Colonization Dynamics of an Encrusting Cyanobacterial Mat in a Hardwater River (Eaulne,France)

Many hardwater rivers of the Seine basin (France) have gradually become affected by extensive stromatolite ( sensu Monty 1973) growth during the last 30 years. In some cases, calcareous buildup development is fast enough to cement large sectors of the stream bed. Although static aspects (mineralogy, morphology, microflora) are well known, few attempts have been made so far to understand the dynamics and regulating factors of buildup development. The formation of an encrusting algal mat was studied from 1996 to 1998 in a hardwater river of Seine-Maritime (France). The structural organization of all stromatolitic buildups was highly homogeneous in all sites. Encrusted microflora settled only on stable, solid substrates and were largely dominated by the cyanobacterial genus Phormidium . Annual dynamics of calcitic structure development closely followed the cyanobacterial microflora growth phases. The main determining factor of encrusting microflora extension appeared to be river-bed stability. Current velocity seems to play a fundamental part on at least two scales. On the "site scale" (100 m²), the discharge variations determine the spatial distribution of the fine-grained, unstable deposits that cannot host encrusting microflora. On the "substrate scale" (0.1 to 1 m²), the maximum cyanobacterial biomass increases with current velocity. The maximal extension rate measured in the present study showed that a stable, cobble river floor could be cemented by cyanobacterially driven calcareous deposits in 6 to 7 months.     

Larger foraminifera as a substratum for encrusting bryozoans (Late Oligocene,Tethyan Seaway,Iran)

Considering the diversity and abundance of larger foraminifera examined from a wide range of Late Oligocene to Early Miocene palaeoenvironments in the Tethyan Seaway, encrusting bryozoans make extremely little use of their tests as substratum. Significant encrustations by bryozoans were exclusively found on large (ø c. 6 cm), undulating tests of Lepidocyclina spp., on which, however, a remarkable 34 taxa of encrusting bryozoans were recorded. This shallow-water fauna of Chattian age was analyzed in respect of the bryozoan taxa present, colony growth type, and mode of budding, colony size, as well as onset of reproduction. Taxic and morphological similarities between the fossil assemblage and modern faunas encrusting mobile substrata indicate a long history of bryozoans as part of the interstitial habitat, while the tests of certain larger foraminifera may have played a significant role in the evolution of shallow-water bryozoans by providing substrata for encrusting species in otherwise unfavorable environments.     

A new encrusting interstitial marine fauna from Brazil

Abstract. This paper reports the second occurrence of a sand‐grain encrusting interstitial epifauna dominated by bryozoans and polychaetes at a site thousands of kilometers from the first described occurrence of such a fauna 20 years ago. Such faunas seem to have gone almost unrecorded in the marine ecological literature, but they are potentially geographically widespread and ecologically significant, deserving recognition and further study by benthic ecologists. Although rooted‐erect and free‐living lunulitiform bryozoans can be abundant in soft‐bottom habitats, the presence of encrusting forms was, until recently, considered to be limited to patches of hard substrata. In 1985 and 1988, a new and seemingly unique habitat for encrusting bryozoans and other organisms on single grains of shell or sand was reported from the coastal waters of Florida, USA. Here we report a second discovery of an interstitial encrusting fauna from the continental shelf off the state of São Paulo, Brazil. In addition to the cupuladriid Discoporella umbellata, several species of bryozoans (9 cheilostomes, 3 ctenostomes, and 1 cyclostome) were found encrusting on or boring into sand grains from the 4 stations examined. Four species were found exclusively on sand to gravel size grains. The most abundant colonies, with ～1300–1500 colonies m^?2, belonged to a new species of Cleidochasma. New species of Trypostega and Reginella, each with up to 200–300 colonies m^?2, were also discovered. The grain‐encrusting bryozoans were characterized by their small size, and by the fact that sexual reproduction was initiated very early in colony growth; brood chambers (for the development of embryos into larvae) occurred in colonies having only a few zooids. Colonies of boring ctenostome and cheilostome bryozoans were even more abundant than those of grain encrusting forms, being present in almost every piece of shell (～5000–5500 colonies m^?2). The fauna also included representatives of other groups of encrusting organisms, especially tubeworms (11,000–13,000 tubes m^?2). Planned work on samples from additional stations on the São Paulo shelf will no doubt yield a larger number of species from various taxa and perhaps show some overlap in sand fauna species between the Brazilian and Floridian sites. In addition to the unique species of single grain encrusters, colonies of bryozoan species characteristic of larger subtidal hard substrata were also found on sand or gravel size grains, indicating that an interstitial refuge may be available to some epifaunal taxa and suggesting that this interstitial refuge, which remains almost completely unknown to benthic ecologists, may play a large role in determining distributions of those taxa.     

