Surface-dependent strategies and energy flux in benthic marine communities or, why corals do not exist in the Mediterranean

Most structure-building organisms in rocky benthic communities are surface-dependent because their energy inputs depend mainly on the surface they expose to water. Two photosynthetic strategies, divided into calcareous and non calcareous algae, strict suspension-feeders and photosynthetic suspension feeders (e.g. hermatypic corals) are the four main strategies evolutively acquired by benthic organisms. Competition between those strategies occur in relation to productivity of the different species, in such a way that, for given environmental conditions, species with a higher growth (P/B ratio) would dominate. At a worldwide scale, littoral marine benthos can he considered to fit into the four fields defined by two main axes: the first, relates to productivity and relies atrophic and oligotrophic waters and the second is defined by the degree of environmental variability or seasonality (from high to low). Coral reefs (marine ecosystems dominated by photosynthetic suspension feeders) develop in the space of oligotrophic areas with low variability, while kelp beds (marine ecosystem dominated by large, non calcareous algae) are to be found only in eutrophic places with a high variability. The space of eutrophic waters with a low variability do not has specially adapted, high structured, benthic marine ecosystems, and in these conditions opportunistic algae and animals predominate. Finally, photophilic mediterranean benthos -devoid of kelps and without hermatypic corals- typifies the field of oligotrophic areas with high variability; in its more genuine aspect, Mediterranean benthos is represented by small algae with a high percentage of calcareous thallii. In all cases strict suspension-feeders compete successfully with photosynthetic organisms only in situations of low irradiances or very high inputs of POM. In its turn, Mediterranean rocky benthos, in spite of its relative uniformity, is geographically organized along the same axes. The Gulf of Lions and the insular bottoms (Balearic Islands, for example) would correspond to the extremes of eutrophic-high variability areas and oligotrophic-low variability areas, respectively. Irradiance, nutrient and POM concentration, and hydrodynamism are the three variables which mainly affect the distribution of the different surface-dependent strategies, and thus, these parameters are of paramount interest for understanding the trophic structure of Mediterranean benthic communities. In environments non limited by light, nutrient availability, defined as the product between nutrient -POM concentration and hydrodynamism, states the dominance of calcareous versus non calcareous algae. Calcareous algae dominate in oligotrophic waters while non-calcareous algae dominate in moderately eutrophic waters. In light-limited environments, passive suspension feeders (octocorallaria, gorgonians) become dominant species if POM availability is enhanced by a high hydrodynamism (strong currents); in waters with a low charge of POM organisms of other groups, mainly active suspension feeders, predominate (sponges, bryozoans, scleractiniarians). In any case, there always exists a very variable bathymetric zone, depending on light attenuation and nutrient-POM availability, where encrusting calcareous algae strongly compete with suspension feeders (coralligenous).

Response of the benthic macroinvertebrate community to a point source in La Tordera stream (Catalonia, NE Spain)

