Natural patterns and anthropogenic disturbance in north Adriatic marine benthic assemblages: descriptive and methodological studies

Marine soft bottom systems show a high variability across multiple spatial and temporal scales. Both natural and anthropogenic sources of disturbance act together in affecting benthic sedimentary characteristics and species distribution. The description of such spatial variability is required to understand the ecological processes behind them. However, in order to have a better estimate of spatial patterns, methods that take into account the complexity of the sedimentary system are required. This PhD thesis aims to give a significant contribution both in improving the methodological approaches to the study of biological variability in soft bottom habitats and in increasing the knowledge of the effect that different process (both natural and anthropogenic) could have on the benthic communities of a large area in the North Adriatic Sea. Beta diversity is a measure of the variability in species composition, and Whittaker’s index has become the most widely used measure of beta-diversity. However, application of the Whittaker index to soft bottom assemblages of the Adriatic Sea highlighted its sensitivity to rare species (species recorded in a single sample). This over-weighting of rare species induces biased estimates of the heterogeneity, thus it becomes difficult to compare assemblages containing a high proportion of rare species. In benthic communities, the unusual large number of rare species is frequently attributed to a combination of sampling errors and insufficient sampling effort. In order to reduce the influence of rare species on the measure of beta diversity, I have developed an alternative index based on simple probabilistic considerations. It turns out that this probability index is an ordinary Michaelis-Menten transformation of Whittaker's index but behaves more favourably when species heterogeneity increases. The suggested index therefore seems appropriate when comparing patterns of complexity in marine benthic assemblages. Although the new index makes an important contribution to the study of biodiversity in sedimentary environment, it remains to be seen which processes, and at what scales, influence benthic patterns. The ability to predict the effects of ecological phenomena on benthic fauna highly depends on both spatial and temporal scales of variation. Once defined, implicitly or explicitly, these scales influence the questions asked, the methodological approaches and the interpretation of results. Problem often arise when representative samples are not taken and results are over-generalized, as can happen when results from small-scale experiments are used for resource planning and management. Such issues, although globally recognized, are far from been resolved in the North Adriatic Sea. This area is potentially affected by both natural (e.g. river inflow, eutrophication) and anthropogenic (e.g. gas extraction, fish-trawling) sources of disturbance. Although few studies in this area aimed at understanding which of these processes mainly affect macrobenthos, these have been conducted at a small spatial scale, as they were designated to examine local changes in benthic communities or particular species. However, in order to better describe all the putative processes occurring in the entire area, a high sampling effort performed at a large spatial scale is required. The sedimentary environment of the western part of the Adriatic Sea was extensively studied in this thesis. I have described, in detail, spatial patterns both in terms of sedimentary characteristics and macrobenthic organisms and have suggested putative processes (natural or of human origin) that might affect the benthic environment of the entire area. In particular I have examined the effect of off shore gas platforms on benthic diversity and tested their effect over a background of natural spatial variability. The results obtained suggest that natural processes in the North Adriatic such as river outflow and euthrophication show an inter-annual variability that might have important consequences on benthic assemblages, affecting for example their spatial pattern moving away from the coast and along a North to South gradient. Depth-related factors, such as food supply, light, temperature and salinity play an important role in explaining large scale benthic spatial variability (i.e., affecting both the abundance patterns and beta diversity). Nonetheless, more locally, effects probably related to an organic enrichment or pollution from Po river input has been observed. All these processes, together with few human-induced sources of variability (e.g. fishing disturbance), have a higher effect on macrofauna distribution than any effect related to the presence of gas platforms. The main effect of gas platforms is restricted mainly to small spatial scales and related to a change in habitat complexity due to a natural dislodgement or structure cleaning of mussels that colonize their legs. The accumulation of mussels on the sediment reasonably affects benthic infauna composition. All the components of the study presented in this thesis highlight the need to carefully consider methodological aspects related to the study of sedimentary habitats. With particular regards to the North Adriatic Sea, a multi-scale analysis along natural and anthopogenic gradients was useful for detecting the influence of all the processes affecting the sedimentary environment. In the future, applying a similar approach may lead to an unambiguous assessment of the state of the benthic community in the North Adriatic Sea. Such assessment may be useful in understanding if any anthropogenic source of disturbance has a negative effect on the marine environment, and if so, planning sustainable strategies for a proper management of the affected area.