Bryozoan assemblages on hard substrata: species abundance distribution and competition for space

Bryozoans are colonial invertebrates that often dominate epibenthic assemblages on the lower surfaces of hard substrata. Competition among neighbouring organisms is usually a critical process regulating biodiversity, and hard substrata have proved to be a suitable model habitat to analyse spatial interactions. We explored the relationships among abundance, species richness, diversity, competitive ability, coverage, available surface, depth and substratum size in an assemblage of bryozoans encrusting pebbles and cobbles in a bank off the eastern mouth of the Strait of Magellan. We also tested whether overgrowth competition can be regarded as hierarchical, and whether the species abundance distribution shows a mode of rare species and a decreasing frequency of increasingly abundant species. Abundance, species richness, diversity and overgrowth competition were highest on the largest substrata. Smaller pebbles tended to be encrusted by the commonest bryozoans, while the rarest species were mainly found on relatively larger clasts. A high proportion of the lower surfaces of most substrata was available for growth. Diversity values of relatively shallow stations were lower than expected under Caswell’s neutral model. Interspecific competition was hierarchical, but the abundance of colonies was not related to the competitive ability of each species. The species abundance distribution was bimodal, with a main mode of rare species and a second one partly composed of relatively abundant bryozoans with poor competitive abilities. This study shows that even in an encrusting assemblage where competition is hierarchical, the weakest competitors persist and the dominant species are far from being capable of monopolizing space.     

Structural and taphonomic analysis of a columnar coralline algal build-up from SE Sicily

Unusual “coralligenous” build-ups were found at the infra–circalittoral boundary from SE Sicily. They are columnar, rising up perpendicularly from the bottom, and consist of an inter-growth of encrusting algae and invertebrates. A selected build-up and its longitudinal section, together with some thin sections of selected parts, were studied to detail the growth structures, composition and fabric of the trapped sediments. The coralline algae show a succession of “concentric”, “fruticulose” and “foliaceous” structures. The framework delineates cavities bounded by algal thalli, sometimes filled by sediments. The organisms constituting the framework were grouped into functional guilds (Fagerström, 1991): “primary frame builders” (mainly coralline algae); “secondary frame builders” (vermetids, serpulids and some bryozoans); “bafflers” (erect, flexible bryozoans, some sponges and soft algae); “binders” (encrusting bryozoans, some crustose coralline algae and serpulids); “destroyers” (a few endolithic algae, rare sponges and bivalves); “dwellers” (brachiopods, foraminifers, bivalves, small bryozoans and some motile organisms). The distribution pattern of taphonomic features (composition, grain size and distribution of sediments filling cavities; presence of cement and its location; mineralogical composition and crystal size), together with interactions between growth structures and functional guilds, allowed one to distinguish three taphofacies, each recording different environmental conditions. They have been tentatively related with cyclic sequences testifying to local hydrodynamic regimes, with subsequent slighter and stronger intensity phases, during the last 2200 years.     

