Static vs dynamic settlement and adhesion of diatoms to ship hull coatings

Many experiments utilize static immersion tests to evaluate the performance of ship hull coatings. These provide valuable data; however, they do not accurately represent the conditions both the hull and fouling organisms encounter while a ship is underway. This study investigated the effect of static and dynamic immersion on the adhesion and settlement of diatoms to one antifouling coating (BRA 640), four fouling-release coatings (Intersleek^® 700, Intersleek^® 900, Hempasil X3, and Dow Corning 3140) and one standard surface (Intergard^® 240 Epoxy). Differences in community composition were observed between the static and dynamic treatments. Achnanthes longipes was present on all coatings under static immersion, but was not present under dynamic immersion. This was also found for diatoms in the genera Bacillaria and Gyrosigma. Melosira moniformis was the only diatom present under dynamic conditions, but not static conditions. Several common fouling diatom genera were present on panels regardless of treatment: Amphora, Cocconeis, Entomoneis Cylindrotheca, Licmophora, Navicula, Nitzschia, Plagiotropis, and Synedra. Biofilm adhesion, diatom abundance and diatom diversity were found to be significantly different between static and dynamic treatments; however, the difference was dependent on coating and sampling date. Several coatings (Epoxy, DC 3140 and IS 700) had significantly higher biofilm adhesion on dynamically treated panels on at least one of the four sampling dates, while all coatings had significantly higher diatom abundance on at least one sampling date. Diversity was significantly greater on static panels than dynamic panels for Epoxy, IS 700 and HX3 at least once during the sampling period. The results demonstrate how hydrodynamic stress will significantly influence the microfouling community. Dynamic immersion testing is required to fully understand how antifouling surfaces will respond to biofilm formation when subjected to the stresses experienced by a ship underway.     

Settlement, early growth and survival of Haliotis rubra in response to different algal species

Five diatom species were isolated from settlementplates at Southern Ocean Mariculture, Victoria,Australia (Navicula sp., Naviculajeffreyi, Cylindrotheca closterium, Cocconeis sp., Amphora sp.) and tested insettlement experiments with Haliotis rubralarvae. Settlement was very low on single speciesdiatom films and varied between 1%–6%. Depending onthe species combination larvae preferred to settle onfilms with mixed diatom species than single speciesfilms. The highest settlement was achieved with amixed film of Navicula sp. and Amphora sp.Five and ten-day-old germlings of Sporolithondurum induced settlement of the abalone Haliotisrubra. However, the settlement rate was significantlylower on germlings than on the whole thallus of thealga. Germlings inoculated with the diatom Navicula sp. induced higher settlement than films ofthe diatom species alone. High settlement of up to52% was also achieved with germlings of the greenalga Ulvella lens. Settlement was reduced onU. lens squares inoculated with the diatom Navicula sp. but higher than on films of the diatomalone. The settlement rate was higher if plates withU. lens were previously grazed by juvenileabalone.Post-larval growth rates were higher on monospecificdiatom films than on U. lens or on S.durum. The best growth rate was obtained with Navicula sp. U. lens and S. durum areboth good settlement inducers, but are notsufficient to support rapid growth of young H.rubra post-larvae. Survival was low on U. lensand on the diatom C. closterium. We suggest thatcommercial nursery plates seeded with U. lenswill result in high and consistent settlement, whilean inoculum with Navicula sp. will ensuresufficient food for rapid growth of the post-larvae.     

