Selective grazing by the mud snail Ilyanassa obsoleta

Summary Mud snails (Ilyanassa obsoleta) starved for 48 h were allowed to feed on sediments in laboratory microcosms. Sediment cores sliced at 2 mm intervals were compared to snail stomach contents for per cent carbon and nitrogen, plant pigment contents and species composition of benthic diatoms. Concentrations of carbon, nitrogen, phaeopigments, phycocyanin and chlorophyll were enriched in the top 2 mm of the sediments compared to 7–10 mm depth by a factor of 2–10. In turn, these materials were 20–40 times more concentrated in snail guts than in the surface sediments. Snail feces were enriched for carbon and nitrogen by 5–7 times over the surface sediments. Bacterial chlorophyll peaked at about 3–4 mm in the sediments and was not detectable in the snail stomach contents. The C/N ratio of the snail stomach contents was only 6 compared to a ratio of 8.5 for their feces and 12 for the surface sediments.The percentage of migratory diatoms (e.g. Nitzschia and Navicula) decreased with depth where non-migratory species, such as Fragilaria pinnata, dominated. These migratory species were more common in the snails than in the sediments on which they were feeding.A comparison of daily ingestion rates to the animal's energy budget shows that this selective ingestion is sufficient to meet Ilyanassa's energy needs.     

171 Light-Induced Growth of Winter Phytoplankton Collected from the Benthic Boundary Layer in Coastal Oregon Waters

The objective of this study was to examine the influences of the adjacent river and surrounding landuse on wetland diatom distributional patterns. Diatoms were identified in surface sediment samples from 35 riverine-impounded wetlands within the Willamette Valley, OR. A total of 221 species were identified and no single species dominated the assemblage at all sites. Diatom richness was high throughout the area (median 71, range 66–75). The Araphidineae:Centrales index, a measure of periphytic to planktonic species ratio, was lowest at sites within the river's annual floodplain zone. A low A:C index is to be expected at wetlands that receive inputs of river planktonic species through regular flooding by adjacent large rivers. Surrounding landuse can confound the influence of riverine flooding on wetland diatom assemblages by influencing water quality. Relative abundance of the two most common species, Aulacoseira crenulata and Fragilaria capusina related negatively to % agricultural landuse surrounding the wetland. These results indicate that riverine wetland diatom assemblages may be influenced by flooding from adjacent rivers and surrounding landuse and therefore wetland sediment diatoms might serve as useful indicators of both historical environmental changes in nearby large rivers and surrounding watersheds.     

Ingestion by deposit-feeding macro-zoobenthos in the aphotic zone does not affect the pool of live pelagic diatoms in the sediment

The ability of deposit feeders to utilise the pool of live planktonic diatoms in the sediment was investigated after the spring bloom in 2001 and 2002 at four sediment sites in the aphotic zone in the Kattegat. Seven species of deposit-feeding bivalves, gastropods and polychaetes were allowed to defecate in containers with filtered seawater. A total of 22 containers were set up holding between 3 and 32 animals, grouped by station and taxa. When defecation was completed, the animals were re-introduced to different stocks of homogenised sediment which were manipulated by addition of luminophores and spores of the diatom Chaetoceros diadema, and the faeces again collected and analysed. Our analysis included the number of germinable planktonic diatoms, measured by the dilution extinction method, and the concentrations of the algae pigments fucoxanthin and chl a, in the faeces samples, in the sediment stocks and in the sediment at the sampling location. Comparison between the sediment composition at the sampling locations, and the composition of the corresponding faeces, showed that the concentration of germinable diatoms averaged about 50,000 g⁻¹ dry weight (DW) in the surface sediment as well as in the faeces in 2001, while in 2002, the average concentration was about 390,000 g⁻¹ DW in both faeces and in the sediment. A similar comparison in the second part of the experiments also showed that the diatoms in the sediment were unaffected by gut passage and the added spores of C. diadema germinated in the same quantity in the faeces as in the sediment. The taxonomic composition of the diatoms in the faeces matched the composition at the respective stations and sediment stocks. Finally, the concentration of pigments did not change significantly during gut passage. The biomass of live planktonic diatoms in the area of study was estimated to be 2-10 g DW/m² or 0.2-5% of the total organic content in the top 3 cm of the sediment. It is concluded that this pool of diatoms is largely unaffected by deposit feeders and seems to be unimportant as a food source. It is hypothesised that the input of the spring bloom should be considered as composed of two fractions playing different roles for the benthic ecosystem. One fraction of dead organic material may be utilised immediately, while the fraction of live planktonic diatoms serve as a stable food buffer, which gradually become available to deposit feeders after the diatoms die and degradation starts.     