Sponge-mediated coral reef growth and rejuvenation

Sponges mediate consolidation of Porites furcata rubble on shallow Caribbean reefs by quickly adhering to rubble and stabilizing it until carbonate secreting organisms can grow and consolidate it to the reef. Experimental investigations demonstrate that the entire cycle from (1) temporary binding of rubble by sponges, through (2) rubble consolidation by encrusting coralline algae, to (3) colonization of consolidated rubble by corals, can be completed within 10 months. Bound rubble both adds to vertical reef growth and also provides stable substrata for colonization by corals. Corals that colonize stabilized rubble are damaged less and survive better than on unstable rubble. Rubble that is not temporarily stabilized by sponges does not become bound to the reef, because continuous movement disturbs the consolidation process, and does not provide suitable substrata for settlement and growth of corals. Sponge-mediated consolidation of rubble may increase rates of reef growth and enhance reef recovery after damage. This new role for sponges in reef growth is not obvious from examination of the internal fabric of a reef frame. Spongemediated consolidation may help to explain geographic and temporal differences in growth and morphology among shallow reefs of ramose corals.     

Influence of Bed‐Sediment Features on the Interstitial Habitat Available for Macroinvertebrates in 15 French Streams

The interstitial space of bed‐sediments is a major component of the stream system as it influences both physical and biological processes and patterns, such as exchanges between surface water and groundwater or the assemblage of invertebrates. To investigate the influence of grain‐size distribution and vertical bed‐packing on the interstitial habitat, we used 99 frozen cores collected from 15 French streams. On our data, overall invertebrate density, density of Chironomidae and taxon richness were more strongly related to effective porosity than to other physical variables describing bed‐sediments. We built two models to quantify the influence of grain‐size distribution and vertical bed‐packing on effective porosity. The effective porosity decreased with increasing grain‐size heterogeneity and was strongly related to grain‐size distribution. These relationships differed with vertical bed‐packing, but were consistent across three geologic types of sediments that varied in particle shape and particle texture. These general relationships suggest that effective porosity could be helpful to better understand the distribution patterns of invertebrates in bed‐sediments.     

Three-year study of benthic algal spring bloom development in a small, Danish lowland stream

Seasonal development of benthic algae was studied over a three-year period in a small, nutrient-rich lowland stream to investigate inter-annual variation in the algal spring bloom and differences in algal biomass regulation on two different substrata: fine-grained sediments and stones. The algal spring bloom was initiated when irradiance at the sediment surface exceeded 7 mol photons m^-2 d^-1 and mean water velocity was concomitantly below the threshold for bed load transport in the stream. Large inter-annual and substratum-dependent differences in peak algal biomass were observed, thus suggesting that different parameters regulate algal biomass development on the two substrata. On fine-grained sediments algal biomass development was predominantly coupled to light availability, while on stony substrata algal composition and peak biomass might be affected by invertebrate grazing.     

Structure and development of a benthic marine microbial mat

Abstract Vertically stratified microbial communities of phototrophic bacteria in the upper intertidal zones of the North Sea island of Mellum were investigated. Growth and population dynamics of the cyanobacterial mat were followed over three successive years. It was concluded that the initial colonization of the sandy sediments was by the cyanobacterium Oscillatoria . In well-established mats, however, the dominant organism was Microcoleus chthonoplastes . The observed succession of cyanobacteria during mat development is correlated with nitrogen fixation. Nitrogen fixation is necessary in this low-nutrient environment to ensure colonization by mat-constructing cyanobacteria. Under certain conditions, a red layer of purple sulfur bacteria developed underneath the cyanobacterial mat in which Chromatium and Thiocapsa spp. dominated, but Thiopedia and Ectothiorhodospira spp. have also been observed. Measurements of light penetrating the cyanobacterial mat indicated that sufficient light is available for the photosynthetic growth of purple sulfur bacteria. Profiles of oxygen, sulfide and redox potential within the microbial mat were measured using microelectrodes. Maximum oxygen concentrations, measured at a depth of 0.7 mm, reached levels more than twice the normal air saturation. Dissolved sulfide was not detected by the microelectrodes. Determination of acid-distilled sulfide, however, revealed appreciable amounts of bound sulfide in the mat. Redox profiles measured in the mat led to the conclusion that the upper 10 mm of the sedimentary sequence is in a relatively oxidized state.     