A comprehensive assessment of environmental ultraviolet radiation induced DNA damage in marine microorganisms

There is evidence that ultraviolet radiation (UVR) is increasing over certain locations on the Earth's surface. Of primary concern is the annual pattern of ozone depletion over Antarctica and the Southern Ocean. Reduction of ozone concentration selectively limits absorption of solar UV-B (290–320 nm), resulting in higher irradiance at the Earth's surface. The effects of ozone depletion on the human population and natural ecosystems, particularly the marine environment, are a matter of considerable concern. Indeed, marine plankton may serve as sensitive indicators of ozone depletion and UV-B fluctuations. Direct biological effects of UVR result from absorption of UV-B by DNA. Once absorbed, energy is dissipated by a variety of pathways, including covalent chemical reactions leading to the formation of photoproducts. The major types of photoproduct formed are cyclobutyl pyrimidine dimer (CPD) and pyrimidine(6-4)pyrimidone dimer [(6-4)PD]. Marine plankton repair these photoproducts using light-dependent photoenzymatic repair or nucleotide excision repair. The studies here show that fluctuations in CPD concentrations in the marine environment at Palmer Station, Antarctica correlate well with ozone concentration and UV-B irradiance at the Earth's surface. A comparison of photoproduct levels in marine plankton and DNA dosimeters show that bacterioplankton display higher resistance to solar UVR than phytoplankton in an ozone depleted environment. DNA damage in marine microorganisms was investigated during two separate latitudinal transects which covered a total range of 140°. We observed the same pattern of change in DNA damage levels in dosimeters and marine plankton as measured using two distinct quantitative techniques. Results from the transects show that differences in photosensitivity exist in marine plankton collected under varying UVR environments. Laboratory studies of Antarctic bacterial isolates confirm that marine bacterioplankton possess differences in survival, DNA damage induction, and repair following exposure to UVR. Results from DNA damage measurements during ozone season, along a latitudinal gradient, and in marine bacterial isolates suggest that changes in environmental UVR correlate with changes in UV-B induced DNA damage in marine microorganisms. Differences in the ability to tolerate UVR stress under different environmental conditions may determine the composition of the microbial communities inhabiting those environments. ^

Surface sediment analyses, bivalve and gastropod taxa of marine sediments from the Banc d'Arguin, Mauritania

High-nutrient tropical carbonate systems are known to produce sediments that, in terms of skeletal composition, are reminiscent of their extra-tropical counterparts. Such carbonate systems and associated carbonate grain assemblages in the tropics are rare in the present-day world. Nonetheless, it is crucial to gain a better understanding of those ecosystems, including their drivers and players because such settings potentially represent models for ancient depositional systems as well as for predicted future environmental conditions. One of the modern occurrences of eutrophic tropical carbonate systems is the northern Mauritanian Shelf. The marine environment is characterized by an eastern boundary upwelling system that pushes cool and nutrient-rich intermediate waters onto a wide epicontinental platform (Golfe d'Arguin) where the waters warm up to tropical temperatures. The resulting facies is mixed carbonate-siliciclastic with a dominant foramol association grading into bimol and barnamol grain assemblages in the shallowest areas forming the Banc d'Arguin. Besides this cool water-related heterozoan association, the carbonate sediment is characterized by tropical molluskan species, while chlorozoan biota (e.g., corals and algal symbiont-bearing foraminifers) are entirely absent. We here present a first comprehensive facies analysis of this model example of eutrophic tropical carbonates. Furthermore, we reconstruct the loci of carbonate production and provide a conclusive depositional model of the Banc d'Arguin that received little attention to date due to its poorly accessible nature.