Short and long-term movement and site fidelity of juvenile Haemulidae in back-reef habitats of a Caribbean embayment

In the context of a main project that aims to recover modern data on diatom distribution applicable to paleosalinity reconstructions in coastal areas of Southern South America, the composition and distribution of dead diatom assemblages in the littoral zone of the Quequén Salado estuary (Argentina) were studied. Diatom zones were defined along the estuarine gradient by cluster analysis and related to the salinity range and sediment composition by Canonical Correspondence Analysis. Four diatom zones were identified. A mixture of marine, brackish and freshwater diatoms, probably allochthonous, characterized the inlet (zone I). Marine/brackish taxa, represented mainly by Paralia sulcata dominated zone II, characterized by polyhaline conditions and sandy sediments. Zone III was characterized by mesohaline conditions, muddy sediments and the dominance of the estuarine diatom Amphora helenensis. Brackish/freshwater and freshwater diatoms dominated the headwaters (zone IV), where salinity was always below 5‰. The comparison of Quequén Salado diatom assemblages with previous results from the Quequén Grande estuary showed a similar taxonomic composition between both estuaries. However, differences in the salinity ranges of the estuaries (related to differences in the degree of human impact and tidal range) lead to a displacement in their spatial distribution along the longitudinal estuarine axis. This paper contributes to the knowledge of the ecological requirements of South American estuarine diatoms and provides useful data for paleosalinity reconstructions in the region. Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users Handling editor: J. Padisak     

Spatial variations in the water quality of Lough Erne, Northern Ireland, on the basis of surface sediment diatom analysis

SUMMARY. 1. Constrained cluster analysis of thirty-five surface sediment diatom assemblages shows that Lough Erne, a complex multi-basin lake in Northern Ireland, can be sub-divided into four main zones.
2. All areas of the lake system have planktonic diatoms typical of eutrophic lakes but the lack of small Stephanodiscus species in zone 4 suggests that this area of the system is least productive.
3. Surface sediment diatom assemblages can be used to assess spatial variations in complex lake systems and can thereby be used as a guide to the choice of sites for water quality monitoring and sediment coring.     

南海某些表层沉积硅藻的分布及其古环境意义

本文通过对南海表层沉积物样品的硅藻分析,试图揭示南海某些表层沉积硅藻的分布规律,以便为南海古海洋学研究提供科学依据。研究发现,Thalassionema nitzshioides为南海地区表层沉积硅藻的优势种,且其百分含量无明显的区域差异,体现了南海与周围海洋水体交换相对较弱的特征。除此之外,南海表层沉积硅藻的分布主要受到海洋环流的影响,表现在黑潮暖流、印度洋暖水的入侵以及沿岸流对南海表层沉积硅藻分布的影响。其中,Nitzshia marina, Azpeitia neocrenulata , Azpeitia africana , Rhizosolenia bergonii等暖水硅藻可作为黑潮暖流及印度洋暖水入侵南海强度的指标种。而Cyclotella stylorum , Cyclotella striata , Diploneis bombus , Traychneis aspera , Tabularia tabulata等则可看作判断沿岸流对南海水体影响强度的标志种。     

