Benthic responses to organic matter variation in a subtropical coastal area off SE Brazil

Organic matter quality, expressed as the proportion of chlorophyll a (Chl a) to degraded organic material (i.e. phaeopigments), is known to influence the structure of benthic associations and plays an important role in the functioning of the ecosystem. This study investigates the vertical distribution of microbial biomass, meiofauna and macrofauna with respect to organic matter variation in Ubatuba, Brazil, a southeastern, subtropical coastal area. On three occasions, samples were collected in exposed and sheltered stations, at high and low hydrodynamic conditions. We hypothesize that benthic assemblages will have high meio- and macrofaunal densities and high microbial biomass at the sediment surface at the sheltered site, and lower and vertically homogeneous microbial biomass and densities of meio- and macrofauna are expected at the exposed site. The accumulation of fresh organic matter at the sediment surface was observed at both stations over the three sampling dates, which contributed to the higher densities of meiofauna in the first layers of the sediment column. Macrofauna followed the same trend only at the exposed station, but changes in the number of species, biodiversity and feeding groups were registered for both stations. Microbial biomass increased at the sheltered station over the three sampling dates, whereas at the exposed station, microbial biomass was nearly constant. Physical exposure did not influence organic matter loading at the sites and therefore did not affect overall structure of benthic assemblages, which negates our original hypothesis. Most of the benthic system components reacted to organic matter quality and quantity, but relationships between different-sized organisms (i.e. competition and/or predation) may explain the unchanged microbial profiles at the exposed site and homogeneous vertical distribution of macrofauna at the sheltered site. In conclusion, the high quality of organic matter was a crucial factor in sustaining and regulating the benthic system, but coupled results showed that interactions between micro-, meio- and macrofauna can be highly complex.

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.

Late Campanian and Maastrichtian benthic foraminifera of El Kef, Tunesia

Late Campanian through Maastrichtian sea-level changes are examined based on lithology, macrofossils and benthic foraminifera at the Elles and El Kef sections in Tunisia. Six major sea-level regressions are identified during the late Campanian (74.4-74.2 Ma, 74.0-72.5 Ma), the Campanian-Maastrichtian transition (72.2-70.3 Ma), early Maastrichtian (69.6-69.3 Ma, 68.9-68.3 Ma), and late Maastrichtian (~65.5 Ma). Correlation of the Maastrichtian sea-level regressions with the oxygen isotope record of DSDP Site 525 in the middle latitude South Atlantic reveals that they coincide with episodes of high latitude cooling and appear to be of eustatic origin.

Benthic respiration and physical oceanography on two contrasting continental margins in the western Indian Ocean: the Yemen-Somali upwelling region and the margin off Kenya

During the Netherlands Indian Ocean Project (NIOP, 1992-1993) sediment community oxygen consumption (SCOC) was measured on two continental margins in the Indian Ocean with different productivity: the productive upwelling region off Yemen-Somalia and the supposedly less productive Kenyan margin, which lacks upwelling. The two margins also differ in terms of river input (Kenya) and the more severe oxygen minimum in the Arabian Sea. Simultaneously with SCOC, distributions of benthic biomass and phytodetritus were studied. Our expectation was that benthic processes in the upwelling margin of the Arabian Sea would be relatively enhanced as a result of the higher productivity. On the Kenyan margin, SCOC (range 1-36 mmol/m**2/d) showed a clear decrease with increasing water depth, and little temporal variation was detected between June and December. Highest SCOC values of this study were recorded at 50 m depth off Kenya, with a maximum of 36 mmol/m**2/d in the northernmost part. On the margin off Yemen-Somalia, SCOC was on average lower and showed little downslope variation, 1.8-5.7 mmol/m**2/d, notably during upwelling, when the zone between 70 and 1700 m was covered with low O2 water (10-50 µM). After cessation of upwelling, SCOC at 60 m depth off Yemen increased from 5.7 to 17.6 mmol/m**2/d concurrently with an increase of the near-bottom O2 concentration (from 11 to 153 µM), suggesting a close coupling between SCOC and O2 concentration. This was demonstrated in shipboard cores in which the O2 concentration in the overlying water was raised after the cores were first incubated under in situ conditions (17 µM O2). This induced an immediate and pronounced increase of SCOC. Conversely, at deeper stations permanently within the oxygen minimum zone (OMZ), SCOC showed little variation between monsoon periods. Hence, organic carbon degradation in sediments on a large part of the Yemen slope appears hampered by the oxygen deficiency of the overlying water. Macrofauna biomass and the pooled biomass of smaller organisms, estimated by the nucleic acid content of the sediment, had comparable ranges in the two areas in spite of more severe suboxic conditions in the Arabian Sea. At the Kenyan shelf, benthic fauna (macro- and meiofauna) largely followed the spatial pattern of SCOC, i.e. high values on the northern shelf-upper slope and a downslope decrease. On the Yemen-Somali margin the macrofauna distribution was more erratic. Nucleic acids displayed no clear downslope trend on either margin owing to depressed values in the OMZ, perhaps because of adverse effects of low O2 on small organisms (meiofauna and microbes). Phytodetritus distributions were different on the two margins. Whereas pigment levels decreased downslope along the Kenya margin, the upper slope off Yemen (800 m) had a distinct accumulation of mainly refractory carotenoid pigments, suggesting preservation under low 02. Because the accumulations of Corg and pigments on the Yemen slope overlap only partly, we infer a selective deposition and preservation of labile particles on the upper slope, whereas refractory material undergoes further transport downslope.

Abundance and biomass of in- and epibenthic invertebrate macrofauna in the German Bight from 1969 to 2000

In order to examine the long-term development of offshore macrozoobenthic soft-bottom communities of the German Bight, four representative permanent stations (MZB-SSd, -FSd, -Slt, -WB) have been sampled continuously since 1969. Inter-annual variability and possible long-term trends were analysed based on spring-time samples from 1969 until 2000. This is part of the ecological long-term series of the AWI and is supplemented by periodic large-scale mapping of the benthos. The main factors influencing the development of the benthic communities are biological interactions, climate, food supply (eutrophication) and the disturbance regime. The most frequent disturbances are sediment relocations during strong storms or by bottom trawling, while occasional oxygen deficiencies and extremely cold winters are important disturbance events working on a much larger scale. Benthic communities at the sampling stations show a large inter-annual variability combined with a variation on a roughly decadal scale. In accordance with large-scale system shifts reported for the North Sea, benthic community transitions occurred between roughly the 1970ies, 80ies and 90ies. The transitions between periods are not distinctly marked by strong changes but rather reflected in gradual changes of the species composition and dominance structure.

Abiotic parameters and abundances, dominance and diversity of benthic macro- and meiofauna in Kongsfjord, Spitsbergen

The intertidal and subtidal soft bottom macro- and meiofauna of a glacier fjord on Spitsbergen was studied after complete ice melt in June 2003. The abundances of the benthic fauna were within the range reported from estuaries and similar intertidal areas of boreal regions. The high proportion of juveniles in the eulittoral zone indicated larval recruitment from subtidal areas. The macrobenthic fauna can be divided into an intertidal and a subtidal community, both being numerically dominated by annelids. Deposit feeders were numerically predominant in intertidal sites, whereas suspension feeders were most abundant in the subtidal area. Among the meiofauna, only the benthic copepods were identified to species, revealing ecological adaptations typical for intertidal species elsewhere.