Resource space partitioning by the Bryozoa of a Fucus serratus L. Community

The spatial distributions of five Bryozoa species encrusting Fucus serratus L. plants in Strangford Lough, Northern Ireland, have been examined in two cross-sectional samples yielding 80 plants and 1326 sampling units drawn from 13 environmentally diverse sites. The co-occurences of Flustrellidra hispida (Fabricius), Alcyonidium hirsutum (Fleming), Electra pilosa (L.), Membranipora membranacea (L.), and Celleporella hyalina (L.) on individual Fucus serratus plants were consistent with independent settlement onto each plant, but their joint distributions within the plants were markedly non-random. Measurements of behavioural interactions between species showed that competitive abilities declined through the sequence Flustrellidra, Alcyonidium, Electra, Membranipora, which parallels the sequence of competition (Levins' α) coefficients calculated from spatial distribution data. Competitive ability varied with position on the plant. Flustrellidra and Alcyonidium reversing their relative competitive superiority between basal and distal regions of the fronds. Flustrellidra and Electra were strongly associated under all conditions. Alcyonidium was also frequently present with this pair, though its numerical abundance was inversely related to that of Flustrellidra. Celleporella tended to co-occur with Electra whilst avoiding Flustrellidra and Alcyonidium. Membranipora numbers were inversely related to those of the other species. Analyses of competition coefficients showed that Flustrellidra, Alcyonidium and Electra each peaked in competitive ability against Membranipora at different points along the frond, such that the latterwaas effectively excluded by direct as well as by diffuse competition. Membranipora populations had a greater dependence on the segment area available for colonization than had any other bryozoan species.     

Effect of ageing on survival of benthic diatom propagules

A comparison of viable benthic diatom propagules based on the observations recorded immediately and after 5 years of ageing at 5 °C is presented. The number of viable benthic diatom propagules decreased with ageing. However, they exhibited an apparently longer lag phase. Although diatoms belonging to the genera Amphora, Navicula and Thalassiosira were dominant during immediate observation, only Amphora and Navicula survived the ageing process. The non-viability of Thalassiosira indicates that ageing for five years was beyond its critical period of survival. The other diatom genera that survived the ageing process were Odontella and Grammatophora.     

The composition and structure of the benthic diatom taxocene (Bacillariophyta) near Cape Fiolent (the Crimea,the Black Sea)

For the first time in the waters of the Cape Fiolent Reserve, a high species richness of benthic diatoms was revealed on natural substrates, 290 species and infraspecific taxa (IT), belonging to 281 species, 68 genera, 32 families, 19 orders, and 3 classes. A total of 68 species and 3 genera were previously registered as new for the flora of the Black Sea; 3 species and 1 taxonomic combination were previously described as new to science, as well as 4 species that did not occur in the Black sea in the past 50 and 100 years of research. Navicula (41 species and IT), Amphora (30), Nitzschia (29), Fallacia (20), Cocconeis (17) and Diploneis (16) were the most represented genera. Only one representative was identified in each of the orders Anaulales, Ardissoneales, Thalassionematales, Rhaphoneidales, and Rhopalodiales. The diversity assessment that was carried out using taxonomic distinctness indices, TaxDI, revealed that the structure of the phylogenetic diatom tree was formed by branches with various species saturation levels and different hierarchical subordination, but with the predominance of polyspecific taxa up to at genera level. The TaxDI values (Δ⁺ = 76.71; Λ⁺ = 361.52) indicate the low evenness and high variability of the structure of the benthic diatom taxocene near Cape Fiolent.     

Grazing and UV radiation effects on an Antarctic intertidal microalgal assemblage: a long-term field study

A 15 week field experiment (austral summer Nov–Mar) was carried out in an intertidal hard bottom platform in Antarctica (King George Island). To test whether grazing and ultraviolet radiation (UVR) influenced the succession of a benthic microalgal assemblage, a two-factorial design was used (1) ambient radiation, >280 nm; (2) ambient minus UV-B, >320 nm; (3) ambient minus UVR, >400 nm versus grazer–no grazer). On four sampling occasions microalgae were identified, counted and carbon contents were calculated. The assemblage was dominated by the diatom genera Navicula and Cocconeis. Biomass was generally low in all treatments but was significantly reduced by grazing throughout the experiment. No significant UV effects were found. Grazer absence particularly favoured diatoms of the genus Cocconeis. We conclude that the Antarctic microalgal assemblage was unaffected by present day UVR whereas grazers acted as important drivers on the intertidal microalgal community structure.     