Highly dealkylated copper and nickel etioporpyfrins in marine sediments

Copper porphyrins have been recognized as natural constituents of marine sediments only within the past 5 years (Palmer and Baker, 1978, Science201, 49-51). In that report it was suggested that these pigments may derive from and be markers for oxidized terrestrial organic matter redeposited in the marine environment. In the present study we describe the distribution of copper porphyrins in sediments from several north Pacific and Gulf of California DSDP/IPQD sites (Legs 56,63,64). These allochthonous pigments have now been found to be accompanied by identical arrays of highly dealkylated nickel etioporphyrins. Evaluation of data from this and past studies clearly reveals that there is a strong carbon-number distribution similarity betweeen coincident Cu and Ni etioporphyrins. This homology match is taken as reflecting a common source for the tetrapyrrole ligands of this population of Cu and Ni chelates. Predepositional generation of these highly dealkylated etioporphyrins is concluded from the occurrence of these pigments in sediments continuing essentially all stages of in situ chlorophyll diagenesis (cf. Baker and Louda, 1983). That is, their presence is not regulated by the in situ diagenetic continuum. Thus, the highly dealkylated Cu and Ni etioporphyrins represent an 'allochthonous' background over which 'autochthonous' (viz. marine produced) chlorophyll derivatives are deposited and are undergoing in situ diagenesis.

Distribution of marine invertebrate larvae in the southern Kara Sea (Russian Arctic)

Within the generally oligotroph Arctic marine environment river outlets are favoured by many planktonic and benthic organisms due to their high input of organic carbon. The retention of pelagic larvae within nursery grounds and/or the ability to return to their parental grounds prior to settlement is one important factor for the persistence of benthic communities in such river influenced areas. The southern Kara Sea is strongly controlled by high freshwater inputs from the Ob and Yenisei Rivers, which create a pronounced bi-layered pycnocline with a warm fresh/brackish water layer on top and a cold high saline marine layer below. The dispersal of five meroplanktonic species and settled juveniles (the brittle star Ophiocten sericeum, and the polychaetes Micronephtys minuta, Nereimyra aphroditoides, Phyllodoce groenlandica and Prionospio cirrifera) in relation to the adult distribution patterns was investigated. For all apart from P. cirrifera the highest densities of larvae were found in the upper brackish water layer. To assess size-at-settlement, the body sizes of larvae and newly settled juveniles were estimated and compared. Dispersal patterns ranged from virtually no adaption to river run-off as in the common, stenohaline O. sericeum and M. minuta (7 ind./m**3, 459 µm) to local retention as in N. aphroditoides (7 ind./m**3, 541 µm) and P. groenlandica (0.5 ind./m**3, 1121 µm) retained by horizontal eddies created by the outflow. Adults of P. cirrifera, which were exclusively restricted to the estuary of the Yenisei River, showed a well adapted reproductive behaviour to ensure a high retention potential of their progenies. The larvae (1.5 ind./m**3, 1513 µm) were only present in the lower water layers, most probably taking advantage of the prevailing near bottom counter current retaining them within their hatching areas.

Palynology, source vegetation, environment and age of ODP Site 188-1166 sediments