170 Wetland Diatom Assemblages as Indicators of Flooding Influence and Surrounding Landuse in the Willamette Valley,OR

The objective of this study was to examine the influences of the adjacent river and surrounding landuse on wetland diatom distributional patterns. Diatoms were identified in surface sediment samples from 35 riverine‐impounded wetlands within the Willamette Valley, OR. A total of 221 species were identified and no single species dominated the assemblage at all sites. Diatom richness was high throughout the area (median 71, range 66–75). The Araphidineae:Centrales index, a measure of periphytic to planktonic species ratio, was lowest at sites within the river's annual floodplain zone. A low A:C index is to be expected at wetlands that receive inputs of river planktonic species through regular flooding by adjacent large rivers. Surrounding landuse can confound the influence of riverine flooding on wetland diatom assemblages by influencing water quality. Relative abundance of the two most common species, Aulacoseira crenulata and Fragilaria capusina related negatively to % agricultural landuse surrounding the wetland. These results indicate that riverine wetland diatom assemblages may be influenced by flooding from adjacent rivers and surrounding landuse and therefore wetland sediment diatoms might serve as useful indicators of both historical environmental changes in nearby large rivers and surrounding watersheds.     

Diatom assemblages in superficial sediments from the Gulf of Riga,eastern Baltic Sea

Twenty one superficial sediment samples from areas of high and moderate eutrophication in the Gulf of Riga were studied with respect to siliceous microfossils, mainly diatoms.The results seem to imply that the number of taxa and the abundance of the frustules are affected by runoff from rivers and the degree of eutrophication in different parts of the Gulf. Areas with high eutrophication, e.g. the river estuaries, have diatom assemblages of varying composition, while in areas with moderate eutrophication the composition is almost constant and the influx of freshwater diatoms and littoral periphyton small. The high abundance and low diversity of brackish-marine and brackish, planktonic diatoms seem to be a result of an influx of nutrients and pollutants in the water and bottom sediments. The sea ice diatoms occurring in the Baltic waters and also in the Gulf of Riga tend to be resistant to eutrophication or are even favoured by it.     

The effect of feeding by mud snails, Ilyanassa obsoleta (Say), on the structure and metabolism of a laboratory benthic algal community

Eastern mud snails, Ilyanassa obsoleta (Say), in densities of 0, 80, and 160 snails · m^?2 were placed in flow-through laboratory microcosms containing 5 cm of frozen and sieved sediments. Other microcosms were raked once daily to a depth of 10 mm. All these containers were incubated for 5 wk and regularly sampled for plant pigments and light and dark transfer of oxygen and carbon dioxide. Feeding at the low density significantly increased chlorophyll standing stock. Respiration and gross photosynthesis increased by an even greater percentage compared to ungrazed controls. Standing stocks of algal pigments, respiration, and photosynthesis were depressed in microcosms which received the 160-snail or raking treatments.The dominant benthic algae in the containers were pennate diatoms. Grazed containers contained a larger percentage of non-motile as compared to motile forms.Sediments fertilized with ammonium at a rate equivalent to excretion of six snails, showed increased chlorophyll content equal to the 80-snail treatment. Daily raking inhibited this effect.We conclude that low densities of Ilyanassa obsoleta stimulate algal growth by accelerating nitrogen cycling and selectively removing specific components of the diatom community. At high snail densities these effects are overwhelmed by overgrazing and sediment stirring.     

Fate of Bacillus sphaericus 2362 spores following ingestion by nontarget invertebrates.

Elimination of Bacillus sphaericus spores ingested by midge larvae, snails, and oysters was most rapid among midge larvae. Spores remained in oysters up to 21 days and in snails up to 49 days. Viable spores were recovered in snail and oyster feces for these same periods. There was no indication of actively growing B. sphaericus in the animals. Passage through oyster gut detoxified the B. sphaericus mosquito larval toxin, but there was a 33% retention of toxicity following snail gut passage. Midge larvae reared to adults in spore-containing water carried spores in/on the adult body. This suggests that these animals could carry the bacteria to sites beyond the application area.     