Diatom community structure on in-service cruise ship hulls

Diatoms are an important component of marine biofilms found on ship hulls. However, there are only a few published studies that describe the presence and abundance of diatoms on ships, and none that relate to modern ship hull coatings. This study investigated the diatom community structure on two in-service cruise ships with the same cruise cycles, one coated with an antifouling (AF) system (copper self-polishing copolymer) and the other coated with a silicone fouling-release (FR) system. Biofilm samples were collected during dry docking from representative areas of the ship and these provided information on the horizontal and vertical zonation of the hull, and intact and damaged coating and niche areas. Diatoms from the genera Achnanthes, Amphora and Navicula were the most common, regardless of horizontal ship zonation and coating type. Other genera were abundant, but their presence was more dependent on the ship zonation and coating type. Samples collected from damaged areas of the hull coating had a similar community composition to undamaged areas, but with higher diatom abundance. Diatom fouling on the niche areas differed from that of the surrounding ship hull and paralleled previous studies that investigated differences in diatom community structure on static and dynamically exposed coatings; niche areas were similar to static immersion and the hull to dynamic immersion. Additionally, diatom richness was greater on the ship with the FR coating, including the identification of several new genera to the biofouling literature, viz. Lampriscus and Thalassiophysa. These results are the first to describe diatom community composition on in-service ship hulls coated with a FR system. This class of coatings appears to have a larger diatom community compared to copper-based AF systems, with new diatom genera that have the ability to stick to ship hulls and withstand hydrodynamic forces, thus creating the potential for new problematic species in the biofilm.     

Invasive species as a new food source: does a nudibranch predator prefer eating an invasive bryozoan?

Membranipora membranacea is an invasive bryozoan that was first found in the Gulf of Maine in 1987 and within two years became the dominant organism living on kelps. Membranipora may have become dominant so quickly because it had little competition in a relatively unoccupied niche; however, lack of predation has also probably played a major role. Where Membranipora is native, there is usually a specialist nudibranch predator that keeps the population in check. For example, in European populations, the nudibranch Polycera quadrilineata prefers Membranipora while Onchidoris muricata is known to prefer another bryozoan, Electra pilosa. Electra, Membranipora, and Onchidoris are all now found in the Gulf of Maine while Polycera is not. We tested whether Onchidoris would (1) eat Membranipora at all, (2) eat Membranipora and Electra at different rates, and (3) show a preference for eating Membranipora or Electra when given a choice. We found that Onchidoris does eat Membranipora, and it generally eats Membranipora faster than Electra. However, when given a choice, Onchidoris prefers Electra. Onchidoris typically reproduces in the spring and grows over the fall and winter, but has recently been found reproducing in the winter in New Hampshire. Although it does not survive the winter as well as Electra, Membranipora is the dominant organism living on many macroalgae in the late summer and fall. Thus, the large Membranipora food source now available in the summer and fall may allow Onchidoris to reproduce earlier.     

Biofilm diatom community structure: Influence of temporal and substratum variability

Abstract

Diatoms, which are early autotrophic colonisers, are an important constituent of the biofouling community in the marine environment. The effects of substratum and temporal variations on the fouling diatom community structure in a monsoon-influenced tropical estuary were studied. Fibreglass and glass coupons were exposed every month for a period of 4 days and the diatom population sampled at 24 h intervals, over a period of 14 months. The planktonic diatom community structure differed from the biofilm community. Pennate diatoms dominated the biofilms whilst centric diatoms were dominant in the water column. Among the biofilm diatoms, species belonging to the genera Navicula, Amphora, Nitzschia, Pleurosigma and Thalassionema were dominant. On certain occasions, the influence of planktonic blooms was also seen on the biofilm community. A comparative study of biofilms formed on the two substrata revealed significant differences in density and diversity. However species composition was almost constant. In addition to substratum variations, the biofilm diatom community structure also showed significant seasonal variations, which were attributed to physico-chemical and biological changes in both the water and substratum. Temporal variations in the tychopelagic diatoms of the water were also observed to exert an influence on the biofilm diatom community. Variations in diatom communities may determine the functional ecosystem of the benthic environment.     