Vam monitoritzar paràmetres físics i químics, macroinvertebrats bentònics, clorofil·la a, productors primaris i matèria orgànica durant un any (2001-2002) per examinar els efectes d'una font puntual sobre la composició taxonòmica, la estructura de la comunitat, l'organització funcional, la utilització de l'habitat i la estoquiometria al riu la Tordera (Catalunya). Aigües avall de la font puntual, concentració de nutrients, cabal i conductivitat eren majors que al tram d'aigües amunt, mentre que oxigen dissolt era menor. La densitat de macroinvertebrats era més elevada al tram d'aigües avall però la biomassa era similar als dos trams. La riquesa taxonòmica al tram de dalt era un 20% més alt que al tram de baix. Els anàlisis d'ordenació separen clarament els dos trams en el primer eix, mentre que els dos trams presentaven una pauta temporal similar en el segon eix. La similaritat entre els dos trams en composició taxonòmica, densitats i biomasses després de les crescudes d'abril i maig de 2002, indiquen que les pertorbacions del cabal poden actuar com a un mecanisme de reinici de la comunitat bentònica i jugar un paper important per a la restauració d'ecosistemes fluvials. Els dos trams presentaven una biomassa de perifiton, plantes vasculars, CPOM i FPOM similars, mentre que clorofil·la a, algues filamentoses, molses i SPOM eren majors al tram d'aigües avall. La densitat relativa de trituradors era menor sota la font puntual mentre que col·lectors i filtradors van ser afavorits. La biomassa relativa de trituradors també era menor sota la font puntual, però la biomassa de col·lectors i depredadors va augmentar. Les relacions entre densitat de grups tròfics i els seus recursos eren rarament significatives. La relació s'explicava millor amb la biomassa de macroinvertebrats. Els dos trams compartien la mateixa relació per raspadors, col·lectors i filtradors però no per trituradors i depredadors. La densitat i la biomassa de macroinvertebrats es trobaven positivament correlacionades amb la quantitat de recursos tròfics i la complexitat d'habitat, mentre que la riquesa taxonòmica es trobava negativament relacionada amb paràmetres hidràulics. La influència dels substrats inorgànics prenia menor rellevància per a la distribució dels macroinvertebrats. Els anàlisis d'ordenació mostren com les variables del microhabitat de major rellevància eren CPOM, clorofil·la a, algues filamentoses i velocitat. La cobertura de sorra només era significativa per al tram d'aigües amunt i les molses, al d'aigües avall. El número de correlacions significatives entre macroinvertebrats i les variables del microhabitat era més elevat per al tram de dalt que per al de baix, bàsicament per diferències en composició taxonòmica. La biomassa de macroinvertebrats va aportar una informació semblant a la obtinguda per la densitat. Perifiton i molses tenien uns continguts de nutrients similars en els dos trams. Els %C i %N d'algues filamentoses també eren similars en els dos trams però el %P sota la font puntual era el doble que al tram de dalt. Les relacions estoquiomètriques en CPOM, FPOM i SPOM eren considerablement menors sota la font puntual. Els continguts elementals i relacions van ser molt variables entre taxons de macroinvertebrats però no van resultar significativament diferents entre els dos trams. Dípters, tricòpters i efemeròpters presentaven una estoquiometria similar, mentre que el C i el N eren inferiors en moluscs i el P en coleòpters. Els depredadors presentaven un contingut en C i N més elevat que la resta de grups tròfics, mentre que el P era major en els filtradors. Els desequilibris elementals entre consumidors i recursos eren menors en el tram d'aigües avall. A la tardor i l'hivern la major font de nutrients va ser la BOM mentre que a la primavera i a l'estiu va ser el perifiton.

Calcareous algae from Upper Albian - Cenomanian strata of the Potiguar basin (NE Brazil)

Calcareous Albian-Cenomanian strata in the Potiguar basin yield a number of microfossils, among which are calcareous algae (Dasycladales and Corallinales). This data, together with an inventory of previous discoveries, enabled the discussion of their biogeography. These benthic organisms probably came from the early Central Atlantic Ocean through its northwestern neck between Africa and South America and thus reached the early South Atlantic platforms.

New drugs with antiprotozoal activity from marine algae: a review

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Trophic structure on the West Antarctic Peninsula shelf: Detritivory and benthic inertia revealed by delta(13)C and delta(15)N analysis

Summer bloom-derived phytodetritus settles rapidly to the seafloor on the West Antarctic Peninsula (WAP) continental shelf, where it appears to degrade relatively slowly, forming a sediment ""food bank"" for benthic detritivores. We used stable carbon and nitrogen isotopes to examine sources and sinks of particulate organic material (POM) reaching the WAP shelf benthos (550-625 m depths), and to explore trophic linkages among the most abundant benthic megafauna. We measured delta(13)C and delta(15)N values in major megafaunal taxa (n = 26) and potential food sources, including suspended and sinking POM, ice algae, sediment organic carbon, phytodetritus, and macrofaunal polychaetes. The range in delta(13)C values (> 14 parts per thousand) of suspended POM was considerably broader than in sedimentary POC, where little temporal variability in stable isotope signatures was observed. While benthic megafauna also exhibited a broad range of VC values, organic carbon entering the benthic food web appeared to be derived primarily from phytoplankton production, with little input from ice algae. One group of organisms, primarily deposit-feeders, appeared to rely on fresh phytodetritus recovered from the sediments, and sediment organic material that had been reworked by sediment microbes. A second group of animals, including many mobile invertebrate and fish predators, appeared to utilize epibenthic or pelagic food resources such as zooplankton. One surface-deposit-feeding holothurian (Protelpidia murrayi) exhibited seasonal variability in stable isotope values of body tissue, while other surface- and subsurface-deposit-feeders showed no evidence of seasonal variability in food source or trophic position. Detritus from phytoplankton blooms appears to be the primary source of organic material for the detritivorous benthos; however, seasonal variability in the supply of this material is not mirrored in the sediments, and only to a minor degree in the benthic fauna. This pattern suggests substantial inertia in benthic-pelagic coupling, whereby the sediment ecosystem integrates long-term variability in production processes in the water column above. Published by Elsevier Ltd.