Sedimentary diatom assemblages and their utility in computing diatom-inferred pH in Sudbury Ontario lakes

Surface sediment diatoms from 30 Sudbury lakes were analyzed to establish a relationship between diatoms and limnological characteristics of these lakes. Factor analysis of various chemical and physical variables suggested that most of the variance was associated with the pH, pH-metal, and conductivity factors. Factor analysis of common diatom taxa indicated that the distribution of most of these diatoms is closely related to lake water pH and/or pH related factors. Regression analysis of various pH indicator assemblages with measured lake water pH also demonstrated significant relationships. Among the transfer functions employed to compute diatom-inferred pH, multiple regressions using pH indicator assemblages proved to be the best. Using this method an accuracy of about 0.3 pH unit can be achieved, even for lakes which have received large trace metal inputs concurrent with increase in acidity. The presence of planktonic diatoms in acidic lakes suggests that the widely accepted hypothesis that planktonic diatoms are greatly reduced in waters at pH below 5.6 is not always true.     

A change in the distribution pattern of diatoms above and below the halocline in bottom sediments of the central Arkona Basin,southern Baltic

Surface sediment samples from the central Arkona Basin, southern Baltic, were investigated by diatom analysis. The samples, taken by a Niemistö sampler, included the sediment-water interface and about 2–3 centimetres of the very water-saturated sediment. The samples represent the sediments deposited in the transition layer of the halocline in the Arkona Basin and immediately above and beneath it. The distribution pattern of diatoms and diatom pieces changed, reflecting the position of the halocline.Diatom analysis is an important tool in identifying Holocene Baltic sediments, and the knowledge of recent hydrological and chemical factors and their impact on the distribution pattern of diatoms is one way to evaluate the usefulness of diatom records in postglacial Baltic marine sediments. The surface-sediment diatom assemblages can be compared with the data from postglacial Baltic offshore cores. They give prelimary indications where diatom records from Littorina or Yoldia stages could be expected.     

Diatom succession in the post-glacial sediments of the Komořany Lake,North-West Bohemia,Czechoslovakia

The Late-glacial and Postglacial sediments of the former Komo?any Lake in North-West Bohemia were studied by means of diatom analysis to trace the pattern of past environmental change in the lake. Several phases of lake development could be distinguished. Originally the Komo?any Lake was a eutrophic shallow water basin. The diatom succession in the early Postglacial sediment indicates alternating alkaliphilic, epiphytic and planktonic diatom communities associated with changes in water level in the basin. At the beginning of the Subatlantic period the water level began to rise and the development of planktonic diatoms, typical of small eutrophic lakes, occurred. Later, the basin became shallow again and the number of indifferent and acidophilous diatoms increased. This is typical for the final phase of the infilling process of a lake before its transformation to peatland     

Hidden diversity in diatoms of Kenyan Lake Naivasha: a genetic approach detects temporal variation

This study provides insights into the morphological and genetic diversity in diatoms occurring in core sediments from tropical lakes in Kenya. We developed a genetic survey technique specific for diatoms utilizing a short region (76-67 bp) of the ribulose-1,5-bisphosphate carboxylase/oxygenase large subunit (rbcL) gene as genetic barcode. Our analyses (i) validated the use of rbcL as a barcoding marker for diatoms, applied to sediment samples, (ii) showed a significant correlation between the results obtained by morphological and molecular data and (iii) indicated temporal variation in diatom assemblages on the inter- and intra-specific level. Diatom assemblages from a short core from Lake Naivasha show a drastic shift over the last 200 years, as littoral species (e.g. Navicula) are replaced by more planktonic ones (e.g. Aulacoseira). Within that same period, we detected periodic changes in the respective frequencies of distinct haplotype groups of Navicula, which coincide with wet and dry periods of Lake Naivasha between 1820 and 1938 AD. Our genetic analyses on historical lake sediments revealed inter- and intra-specific variation in diatoms, which is partially hidden behind single morphotypes. The occurrence of particular genetic lineages is probably correlated with environmental factors.     