DIATOMS AS HOSTS FOR OTHER DIATOMS: OBSERVATIONS FROM SOUTHERN CALIFORNIA

Large epiphytic and epilithic diatom species hosting other diatoms were observed in several fresh‐ and brackish water sites in Southern California. The most commonly encountered hosts were species forming long filaments attached to rocks or macroalgae, Hydrosera whampoensis (Schwarz) Hendey, Melosira varians Agardh, Pleurosira laevis (Ehrenberg) Compere and Terpsinoe musica Ehrenberg. These large diatoms often had smaller diatoms attached, usually to the girdle bands and occasionally to the mucilage pads connecting the cells. For example, cells of T. musica were observed supporting growth of a diverse diatom assemblage composed of species of the genera Achnanthes, Achnanthidium, Amphora, Cocconeis and Tabularia; Synedra sp. was attached to M. varians and B. paxillifer; and Cocconeis placentula was seen on H. whampoensis. Thus, large epiphytic and epilithic diatoms seem to provide suitable sites for attachment of small epiphytic diatom species, and it appears that this phenomenon is more common than previously thought.     

Adhesion and motility of fouling diatoms on a silicone elastomer

Recent demands for non-toxic antifouling technologies have led to increased interest in coatings based on silicone elastomers that ‘release’ macrofouling organisms when hydrodynamic conditions are sufficiently robust. However, these types of coatings accumulate diatom slimes, which are not released even from vessels operating at high speeds ( > 30 knots). In this study, adhesion strength and motility of three common fouling diatoms (Amphora coffeaeformis var. perpusilla (Grunow) Cleve, Craspedostauros australis Cox and Navicula perminuta Grunow) were measured on a polydimethylsiloxane elastomer (PDMSE) and acid-washed glass. Adhesion of the three species was stronger to PDMSE than to glass but the adhesion strengths varied. The wall shear stress required to remove 50% of cells from PDMSE was 17 Pa for Craspedostauros, 24 Pa for Amphora and >> 53 Pa for Navicula; the corresponding values for glass were 3, 10 and 25 Pa. In contrast, the motility of the three species showed little or no correlation between the two surfaces. Craspedostauros moved equally well on glass and PDMSE, Amphora moved more on glass initially before movement ceased and Navicula moved more on PDMSE before movement ceased. The results show that fouling diatoms adhere more strongly to a hydrophobic PDMSE surface, and this feature may contribute to their successful colonization of low surface energy, foul-release coatings. The results also indicate that diatom motility is not related to adhesion strength, and motility does not appear to be a useful indicator of surface preference by diatoms.     

Microalgal–meiofaunal interactions in a sublittoral site of the Gulf of Trieste (northern Adriatic Sea, Italy): A three-year study

Monthly sediment sampling was carried out from February 1999 to December 2001 at a 21-m deep station. Benthic diatom biomass and meiobenthic abundance were estimated together with abiotic parameters (PAR irradiance, temperature, salinity and dissolved oxygen). During the three-year study average microphytobenthic abundance (ABU) and biomass (BIOM) were 4.7 ± 2.6 × 10⁴ cell cm^− 3 and 14.3 ± 8.1 µg C cm^− 3, respectively, while the mean meiofaunal abundance was 923 ± 210 ind 10 cm^− 2. The microphytobenthic community was mainly composed of Bacillariophyceae (99.3%) with a smaller percentage of phytoflagellates (0.7%). Among diatoms 39 genera were observed with a total of 110 taxa and 100 species. The main benthic diatom genera were Navicula, Paralia, Nitzschia and Diploneis. A total of 18 meiobenthic taxa were detected: 7 taxa belonged to permanent meiofauna (Nematoda, Copepoda Harpacticoida and their nauplius stages, Kinorhyncha, Turbellaria, Gastrotricha, Ostracoda and Acarina). The remaining 11 taxa belonged to temporary meiofauna (juvenile macrofauna: Polychaeta, Cumacea, Amphipoda, Isopoda, Nemertea, Decapoda, Sipunculida, Ophiurida, Gastropoda, Bivalvia and Cnidaria). Positive correlations between PAR and the biomass of the major diatom genera were revealed. Temperature did not seem to influence the microphytobenthic community as a whole, although we found statistically significant correlation between temperature and the genera Nitzschia and Amphora. Salinity showed significant positive correlation only with the genus Pleurosigma, in contrast, dissolved oxygen did not seem to affect the microalgal community. The major meiofaunal taxa were not correlated with abiotic variables, but were positively correlated with most diatom genera. Paralia was negatively correlated with the three main meiofaunal taxa. The three-point moving average applied to our biotic data allowed us to better the signal without the background noise which hid the actual variations of the studied communities. Applying the three-point moving average to the main taxa, it resulted that microphyto- and meiobenthos were in phase. The principal component analysis (PCA), constructed considering both biotic and abiotic variables, accounted for 58% of the total variance. PC1 axis explained 39.74% of the total variance and was correlated with Navicula, Diploneis and Nitzschia. PC2 axis explained 18.40% of the remaining variance and was correlated with Copepoda, Paralia and PAR. In contrast to 1999 and 2001, a mucilage event occurred in 2000, which resulted in the formation of a false bottom and a decrease in the abundance of microphyto- and meiobenthos.     