Deposit-feeding in benthic macrofauna : Tracer studies from the Baltic Sea

A low content of organic matter, which is largely refractory in nature, is characteristic of most sediments, meaning that aquatic deposit-feeders live on a very poor food source. The food is derived mainly from sedimenting phytodetritus, and in temperate waters like the Baltic Sea, from seasonal phytoplankton blooms. Deposit-feeders are either bulk-feeders, or selective feeders, which preferentially ingest the more organic-rich particles in the sediment, including phytodetritus, microbes and meiofauna. The soft-bottom benthos of the Baltic Sea has low species biodiversity and is dominated by a few macrobenthic species, among which the most numerous are the two deposit-feeding amphipods Monoporeia affinis and Pontoporeia femorata, and the bivalve Macoma balthica. This thesis is based on laboratory experiments on the feeding of these three species, and on the priapulid Halicryptus spinulosus. Feeding by benthic animals is often difficult to observe, but can be effectively studied by the use of tracers. Here we used the radioactive isotope 14C to label food items and to trace the organic matter uptake in the animals, while the stable isotopes 13C and 15N were used to follow feeding on aged organic matter in the sediment. The abundance of M. balthica and the amphipods tends to be negatively correlated, i.e., fewer bivalves are found at sites with dense populations of amphipods, with the known explanation that newly settled M. balthica spat are killed by the amphipods. Whether the postlarvae are just accidentally killed, or also ingested after being killed was tested by labelling the postlarvae with 14C and Rhodamine B. Both tracer techniques gave similar evidence for predation on and ingestion of postlarval bivalves. We calculated that this predation was likely to supply less than one percent of the daily carbon requirement for M. affinis, but might nevertheless be an important factor limiting recruitment of M. balthica. The two amphipods M. affinis and P. femorata are partly vertically segregated in the sediment, but whether they also feed at different depths was unknown. By adding fresh 14C-labelled algae either on the sediment surface or mixed into the sediment, we were able to distinguish surface from subsurface feeding. We found M. affinis and P. femorata to be surface and subsurface deposit-feeders, respectively. Whether the amphipods also feed on old organic matter, was studied by adding fresh 14C-labelled algae on the sediment surface, and using aged, one-year-old 13C- and 15N-labelled sediment as deep sediment. Ingestion of old organic matter, traced by the stable isotopes, differed between the two species, with a higher uptake for P. femorata, suggesting that P. femorata utilises the older, deeper-buried organic matter to a greater extent. Feeding studies with juveniles of both M. affinis and P. femorata had not been done previously. In an experiment with the same procedure and treatments as for the adults, juveniles of both amphipod species were found to have similar feeding strategies. They fed on both fresh and old sediment, with no partitioning of food resources, making them likely to be competitors for the same food resource. Oxygen deficiency has become more wide-spread in the Baltic Sea proper in the last half-century, and upwards of 70 000km2 are now devoid of macrofauna, even though part of that area does not have oxygen concentrations low enough to directly kill the macrofauna. We made week-long experiments on the rate of feeding on 14C-labelled diatoms spread on the sediment surface in different oxygen concentrations for both the amphipod species, M. balthica and H. spinulosus. The amphipods were the most sensitive to oxygen deficiency and showed reduced feeding and lower survival at low oxygen concentrations. M. balthica showed reduced feeding at the lowest oxygen concentration, but no mortality increase. The survival of H. spinulosus was unaffected, but it did not feed, showing that it is not a surface deposit-feeder. We conclude that low oxygen concentrations that are not directly lethal, but reduce food intake, may lead to starvation and death in the longer term.