Colonization rates and community structure of diatoms on three different rock substrata in a lotic system

Diatom assemblages were monitored at weekly intervals over a 5 week period on Verde limestone, Supai sandstone, and Andesitic basalt substrata in a mountain stream in northern Arizona, U.S.A. Density, Shannon-Weiner diversity, evenness, and community similarity (SIMI) were used to compare colonization patterns and community structure between individual substratum types. Average standing crop values were nearly two-fold higher on sandstone than on either basalt or limestone substrata after the first week of the study. It is proposed that differences in micro-surface features between substrata and possibly the rate of substratum solubilization may cause these differences in density early in the colonization period. Following the initial week, standing crop and community structure were significantly similar on all substrata for the remainder of the study period. Maximum densities were attained by the third week and remained relatively constant on all substrata for the remainder of the study.

SEM micrographs demonstrated that surfaces of submerged substrata in streams are modified after the first week by the accumulation of organic aggregates. The establishment of an “organic matrix” early in the colonization process may provide relatively similar attachment surfaces for microbial invasion. This appears to reduce the initial microtopographic differences displayed by substrata and allows for a more uniform colonization pattern.     

Modern bryomol-sediments in a cool-water, high-energy setting: the inner shelf off Northern Brittany

Summary The inner shelf off Roscoff/North Brittany, France situated in the Western English Channel, provides a unique facility for studying modern cold water BRYOMOL-carbonate deposits in the North Atlantic. High tidal gauge with extreme strong tidal currents, which effect the well-mixed water column throughout the year, and strong, seasonal storms characterize the conditions of the area. The shelf is seperated in two carbonate production zones: Boulder fields and sand fields. The main production areas are the sand fields with an intensive epifaunal and infaunal colonization by bivalves and the bryozoan thickets ofCellaria spp. and their associated epifauna. The most beneficial and dominant strategies of the sessile benthos are erect flexible and encrusting growth habits. The study area is partitioned in two zones: (1) the coarse sediment blanket typified by active sediment generation and little accumulation, and (2) the shell dune Trezen ar Skoden, characterized by accumulated sediments. Sediment transport and distribution of facies areas are controlled by the strong semidiurnal tidal current regime and episodically severe storms. As a result of these high energy processes of redeposition the autochthonous sediment particles are physically reworked and redeposited while in calmer periods deposition and biological destruction of the components occur. The Holocene development of the sea level has been of crucial influence to the costal morphology and the establishment of different carbonate production centers. The benthic communities produce with their carbonate skeletons the first biogenic sediment and provide substrates for colonization of epi- and endofauna. Changes in the current patterns and the morphology of the sea bottom resulted in the origin of the shell dune Trezen ar Skoden.     

Jurassic corals as emersion indicators

In Recent coral reefs, emersion has resulted in the development of special growth forms of colonial corals which have been named microatolls. These colonies grow almost exclusively in a horizontal direction. In the upper part of the colony, subaerial exposure leads to the decay of the living body. The growth then stops and the skeleton is later colonized by various boring and encrusting organisms. Here we show the first record of a massive Mesozoic coral colony displaying the main features of emersion. The colony has been collected in perireefal Oxfordian (Jurassic) limestones from the Jura Mountains (France). It is situated exactly at the expected place in a shallowing-upward sequence between infralittoral buildups and supralittoral limestones. We suggest that such growth structures could be more common than previously thought in ancient coralliferous sediments and add some new details to discriminate between these colonies.     

Size-related shifts in the habitat associations of young-of-the-year winter flounder (Pseudopleuronectes americanus): field observations and laboratory experiments with sediments and prey

Field surveys and laboratory studies were used to determine the role of substrata in habitat selection by young-of-the year winter flounder. A synoptic field survey of winter flounder and sediments in the Navesink River-Sandy Hook Bay estuarine system in New Jersey demonstrated that winter flounder distribution was related to sediment grain size. Analysis using a generalized additive model indicated that the probability of capturing 10-49 mm SL winter flounder was high on sediments with a mean grain diameter of /=40 mm SL) preferred coarse-grained sediments. Burying ability increased with size and all flounders avoided sediments that prevented burial. Subsequent laboratory experiments revealed that the presence of live prey (Mya arenaria) can over-ride sediment choice by winter flounder (50-68 mm SL) indicating the complexity of interrelated factors in habitat choice.     