Reproductive strategies of epialgal bryozoans

Summary

Bryozoans are common encrusting organisms in many shallow-water marine environments. Although reproducing sexually, their success as space occupiers resides largely in their capacity for colonial growth by zooidal budding (regarded by some as a form of asexual reproduction). This paper examines the reproductive strategies of several bryozoan species commonly associated with the fronds of coastal macroalgae. These range from ephemeral species that grow rapidly, reproduce and die (Electra pilosa, Celleporella hyalina, Membranipora membranacea) to more or less annual species with well-developed reproductive and growth cycles (Flustrellidra hispida, Alcyonidium spp.). Whilst many of these bryozoans brood relatively few short-lived lecithotrophic larvae, others produce large numbers of longer-lived planktotrophic larvae. The seasonal and daily patterns of larval release are described for selected species. Resource allocation to sexual and colonial functions is considered in the context of environment-genotype interactions. Reproductive strategy is especially important in determining dispersal and colonising ability, and these in turn are major determinants of ecological pattern in epialgal bryozoan communities.     

Density effects in a colonial monoculture: experimental studies with a marine bryozoan (Membranipora membranacea L.)

Summary Naturally occurring monocultures of plants and animals are not common, despite recent emphasis on the analysis of density effects in artificial plant monocultures. In natural populations, Membranipora membranacea, an encrusting marine bryozoan, usually forms monospecific, nearly even-aged stands on kelp blades. We experimentally manipulated the density of M. membranacea colonies and monitored the responses of individual colonies on settling panels. Colonies undergo a sub-annual cycle of growth, stasis and reproduction, shrinkage, and death. However, crowding by conspecifics accelerates the transition to stasis, triggers early onset of reproduction, and results in increased stage-specific mortality. Unlike many interactions involving colonial invertebrates, overgrowth rarely occurs at boundaries of M. membranacea colonies. Instead, colonies stop growing when they contact conspecifics; therefore more dense assemblages are populated with smaller individual colonies. At the peak in colony size during August, the mean size among colonies grown at high population densities was 300 mm² less than colonies grown at low densities or approximately 62% smaller. Mortality was concentrated in small size classes; at the end of the season colonies gradually shrank to the smallest size classes and then died. We summarized the demography of M. membranacea colonies on low- and high-density panels using size-classified transition matrices and used loglinear analysis to examine the effects of density and time on the transition patterns. As the amount of free space on panels declined, so did the frequency of upward size-class transitions. Our analysis revealed that free space declined more rapidly on panels in the high density treatment and that the transitional probabilities were sensitive to density of conspecifics and seasonal change, but only for some size classes and during some time periods.     

The Diatom Attachment Scar Ophthalmichnus lyolithon igen. et isp. n.

A number of benthic diatoms (including species of the genera Cocconeis, Achnanthes, and Amphora) etch shallow attachment scars in carbonate hard substrates. The morphology of these microbioerosion traces mirrors the elliptical to biconvex outline of the diatoms, which, in its most common appearance, is expressed as a distinct ring-shaped groove. The traces are established as new ichnogenus and -species Ophthalmichnus lyolithon. The diatoms are assumed to form the etching scars by means of their adhesive mucilage composed of acidic polysaccharides, probably in order to enhance adhesion.     