Effects of Paramuricea clavata (Risso, 1826) (Anthozoa: Plexauridae) forests on settlement of benthic species: an experimental approach

Rationale: Coralligenous habitat is considered the second most important subtidal “hot spot” of species diversity in the Mediterranean Sea after the Posidonia oceanica meadows. It can be defined as a typical Mediterranean biogenic hard bottom, mainly produced by the accumulation of calcareous encrusting algae that, together with other builder organisms, form a multidimensional framework with a high micro-spatial variability. The development of this habitat depends on physical factors (i.e. light, hydrodynamism, nutrients, etc.), but also biologic interactions can play a relevant role in structuring the benthic assemblages. This great environmental heterogeneity allows several different assemblages to coexist in a reduced space. One of the most beautiful is that characterised by the Mediterranean gorgonian Paramuricea clavata (Risso, 1826) that can contribute to above 40% of total biomass of the community and brings significant structural complexity into the coralligenous habitat. In sites moderately exposed to waves and currents, P. clavata can form high-density populations (up to 60 colonies m-2) between 20 – 70 m in depth. Being a suspension feeder, where it forms dense populations, P. clavata plays a significant role in transferring energy from planktonic to benthic system. The effects of the branched colonies of P. clavata could be comparable to those of the forests on land. They can affect the micro scale hydrodynamism and light, promoting or inhibiting the growth of other species. Unfortunately, gorgonians are threatened by several anthropogenic disturbance factors (i.e. fishing, pollution, tourism) and by climatic anomalies, linked to the global changes, that are responsible of thermal stress, development of mucilage and enhanced pathogens activity, leading to mass mortality events in last decades. Till now, the possible effects of gorgonian forest loss are largely unknown. Our goal was to analyse the ecological role of these sea fan forests on the coralligenous benthic assemblages. Experimental setup and main results: The influence of P. clavata in the settlement and recruitment of epibenthic organisms was analysed by a field experiment carried out in two randomly selected places: Tavolara island and Portofino promontory. The experiment consisted in recreate the presence and absence of the gorgonian forest on recruitment panels, arranged in four plots per type (forested and non-forested), interspersed each other, and deployed at the same depth. On every forested panel 3 gorgonian colonies about 20 cm height were grafted with the use of Eppendorf tubes and epoxy resin bicomponent simulating a density of 190 sea fans per m-2. This density corresponds to a mean biomass of 825 g DW m-2,3 which is of the same order of magnitude of the natural high-density populations. After about 4 months, the panels were collected and analysed in laboratory in order to estimate the percent cover of all the species that have colonized the substrata. The gorgonian forest effects were tested by multivariate and univariate permutational analyses of the variance (PERMANOVA). Recruited assemblages largely differed between the two study sites, probably due to different environmental conditions including water quality and turbidity. On overall, the presence of P. clavata reduced the settlement and recruitment of several algae: the shadow caused by the gorgonian might reduce light availability and therefore their growth. This effect might be greater in places where the waters are on average more clear, since at Portofino it is less visible and could be masked by the high turbidity of the water. The same pattern was registered for forams, more abundant outside gorgonian forest, probably linked with algal distribution, shadowing effect or alimentary competition. The last one hypothesis could be valid also for serpulids polychaetes that growth mainly on non-forested panels. An opposite trend, was showed by a species of bryozoan and by an hydroid that is facilitated by the presence of P. clavata, probably because it attenuates irradiance level and hydrodynamism. Species diversity was significantly reduced by the presence of P. clavata forests at both sites. This seems in contrast with what we expected, but the result may be influenced by the large algal component on non-forested panels. The analysis confirmed the presence of differences in the species diversity among plots and between sites respectively due to natural high variability of the coralligenous system and to different local environment conditions. The reduction of species diversity due to the presence of gorgonians appeared related to a worst evenness rather than to less species richness. With our experiment it is demonstrated that the presence of P. clavata forests can significantly alter local coralligenous assemblages patterns, promoting or inhibiting the recruitment of some species, modifying trophic relationships and adding heterogeneity and complexity to the habitat. Moreover, P. clavata could have a stabilising effect on the coralligenous assemblages.