Management-free techniques for restoration of Eisenia and Ecklonia beds along the central Pacific coast of Japan

Because of the importance of seaweed resources for conservation of thecoastal environment, the provision of habitats of commercially importantspecies and because of use as human food, a study was carried out on bedsof Eisenia bicyclis and Ecklonia cava. This included the mostimportant environmental factors and the development of techniques forrestoration and long-term maintenance of the beds. At a site chosen fordetailed study, the Eisenia and Ecklonia bed was influenced bymany factors, including desiccation, rainfall, low salinity, low light intensity,turbidity, accumulation of suspended sediments, cover by adhering animals,wave and current action and grazing pressure. The distribution of suchseaweed beds is also limited locally by the size and hardness of rock on thebottom, and by fluctuations in the sand level at the base of the reefs.The management-free technique of creating Eisenia and Ecklonia beds is based on an understanding of the factors limiting the localdistribution of the beds. Conditions for seaweed growth can then beartificially enhanced by coastal engineering. Suggestions to enhance Eisenia and Ecklonia bed restoration using management-freetechniques include raising the bottom, elevating substrata above the sandybottom, and providing substrata of a shape suitable for the attachment ofkelp.     

Benthic algal response to N and P enrichment along a pH gradient

Nutrient enrichment and its effect on benthic algal growth, community composition, and average cell size was assessed across two sites of differing pH within a single habitat. Nutrients were added using in situ substrata, which released either N, P, or no additional nutrients (controls) at each site for 21 days. Upon collection, chlorophyll and biovolume standing stocks of the attached algal microflora were measured. Chlorophyll concentration was different among all treatments, accumulating greatest on P, followed by N, and the least on C substrata (P < 0.001) and was highest at site-2 (P < 0.001), while total algal biovolume was highest on P compared to both N and C substrata (P < 0.05) and did not vary between sites. Increased growth on P substrata was due to the enhanced biovolume of filamentous green algae, although the affected taxa varied between sites. Biovolume to cell density ratios (as a measure of average cell size) were highest on P substrata over both N-enriched and control substrata (P < 0.05) and this pattern was similar between sites. Progression towards a community composed of larger cells following P enrichment observed along this pH gradient, seems to be related to the dominance of larger celled filamentous green algae. Thus, nutrients exhibited greater control on benthic algal growth than did changes in hydrogen ion concentration.Contribution number 581, Great Lakes Environmental Research LaboratoryContribution number 581, Great Lakes Environmental Research Laboratory     

Isolation and enumeration of marine actinomycetes from seawater and sediments in Alexandria

Actinomycetes were isolated from near-shore marine sediments and water at four different sites in Alexandria. Statistical analysis revealed that variation in temperature, pH, and dissolved phosphate were of insignificant values, but that variation in total nitrogen and organic matter were significant. The treatment of sediments and water samples by heat resulted in a selective reduction of the nonactinomycetal heterotrophic microflora. Four selective culture media were used for counting actinomycetes in marine water and sediments. The starch nitrate medium favored the growth of these microorganisms. The diversity and counts of actinomycetes varied with the seasonal variation, and the highest counts were detected in dry warm seasons. The numbers of this bacterial group in sediments exceeded by far their numbers in seawater. A positive correlation was found between population size and location. Actinomycetes were found in the highest numbers in the upper layers (0-20 cm depth). In a few cases, the counts of actinomycetes showed bimodal maxima 0-20 and 60-100 cm deep. Sediments were the best source of marine actinomycetes, and their distribution varied depending on the depth from which samples were collected. The ratio of actinomycetes to the total microflora ranged from 0.48 to 2.29, depending on location.     

Behavioural interactions between ecosystem engineers control community species richness

Behavioural interactions between ecosystem engineers may strongly influence community structure. We tested whether an invasive ecosystem engineer, the alga Caulerpa taxifolia , indirectly facilitated community diversity by modifying the behaviour of a native ecosystem engineer, the clam Anadara trapezia , in southeastern Australia . In this study, clams in Caulerpa -invaded sediments partially unburied themselves, extending >30% of their shell surface above the sediment, providing rare, hard substrata for colonization. Consequently, clams in Caulerpa had significantly higher diversity and abundance of epibiota compared with clams in unvegetated sediments. To isolate the role of clam burial depth from direct habitat influences or differential predation by habitat, we manipulated clam burial depth, predator exposure and habitat ( Caulerpa or unvegetated) in an orthogonal experiment. Burial depth overwhelmingly influenced epibiont species richness and abundance, resulting in a behaviourally mediated facilitation cascade. That Caulerpa controls epibiont communities by altering Anadara burial depths illustrates that even subtle behavioural responses of one ecosystem engineer to another can drive extensive community-wide facilitation.     