Species- and site-specific bacterial communities associated with four encrusting bryozoans from the North Sea, Germany

Epi- and endozoic bacterial communities associated with four bryozoan species from the Jade (North Sea, Germany) were investigated by the combined application of molecular tools and electron microscopic visualisation of zooids. Denaturing gradient gel electrophoresis of PCR-amplified 16S rRNA gene fragments of associated bacteria displayed specific bacterial community profiles in the examined species Aspidelectra melolontha, Conopeum reticulum, Electra monostachys and Electra pilosa. Actual bacterial epibiosis was only observed on C. reticulum whilst the other bryozoans under investigation were largely free of microbial epibionts. These observations indicated that bryozoan-associated bacteria identified by molecular methods originated from internal cavities of bryozoan zooids. Cluster analysis of DGGE band patterns revealed species-specific bacterial communities in A. melolontha, E. monostachys and E. pilosa. Bacteria associated with C. reticulum were seemingly influenced by site-specific parameters. A comparison of bacterial community profiles between reference and invertebrate surfaces allowed for an interpretation of conspicuous group-specific differences. Operational taxonomic units (OTUs) obtained from a single set of bryozoan replicates that were absent on the inorganic reference samples (mussel shells) were hypothesized to be favourable endobionts. Contrary, OTUs present in the references but absent in bryozoan samples could be assumed to stem from bryozoan-specific defenses against ubiquitous bacterial colonizers. Although there was no experimental evidence for a mutual relationship between prokaryotes and their eukaryotic bryozoan hosts, this study demonstrated that in three out of four bryozoans under investigation associated bacterial communities were characteristically shaped by host attributes.     

Seasonal variations in the fouling diatom community structure from a monsoon influenced tropical estuary

Seasonal variations in the fouling diatom community from a monsoon influenced tropical estuary were investigated. The community composition did not differ significantly between stainless steel and polystyrene substrata due to dominance by Navicula spp. The experimental evidence suggests that Amphora, which is a dominant fouling diatom in temperate waters, ultimately dominates the community in tropical regions when conditions are favourable. These observations reveal that a faster onset of macrofouling interferes with the microfouling community wherein the faster recruiters that have a higher abundance in ambient waters, dominate the community. Seasonal variations were observed in the fouling diatom community. Navicula delicatula dominated during the post-monsoon and tychopelagic diatoms (Melosira and Odontella) were observed during the monsoon. Low diatom abundance was recorded during the pre-monsoon season. The results indicate that although the fouling diatom community composition does not vary between substrata, there is a seasonal change in the community depending on the physical, chemical and biological interactions.     

Colony Structure and Seasonal Differences in Light and Nitrogen Modify the Impact of Sessile Epifauna on the Giant Kelp Macrocystis pyrifera (L.) C Agardh

The presence of the stoloniferous hydroid Obelia geniculata (L.) had no effect on the pigment concentration or nitrogen status of underlying blade tissue of the giant kelp Macrocystis pyrifera (L.) C. Agardh. The sheet-like colonies of the bryozoan Membranipora membranacea (L.) markedly reduced the pigment concentration of colonized blade tissue, but only during winter. Reductions in pigment concentration are most likely a result of damage to underlying tissue due to some factor related to the presence of bryozoan colonies on blade surfaces. Blade tissue colonized by M. membranacea also had higher δ¹⁵N signatures than surrounding bryozoan-free tissue, possibly indicating the provision of nitrogen to M. pyrifera by bryozoan colonies. Results show that seasonal changes in nitrogen and colony size can strongly modify the effect of epifauna on macroalgae they colonize. Unlike bryozoans, hydroid colonies provided no barrier to nitrogen uptake by colonized M. pyrifera tissue and enhanced ammonium uptake was observed for tissue colonized by O. geniculata during nitrogen limitation. Epifauna with stoloniferous growth forms such as hydroids are more likely to have benign or even mutualistic relationships with macroalgae they colonize than the sheet-like colonies of bryozoans.