Twenty-three core catcher samples from Site 1166 (Hole 1166A) in Prydz Bay were analyzed for their palynomorph content, with the aims of determining the ages of the sequence penetrated, providing information on the vegetation of the Antarctic continent at this time, and determining the environments under which deposition occurred. Dinocysts, pollen and spores, and foraminiferal test linings were recovered from most samples in the interval from 142.5 to 362.03 meters below seafloor (mbsf). The interval from 142.5 to 258.72 mbsf yielded palynomorphs indicative of a middle-late Eocene age, equivalent to the lower-middle Nothofagidites asperus Zone of the Gippsland Basin of southeastern Australia. The Prydz Bay sequence represents the first well-dated section of this age from East Antarctica. Dinocysts belonging to the widespread "Transantarctic Flora" give a more confident late Eocene age for the interval 142.5-220.5 mbsf. The uppermost two cores within this interval, namely, those from 142.5 and 148.36 mbsf, show significantly higher frequencies of dinocysts than the cores below and suggest that an open marine environment prevailed at the time of deposition. The spore and pollen component may reflect a vegetation akin to the modern rainforest scrubs of Tasmania and New Zealand. Below 267 mbsf, sparse microfloras, mainly of spores and pollen, are equated with the Phyllocladidites mawsonii Zone of southeastern Australia, which is of Turonian to possibly Santonian age. Fluvial to marginal marine environments of deposition are suggested. The parent vegetation from this interval is here described as "Austral Conifer Woodland." The same Late Cretaceous microflora occurs in two of the cores above the postulated unconformity at 267 mbsf. In the core at 249.42 mbsf, the Late Cretaceous spores and pollen are uncontaminated by any Tertiary forms, suggesting that a clast of this older material has been sampled; such a clast may reflect transport by ice during the Eocene. At 258.72 mbsf, Late Cretaceous spores and pollen appear to have been recycled into the Eocene sediments.

Major, trace element and XRF data for sediments and fish tooth samples, from a shallow marine setting with varying redox conditions (Site U1356, Wilkes Land Margin, E.Antarctica)

Fossil fish teeth from pelagic open ocean settings are considered a robust archive for preserving the neodymium (Nd) isotopic composition of ancient seawater. However, using fossil fish teeth as an archive to reconstruct seawater Nd isotopic compositions in different sedimentary redox environments and in terrigenous-dominated, shallow marine settings is less proven. To address these uncertainties, fish tooth and sediment samples from a middle Eocene section deposited proximal to the East Antarctic margin at Integrated Ocean Drilling Program Site U1356 were analyzed for major and trace element geochemistry, and Nd isotopes. Major and trace element analyses of the sediments reveal changing redox conditions throughout deposition in a shallow marine environment. However, variations in the Nd isotopic composition and rare earth element (REE) patterns of the associated fish teeth do not correspond to redox changes in the sediments. REE patterns in fish teeth at Site U1356 carry a typical mid-REE-enriched signature. However, a consistently positive Ce anomaly marks a deviation from a pure authigenic origin of REEs to the fish tooth. Neodymium isotopic compositions of cleaned and uncleaned fish teeth fall between modern seawater and local sediments and hence could be authigenic in nature, but could also be influenced by sedimentary fluxes. We conclude that the fossil fish tooth Nd isotope proxy is not sensitive to moderate changes in pore water oxygenation. However, combined studies on sediments, pore waters, fish teeth and seawater are needed to fully understand processes driving the reconstructed signature from shallow marine sections in proximity to continental sources. This article is protected by copyright. All rights reserved.

Effects of CO2-driven ocean acidification on early life stages of marine medaka (Oryzias melastigma)

The potential effects of elevated CO2 level and reduced carbonate saturation state in marine environment on fishes and other non-calcified organisms are still poorly known. In present study, we investigated the effects of ocean acidification on embryogenesis and organogenesis of newly hatched larvae of marine medaka (Oryzias melastigma) after 21 d exposure of eggs to different artificially acidified seawater (pH 7.6 and 7.2, respectively), and compared with those in control group (pH 8.2). Results showed that CO2-driven seawater acidification (pH 7.6 and 7.2) had no detectable effect on hatching time, hatching rate, and heart rate of embryos. However, the deformity rate of larvae in pH 7.2 treatment was significantly higher than that in control treatment. The left and right sagitta areas did not differ significantly from each other in each treatment. However, the mean sagitta area of larvae in pH 7.6 treatment was significantly smaller than that in the control (p = 0.024). These results suggest that although marine medaka might be more tolerant of elevated CO2 than some other fishes, the effect of elevated CO2 level on the calcification of otolith is likely to be the most susceptibly physiological process of pH regulation in early life stage of marine medaka.