Weak diurnal changes in the biochemical properties and benthic macrofauna of urbanised mangrove forests and mudflats

Diurnal changes in the biochemical properties and the benthic macrofaunal assemblage of sediments in urbanised mangrove forests and their adjacent mudflats in Sydney Harbour were investigated. Behavioural and physiological changes in the microphytobenthos between day and night were predicted to cause diurnal changes in the micro-scale depth distribution of chlorophylls a and b and colloidal carbohydrate. In addition, because macrofauna can alter sediment properties, diurnal changes in the macrofaunal assemblages were investigated. The microphytobenthos at the study sites were predominantly filamentous green algae, although diatoms were present. Samples for biochemical analysis were collected from the top 2 mm of sediment using mini-cryolanders, during low tide in the day and at night. Three biochemical properties of the sediments were measured spectrophotometrically: chlorophylls a and b (surrogate for microphytobenthos biomass) and colloidal carbohydrate. The amount of chlorophylls tended to be less at night than during the day, but site to site variability was large and these differences were generally small and not significant. Depth profiles indicated that there was some redistribution of pigments in the surface 2 mm between day and night, possibly due to migration of microphytobenthos or physiological changes. There was no significant difference in chlorophylls between the mangrove forest and adjacent mudflat, with the exception of chlorophyll b at one sampling time, which was larger in the mangrove forest than on the mudflat. Colloidal carbohydrate was significantly larger in the mangrove forest and significantly less on the mudflat during the day at one site at one time, but otherwise showed no significant differences between day and night or between the mangrove forest and mudflat. Whilst there were some differences in the benthic macrofaunal assemblages between day and night, these differences were only significant for spionids and polychaetes at one time. There were, however, significant differences in assemblages of benthic macrofauna between the mangrove forest and mudflat, probably due to structural differences between these habitats such as the presence of pneumatophores, shade and leaf litter. In summary, there was some minor diurnal variation in the measured biochemical properties of the sediment, but not in the macrofaunal assemblage. Diurnal changes should, therefore, be considered when investigating biochemical properties in these habitats, but they are not a major influence. These findings contrast to previous studies on diatom dominated mudflats in Europe, where stronger diurnal changes in biochemical properties were found. Diurnal changes in the macrofauna assemblages were largely insignificant and therefore could not explain the changes in the biochemical properties. Diurnal effects on the macrofauna in these habitats are more likely to be via altered behaviours and this requires further investigation.     

Microscale imaging sheds light on species-specific strategies for photo-regulation and photo-acclimation of microphytobenthic diatoms

Intertidal microphytobenthic (MPB) biofilms are key sites for coastal primary production, predominantly by pennate diatoms exhibiting photo-regulation via non-photochemical quenching (NPQ) and vertical migration. Movement is the main photo-regulation mechanism of motile (epipelic) diatoms and because they can move from light, they show low-light acclimation features such as low NPQ levels, as compared to non-motile (epipsammic) forms. However, most comparisons of MPB species-specific photo-regulation have used low light acclimated monocultures, not mimicking environmental conditions. Here we used variable chlorophyll fluorescence imaging, fluorescent labelling in sediment cores and scanning electron microscopy to compare the movement and NPQ responses to light of four epipelic diatom species from a natural MPB biofilm. The diatoms exhibited different species-specific photo-regulation features and a large NPQ range, exceeding that reported for epipsammic diatoms. This could allow epipelic species to coexist in compacted light niches of MPB communities. We show that diatom cell orientation within MPB can be modulated by light, where diatoms oriented themselves more perpendicular to the sediment surface under high light vs. more parallel under low light, demonstrating behavioural, photo-regulatory response by varying their light absorption cross-section. This highlights the importance of considering species-specific responses and understanding cell orientation and photo-behaviour in MPB research.     

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.     