Harmful algae and their potential impacts on the coastal ecosystem: growth and toxin production dynamics

The main goal of the present thesis was to study some harmful algal species which cause blooms in Italian coastal waters, leading to consequences for human health, coastal ecosystem, fishery and tourism. In particular, in the first part of this thesis the toxicity of Adriatic strains of the raphidophyte Fibrocapsa japonica was investigated. Despite several hypotheses have been proposed for the toxic mechanism of the raphidophytes, especially for the species Chattonella antiqua and C. marina, which have been studied more extensively, just a few studies on the toxic effects of these species for different organisms were reported. Moreover, a careful reading of the literature evidenced as any ichthyotoxic events reported worldwide can be linked to F. japonica blooms. Although recently several studies were performed on F. japonica strains from the USA, Japan, Australia, New Zealand, the Netherlands, Germany, and France in order to characterize their growth and toxicity features, the work reported in this thesis results one of the first investigation on the toxic effects of F. japonica for different organisms, such as bacteria, crustaceans and fish. Mortality effects, together with haemolysis of fish erythrocytes, probably due to the relatively high amount of PUFAs produced by this species, were observed. Mortality for fish, however, was reported only at a high cell density and after a long exposition period (9-10 days); moreover a significant increase of H2O2 obtained in the tanks where sea basses were exposed to F. japonica was also relevant. This result may justify the absence of ichthyotoxic events in the Italian coasts, despite F. japonica blooms detected in these areas were characterized by high cell densities. This work reports also a first complete characterization of the fatty acids produced and extracellularly released by the Adriatic F. japonica, and results were also compared with the fatty acid profile of other strains. The absence of known brevetoxins in F. japonica algal extracts was also highlighted, leading to the hypothesis that the toxicity of F. japonica may be due to a synergic effect of PUFAs and ROS. Another microalgae that was studied in this thesis is the benthic dinoflagellate Ostreopsis cf. ovata. This species was investigated with the aim to investigate the effect of environmental parameters on its growth and toxicity. O. cf. ovata, in fact, shows different blooming periods along the Italian coasts and even the reported toxic effects are variable. The results of this work confirmed the high variability in the growth dynamic and toxin content of several Italian strains which were isolated in recent years along the Adriatic and Tyrrhenian Seas. Moreover, the effects of temperature and salinity on the behaviour of the different isolates are in good agreement with the results obtained from field surveys, which evidence as the environmental parameters are important factors modulating O. cf. ovata proliferation. Another relevant result that was highlighted is the anomaly in the production of palytoxin-like compounds reported by one of the studied isolate, in particular the one isolated in 2008 in Ancona (Adriatic Sea). Only this strain reported the absence of two (ovatoxin-b and –c) of the five ovatoxins so far known in the toxin profile and a different relative abundance of the other toxins. The last aspect that was studied in this thesis regards the toxin biosythesis. In fact, toxins produced (palytoxin-like compounds) or supposed to be produced (brevetoxin-like compounds) by O. cf. ovata and F. japonica, respectively, are polyketides, which are highly oxygenated compounds synthesized by complex enzymes known as polyketide synthase (PKS) enzymes. These enzymes are multi-domain complexes that structurally and functionally resemble the fatty acid synthases (FASs). This work reports the first study of PKS proteins in the dinoflagellates O. cf. ovata, C. monotis and in the raphidophyte F. japonica. For the first time some PKSs were identified in these species, confirming the presence of PKS proteins predicted by the in silico translation of the transcripts found in K. brevis also in other species. The identification of O. cf. ovata PKSs and the localization of the palytoxin-like compounds produced by this dinoflagellate in a similar location (chloroplast) as that observed for other dinoflagellate and cyanobacterial toxins provides some indication that these proteins may be involved in polyketide biosynthesis. However, their potential function as fatty acid synthases cannot be ruled out, as plant fatty acid synthesis also occurs within chloroplasts. This last hypothesis is also supported by the fact that in all the investigated species, and in particular in F. japonica, PKS proteins were present. Therefore, these results provide an important contribution to the study of the polyketides and of the involvement of PKS proteins in the toxin biosynthesis.