Marine debris occurrence and treatment: A review

Marine debris produces a wide variety of negative environmental, economic, safety, health and cultural impacts. Most marine litter has a very low decomposition rate (as plastics, which are the most abundant type of marine debris), leading to a gradual, but significant accumulation in the coastal and marine environment. Along that time, marine debris is a significant source of chemical contaminants to the marine environment. Once extracted from the water, incineration is the method most widely used to treat marine debris. Other treatment methods have been tested, but they still need some improvement and so far have only been used in some countries. Several extraction and collection programs have been carried out. However, as marine debris keep entering the sea, these programs result insufficient and the problem of marine debris will continue its increase. The present work addresses the environmental impact and social aspects of the marine debris, with a review of the state of the art in the treatments of this kind of waste, together with an estimation of the worldwide occurrence and characteristics.

Age determination of Antarctic marine sediments

We present the first circum-East Antarctic chronology for the Holocene, based on 17 radiocarbon dates generated by the accelerator method. Marine sediments from around East Antarctica contain a consistent, high-resolution record of terrigenous (ice-proximal) and biogenic (open-marine) sedimentation during Holocene time. This record demonstrates that biogenic sedimentation beneath the open-marine environment on the continental shelf has been restricted to approximately the past 4 ka, whereas a period of terrigenous sedimentation related to grounding line advance of ice tongues and ice shelves took place between 7 and 4 ka. An earlier period of open-marine (biogenic sedimentation) conditions following the late Pleistocene glacial maximum is recognized from the Prydz Bay (Ocean Drilling Program) record between 10.7 and 7.3 ka. Clearly, the response of outlet systems along the periphery of the East Antarctic ice sheet during the mid-Holocene was expansion. This may have been a direct consequence of climate warming during an Antarctic 'Hypsithermal'. Temperature-accumulation relations for the Antarctic indicate that warming will cause a significant increase in accumulation rather than in ablation. Models that predict a positive mass balance (growth) of the Antarctic ice sheet under global warming are supported by the mid-Holocene data presented herein.

Tetra- and tribromopbenoxyanisoles in Marine Samples from Oceania

Some methoxylated polybrominated diphenyl ethers (MeO-BDEs) are known halogenated natural products (HNPs) and are frequently detected in higher organisms of the marine environment. In this study we demonstrate that a prominent MeO-BDE, previously detected in marine mammals from Australia, is identical to 3,5-dibromo-2-(2',4'-dibromo)phenoxyanisole(BC-3,6-MeO-BDE47). Up to 1.9mg/ kg of 6-MeO-BDE 47 was present in cetaceans from Australia, 0.2-0.3 mg/kg in two crocodile eggs from Australia, but concentrations of 1 or 2 orders of magnitude lower were found in shark liver oil from New Zealand and in marine mammals from Africa and the Antarctic. Concentrations of 6-MeO-BDE47 in samples from Australia were in the same range as anthropogenic pollutants such as PCB 153 and p,p'-DDE. Along with 6-MeO-BDE 47 and the known HNP 4,6-dibromo-2-(2',4'-dibromo)phenoxyanisole (BC-2,2'-MeO-BDE 68), several tribromophenoxyanisoles (MeO-triBDE) were present in tissue of Australian cetaceans. To determine their structure, abiotic debromination experiments were performed using 6-MeO-BDE 47 and 2'-MeO-BDE 68 and superreduced di cyanocobalamine. These experiments resulted in formation of eight MeO-triBDEs, all of which were detected in the cetacean samples. Five of these eight MeO-triBDEs could be identified based on two standard compounds as well as gas chromatographic and mass spectrometric features. It was also shown that the first eluting isomer (compound 1), 6-MeO-BDE 17 (compound 2), and 2-MeO-BDE 39 (compound 5) were the most prominent MeO-triBDEs in the Australian cetacean samples. The concentrations of the MeO-triBDEs in two cetacean samples were 0.20 and 0.36 mg/kg, respectively. Although the reductive debromination with dicyanocobalamine resulted in a different congener pattern than was found in the marine mammals, it could not be excluded that the tribromo congeners of 6-MeO-BDE 47 and 2'-MeO-BDE 68 in the samples were metabolites of the latter.