Microplankton response to environmental conditions in the Alboran Sea (Western Mediterranean): One year sediment trap record

The present work analyses the seasonal evolution of planktonic assemblages and particle fluxes through the water column in the Eastern Alboran Sea (Western Mediterranean) at 35º55.47'N/01º30.77'W. A Sediment trap was deployed below the influence of the Almeria-Oran Front (AOF), a semi-permanent geostrophic front, during July 1997 to June 1998. Overall, the temporal variability of coccolithophore, planktonic foraminifer, diatom, benthic and wind-carried biogenic particle fluxes is linked to the seasonal evolution of sea surface hydrological structures. Maximum planktonic fluxes were found during high-productivity periods and wind-induced upwelling, following a trimodal pattern, with maximum fluxes in July 1997, November–December 1997, and April–May 1998. These periods were characterized by vertical mixing and the full development of anticyclonic gyres in the Alboran Sea. The annual flux of coccolithophores was dominated by the “small Gephyrocapsa Group” and Emiliania huxleyi, whereas Turborotalita quinqueloba and Globigerina bulloides dominated the foraminiferal fluxes, and Chaetoceros Resting Spores (RS) were predominant in the diatom assemblage. Benthic specimens were also collected with the sediment trap, suggesting a variable influence of bottom water activity. Wind-driven particles (phytoliths and fresh-water diatoms) were collected along the year, but their fluxes followed the local wind regime.The high Sea Surface Temperature (SST) during fall due to weaker than usual westerly winds, and the pressure anomaly prevailing in the Alboran Sea during early winter, were reflected in the planktonic assemblages by the proliferation of warm, lower photic layer inhabitants and/or oligotrophic taxa of coccolithophores (Florisphaera profunda), planktonic foraminifers (Globigerinoides ruber and Globorotalia inflata) and diatoms (Leptocylindrus danicus). These unusual climatic conditions in the eastern Alboran Sea must have been caused by the 1997–1998 ENSO event.     

Physical microhabitat requirements of freshwater pearl mussels,Margaritifera margaritifera (L.)

Surface sediment diatoms from the east coast of Lake Tanganyika were analysed using ordination and classification techniques, and compared with assemblages previously described from the northern part of the lake. Grain-size analyses were performed on subsamples. Four groups of diatom assemblages were recognised. The first group clusters samples taken in the north, far from the Rusizi river mouth. The second group comprises samples taken on silty sediment along the Tanzanian coast, including one sample taken near the mouth of the Malagarazi river and those from the northernmost part of the lake. The third group comprises surface sediments along the Burundian coast (near Ramba and Magara), and the fourth is characterised by epipsammic taxa. A sample taken near the central arm of the Malagarazi river is included in the latter group. The impact of small rivers on the diatom assemblages in the surface sediments is restricted to the mouth area.     

Changes in diatom assemblages in the profundal sediments of two large oligohumic lakes in eastern Finland

Sedimentary diatom assemblages in two large oligotrophic clear-water lakes were analysed, to assess their present ecological state and possible eutrophication due to diffuse nutrient loading. The lakes Pyhäjärvi and Puruvesi (Finnish lake district) are proportionally large for their catchment areas which accounts for their long retention times (ca 7 and 11 yr) and oligohumic character. Pyhäjärvi was studied by pairwise comparison of surface sediment diatom assemblages collected in 1985 and 1990 at 12 sites from different parts of the lake. In Puruvesi, the stratigraphy of diatoms was analysed in two short cores from 8 m and 32 m depths.The diatom assemblages of the two lakes are rather similar, and quite distinct from the assemblages of the mesohumic large lakes of the area. Cyclotella kuetzingiana is the most common planktonic dia- tom, but Aulacoseira ambigua abounds in Pyhäjärvi at sites with local sources of eutrophication. A diverse assemblage of benthic forms, especially Fragilaria and Achnanthes spp. characterizes the shallow bottoms in both lakes.There was little change within the short-core diatom profiles of Puruvesi, but the floral composition of the 8-m and 32-m sites differed markedly. The 8-m site, with 60–70% of benthic forms, represents illuminated bottom, on which much of the buried algae have lived in situ, while the deeper site is true profundal, dominated by sedimented planktonic algae.In Pyhäjärvi there was a slight increase in the benthic diatoms from 1985 to 1990, coinciding with increased phosphorus and chlorophyll concentrations as well as Secchi depth lowering. We interprete this observation as a very early step of eutrophication, of which first the sessile algal communities of the illuminated bottom areas have benefited.