¹ and 0.01 g yeast extract L⁻¹ to artificial seawater increases the degradation rate from 12.61% to 75% within 7 d.     

Hydrodynamic accumulation of Karenia off the west coast of Florida

Blooms of the toxic dinoflagellates, Karenia spp. occur nearly annually in the eastern Gulf of Mexico with cell abundances typically >10⁵ cells L⁻¹. Thermal and ocean color satellite imagery shows sea surface temperature patterns indicative of upwelling events and the concentration of chlorophyll at fronts along the west Florida continental shelf. Daily cell counts of Karenia show greater increases in cell concentrations at fronts than can be explained by Karenia's maximum specific growth rate. This is observed in satellite images as up to a 10-fold greater increase in chlorophyll biomass over 1–2 d periods than can be explained by in situ growth. In this study, we propose a model that explains why surface blooms of Karenia may develop even when nutrients on the west Florida shelf are low. In the summer, northward winds produce a net flow east and southeast bringing water and nutrients from the Mississippi River plume onto the west Florida shelf at depths of 20–50 m. This water mass supplies utilizable inorganic and organic forms of nitrogen that promote the growth of Karenia to pre-bloom concentrations in sub-surface waters in the mid-shelf region. In the fall, a change to upwelling favorable winds produces onshore transport. This transport, coupled with the swimming behavior of Karenia, leads to physical accumulation at frontal regions near the coast, resulting in fall blooms. Strong thermal fronts during the winter provide a mechanism for re-intensification of the blooms, if Karenia cells are located north of the fronts. This conceptual model leads to testable hypotheses on bloom development throughout the Gulf of Mexico.     

Long-term differences in annual litter production between alien (Sonneratia apetala) and native (Kandelia obovata) mangrove species in Futian,Shenzhen, China

Annual litter production in alien (Sonneratia apetala) and native (Kandelia obovata) mangrove forests in Shenzhen, China were compared from 1999 to 2010. S. apetala had significantly higher litter production than K. obovata, with mean annual total litter of 18.1 t ha⁻¹ yr⁻¹ and 15.2 t ha⁻¹ yr⁻¹, respectively. The higher litter production in S. apetala forest indicates higher productivity and consequently more nutrient supply to the estuarine ecosystems but may be more invasive due to positive plant-soil feedbacks and nutrient availability to this alien species. Two peaks were recorded in S. apetala (May and October), while only one peak was observed in K. obovata, in early spring (March and April). Leaf and reproductive materials were the main contributors to litter production (>80%) in both forests. These results suggest that the ecological function of S. apetala and its invasive potential can be better understood based on a long-term litter fall analysis.     

Thorium(IV) removal from aqueous medium by citric acid treated mangrove endophytic fungus Fusarium sp. #ZZF51

Thorium(IV) biosorption is investigated by citric acid treated mangrove endophytic fungus Fussarium sp. #ZZF51 (CA-ZZF51) from South China Sea. The biosorption process was optimized at pH 4.5, equilibrium time 90 min, initial thorium(IV) concentration 50 mg L⁻¹ and adsorbent dose 0.6 g L⁻¹ with 90.87% of removal efficiency and 75.47 mg g⁻¹ of adsorption capacity, which is obviously greater than that (11.35 mg g⁻¹) of the untreated fungus Fussarium sp. #ZZF51 for thorium(IV) biosorption under the condition of optimization. The experimental data are analyzed by using isotherm and kinetic models. Kinetic data follow the pseudo-second-order model and equilibrium data agree very well with the Langmuir model. In addition, FTIR analysis indicates that hydroxyl, amino, and carbonyl groups act as the important roles in the adsorption process.     

The new potential invader Linopherus canariensis (Polychaeta: Amphinomidae) in a Mediterranean coastal lake: colonization dynamics and morphological remarks

The newly introduced polychaete Linopherus canariensis Langerhans, 1881 was found in the Lake of Faro (NE Sicily), during a study comparing the macrobenthos in artificial modules with a neighboring sandy bottom assemblage. Of a total of 4465 specimens, almost 6% showed morphological variation related to branchial turfs and mean body size. The sandy bottom exhibited an average density of 41.86 ind L⁻¹ and a wet biomass of 30.35 mg L⁻¹, whereas the artificial substratum reached levels of 205.29 ind L⁻¹ and 318.44 mg L⁻¹. The highest estimated immigration rate was 3.7 ind L⁻¹ d⁻¹ (5.8 mg L⁻¹ d⁻¹). In the artificial microhabitat, 0.4% of the population showed mid-anterior fragmentation, with anterior- (2%), mid- (<1%) and posterior- (1%) regenerating lineages, which contributed significantly to the dispersion ability of this species. L. canariensis was a selective micro-deposit feeder, even under conditions of reduced sediments. Linopherus was found to be a new potential invader of stressed environments that is probably tied to the import of oysters.     

Community composition, distribution, and contribution of microbenthos in offshore sediments from the Yellow Sea

We investigated the spatial distribution and composition of microbenthos in the seafloor sediments from 48 stations in the Yellow Sea using epifluorescence microscopy and quantitative protargol staining techniques. The bacterial abundance ranged from 2.4×10⁸ to 1.9×10⁹ cells cm⁻³ in the wet sediment, about three orders of magnitude higher than that of phototrophic (PNFs, from 6.4×10⁵ to 8.8×10⁶ cells cm⁻³) and heterotrophic nanoflagellates (HNFs, from 5.8×10⁴ to 5.9×10⁶ cells cm⁻³) and four orders of magnitude higher than that of cyanobacteria (from 2.3×10⁴ to 2.3×10⁶ cells cm⁻³) in the upper 5 cm of sediments. The abundance of diatoms varied greatly, from 3-1.1×10⁵ cells cm⁻³ in the upper 8 cm of sediments, whereas those of heterotrophic microflagellates (HMFs, 1-182 cells cm⁻³) and ciliates (1-221 cells cm⁻³) were less varied and lower. The biomass partitioning indicates the primary importance of benthic bacteria (50.3 μg C cm⁻³ on average), followed by PNFs (40.7 μg C cm⁻³), HNFs (19.3 μg C cm⁻³), and finally by cyanobacteria (8.8 μg C cm⁻³). Benthic diatoms (0.8 μg C cm⁻³), ciliates (0.15 μg C cm⁻³), and HMFs (0.03 μg C cm⁻³) contribute relatively small fractions to the total biomass of the microbenthos. About 95% of diatoms, 77% of ciliates, and 56% of HMFs were distributed in the upper 2 cm of sediments, whereas no distinct vertical distributions were observed for bacteria, cyanobacteria, PNFs, and HNFs. The microbenthos are quantitatively important in the shallow seafloor, wherein their main components have an average abundance three orders of magnitude higher than the corresponding planktonic organisms in the same sea area. Our estimates indicate that pico-sized phytobenthos might contribute a large proportion to the primary production. Benthic ciliates and heterotrophic flagellates contribute about 90% to the estimated combined metabolic rate of micro- and meiobenthic consumers in the whole sea area, with nanoheterotrophs accounting for the majority. The data suggest the potential for the rapid primary and secondary production of microbenthos and detrital utilization in the shallow seafloor sediments of the Yellow Sea.