Moreton Bay, Queensland, Australia: An example of the co-existence of significant marine mammal populations and large-scale coastal development

Recent analyses assert that large marine vertebrates such as marine mammals are now 'functionally or entirely extinct in most coastal ecosystems'. Moreton Bay is a large diverse marine ecosystem bordering the fastest growing area in Australia. The human population is over 1.6 million and increasing yearly by between 10% and 13% with resultant impacts upon the adjoining marine environment. Nonetheless, significant populations of three species of marine mammals are resident within Moreton Bay and a further 14 species are seasonal or occasional visitors. This paper reviews the current and historical distributions and abundance of these species in the context of the current management regime and suggests initiatives to increase the resilience of marine mammal populations to the changes wrought by the burgeoning human population in coastal environments. (C) 2004 Elsevier Ltd. All rights reserved.

A review of the potential role of tumour-promoting compounds produced by Lyngbya majuscula in marine turtle fibropapillomatosis

Harmful algal blooms (HABs) have increased in abundance and severity in recent decades. Whereas the implications for human impacts and intoxication resulting from blooms have been extensively studied, the ecological implications of these microalgae are less well understood. Many HAB species produce biologically active, secondary metabolites and the fate of these toxins through the foodweb is generally not well understood unless it culminates in extensive fish mortalities or human poisonings. This review focusses on one HAB species, the cyanobacterium Lyngbya majuscula, and presents a hypothetical role for its involvement in fibro-papillornatosis (FP), a neoplastic disease of marine turtles. FP is expressed as benign tumours that grow both internally and externally on marine turtles, preventing vision, movement and organ function. The aetiology of FP is currently not conclusively understood, but virus material has been associated with tumours and previous studies have suggested a role for naturally produced tumour promoters. In this review, we present a hypothesis regarding the involvement of L. majuscula in FP, either through direct intoxication and action of tumour-promoting compounds or indirectly by causing seagrass loss and compromised immune function, thus leaving the turtles more susceptible to disease.

Offspring size effects mediate competitive interactions in a colonial marine invertebrate

Over the past 30 years, numerous attempts to understand the relationship between offspring size and fitness have been made, and it has become clear that this critical relationship is strongly affected by environmental heterogeneity. For marine invertebrates, there has been a long-standing interest in the evolution of offspring size, but there have been very few empirical and theoretical examinations of post-metamorphic offspring size effects, and almost none have considered the effect of environmental heterogeneity on the offspring size/fitness relationship. We investigated the post-metamorphic effects of offspring size in the field for the colonial marine invertebrate Botrylloides violaceus. We also examined how the relationship between offspring size and performance was affected by three different types of intraspecific competition. We found strong and persistent effects of offspring size on survival and growth, but these effects depended on the level and type of intraspecific competition.. Generally, competition strengthened the advantages of increasing maternal investment. Interestingly, we found that offspring size determined the outcome of competitive interaction: juveniles that had more maternal investment were more likely to encroach on another juvenile's territory. This suggests that mothers have the previously unrecognized potential to influence the outcome of competitive interactions in benthic marine invertebrates. We created a simple optimality model, which utilized the data generated from our field experiments, and found that increasing intraspecific competition resulted in an increase,in predicted optimal size. Our results suggest that the relationship between offspring size and fitness is highly variable in the marine environment and strongly dependent on the density of conspecifics.