The role of colony morphology and substratum inclination in the success of Millepora alcicornis on shallow coral reefs

 Millepora species are conspicuous members of shallow coral reefs where they occupy a variety of substrata and produce morphologically complex skeletons. This study focuses on the roles of growth on vertical and horizontal surfaces and the production of encrusting bases and branches (a “sheet-tree” morphology) for the success of the Millepora alcicornis on coral reefs. The effects of inclination were investigated by comparing the size and growth rates of M. alcicornis on vertical and horizontal surfaces at 3–5 m depth, in St. John, US Virgin Islands. The consequences of morphological complexity were investigated by comparing polyp density, chlorophyll content and biomass between encrusting bases and branches; the role of branches in asexual reproduction was also quantified. Colonies on vertical surfaces had larger encrusting bases, longer perimeters and lower densities of branches compared to those on horizontal surfaces. Growth rates also varied significantly between surfaces, largely because colonies on horizontal surfaces shrank in area while those on vertical surfaces increased in area, albeit slowly. Branches were not specialized in comparison to encrusting bases in terms of the density of dactylozooids and gastrozooids, chlorophyll content and biomass, but they were effective asexual propagules. During one storm, 79% of the branches were removed from colonies of M. alcicornis, and 4% attached to the substratum to produce new colonies at a density of ≈0.5 colonies.m^-2. Anecdotal observations suggest that such storms rarely damaged encrusting bases on vertical surfaces, but often destroyed those on horizontal surfaces. Thus, the encrusting bases on vertical surfaces are likely to be large because of greater age rather than faster growth, while those on horizontal surfaces are likely to be small because they are relatively young and short lived. These findings suggest that the success of M. alcicornis is a result, in part, of the beneficial consequences of their “sheet-tree” morphology, that supports: (a) slow growth and resistance to wave damage of encrusting bases on vertical surfaces, and (b) the use of branches as asexual propagules. Accepted: 24 November 1998     

DIATOM COLONIZATION ON ARTIFICIAL SUBSTRATA IN POOL AND RIFFLE ZONES STUDIED BY LIGHT AND SCANNING ELECTRON MICROSCOPY1

Light and scanning electron microscopy were utilized to quality diatom colonization in Oak Creek, Arizona. Aluminum SEM stubs with and without plexiglass discs were anchored into rocks. Early colonization on five stub microzones was examined at hourly intervals; weekly intervals of up to 3 wk were employed to record community development in pool find riffle. Plexiglass was more suitable for microbial colonization than aluminum. Organic matter and bacteria were important surface pre-conditioning agents while fungi were instrumental in trap/ting cells during early stages of colonization in the riffle. Diatom colonization was initialed within 1 h on the upstream side of substrata in riffles, while the tap face was colonized first in pools. Colonization moved rapidly to the perimeter in each system. Early colonization of-side microzones was considerably more asymmetric in the riffle than, pool. At Idler stages (2 wk) diatoms with their associated mucilage and algal filaments contributed to the stability of the microbial communities. Horizontally positioned species (Achnanthes, Cocconeis) were early colonizers in both systems while vertically positioned species (Gomphonema, Nitzschia) were more important in later successional stages (3 wk) in the riffle. Horizontally positioned species remained dominant throughout the 3 wk period in the pool. After 3 wk, diversity was normally greater in the pool while density was higher in the riffle. Detrital microcosms containing viable microbiol assemblages frequently collected on tin-upstream face of substrata in the riffle. The random nature by which these detrital microcoms contact downstream substrata greatly contribute to the spatial variation of periphyton in streams. These detrital microcosms expedite repeated colonization in lotic systems.