Dauphin Island Sea Lab

Coastal ecosystems and the services they provide are adversely affected by a wide variety of human activities. In particular, seagrass meadows are negatively affected by impacts accruing from the billion or more people who live within 50 km of them. Seagrass meadows provide important ecosystem services, including an estimated $1.9 trillion per year in the form of nutrient cycling; an order of magnitude enhancement of coral reef fish productivity; a habitat for thousands of fish, bird, and invertebrate species; and a major food source for endangered dugong, manatee, and green turtle. Although individual impacts from coastal development, degraded water quality, and climate change have been documented, there has been no quantitative global assessment of seagrass loss until now. Our comprehensive global assessment of 215 studies found that seagrasses have been disappearing at a rate of 110 km(2) yr(-1) since 1980 and that 29% of the known areal extent has disappeared since seagrass areas were initially recorded in 1879. Furthermore, rates of decline have accelerated from a median of 0.9% yr(-1) before 1940 to 7% yr(-1) since 1990. Seagrass loss rates are comparable to those reported for mangroves, coral reefs, and tropical rainforests and place seagrass meadows among the most threatened ecosystems on earth.

ABSTRACT Seagrasses, marine flowering plants, have a long evolutionary history but are now challenged with rapid environmental changes as a result of coastal human population pressures. Seagrasses provide key ecological services, including organic carbon production and export, nutrient cycling, sediment stabilization, enhanced biodiversity, and trophic transfers to adjacent habitats in tropical and temperate regions. They also serve as “coastal canaries,” global biological sentinels of increasing anthropogenic influences in coastal ecosystems, with large-scale losses reported worldwide. Multiple stressors, including sediment and nutrient runoff, physical disturbance, invasive species, disease, commercial fishing practices, aquaculture, overgrazing, algal blooms, and global warming, cause seagrass declines at scales of square meters to hundreds of square kilometers. Reported seagrass losses have led to increased awareness of the need for seagrass protection, monitoring, management, and restoration. However, seagrass science, which has rapidly grown, is disconnected from public awareness of seagrasses, which has lagged behind awareness of other coastal ecosystems. There is a critical need for a targeted global conservation effort that includes a reduction of watershed nutrient and sediment inputs to seagrass habitats and a targeted educational program informing regulators and the public of the value of seagrass meadows.

The conservation status of 845 zooxanthellate reef-building coral species was assessed by using International Union for Conservation of Nature Red List Criteria. Of the 704 species that could be assigned conservation status, 32.8% are in categories with elevated risk of extinction. Declines in abundance are associated with bleaching and diseases driven by elevated sea surface temperatures, with extinction risk further exacerbated by local-scale anthropogenic disturbances. The proportion of corals threatened with extinction has increased dramatically in recent decades and exceeds that of most terrestrial groups. The Caribbean has the largest proportion of corals in high extinction risk categories, whereas the Coral Triangle (western Pacific) has the highest proportion of species in all categories of elevated extinction risk. Our results emphasize the widespread plight of coral reefs and the urgent need to enact conservation measures.

Impacts of chronic overfishing are evident in population depletions worldwide, yet indirect ecosystem effects induced by predator removal from oceanic food webs remain unpredictable. As abundances of all 11 great sharks that consume other elasmobranchs (rays, skates, and small sharks) fell over the past 35 years, 12 of 14 of these prey species increased in coastal northwest Atlantic ecosystems. Effects of this community restructuring have cascaded downward from the cownose ray, whose enhanced predation on its bay scallop prey was sufficient to terminate a century-long scallop fishery. Analogous top-down effects may be a predictable consequence of eliminating entire functional groups of predators.

Microbial ecology is plagued by problems of an abstract nature. Cell sizes are so small and population sizes so large that both are virtually incomprehensible. Niches are so far from our everyday experience as to make their very definition elusive. Organisms that may be abundant and critical to our survival are little understood, seldom described and/or cultured, and sometimes yet to be even seen. One way to confront these problems is to use data of an even more abstract nature: molecular sequence data. Massive environmental nucleic acid sequencing, such as metagenomics or metatranscriptomics, promises functional analysis of microbial communities as a whole, without prior knowledge of which organisms are in the environment or exactly how they are interacting. But sequence-based ecological studies nearly always use a comparative approach, and that requires relevant reference sequences, which are an extremely limited resource when it comes to microbial eukaryotes [1]. In practice, this means sequence databases need to be populated with enormous quantities of data for which we have some certainties about the source. Most important is the taxonomic identity of the organism from which a sequence is derived and as much functional identification of the encoded proteins as possible. In an ideal world, such information would be available as a large set of complete, well-curated, and annotated genomes for all the major organisms from the environment in question. Reality substantially diverges from this ideal, but at least for bacterial molecular ecology, there is a database consisting of thousands of complete genomes from a wide range of taxa, supplemented by a phylogeny-driven approach to diversifying genomics [2]. For eukaryotes, the number of available genomes is far, far fewer, and we have relied much more heavily on random growth of sequence databases [3],[4], raising the question as to whether this is fit for purpose.

Climate-driven changes in biotic interactions can profoundly alter ecological communities, particularly when they impact foundation species. In marine systems, changes in herbivory and the consequent loss of dominant habitat forming species can result in dramatic community phase shifts, such as from coral to macroalgal dominance when tropical fish herbivory decreases, and from algal forests to 'barrens' when temperate urchin grazing increases. Here, we propose a novel phase-shift away from macroalgal dominance caused by tropical herbivores extending their range into temperate regions. We argue that this phase shift is facilitated by poleward-flowing boundary currents that are creating ocean warming hotspots around the globe, enabling the range expansion of tropical species and increasing their grazing rates in temperate areas. Overgrazing of temperate macroalgae by tropical herbivorous fishes has already occurred in Japan and the Mediterranean. Emerging evidence suggests similar phenomena are occurring in other temperate regions, with increasing occurrence of tropical fishes on temperate reefs.

The importance of restoring filter-feeders, such as the Eastern oyster Crassostrea virginica, to mitigate the effects of eutrophication (e.g. in Chesapeake Bay) is currently under debate. The argument that bivalve molluscs alone cannot control phytoplankton blooms and reduce hypoxia oversimplifies a more complex issue, namely that ecosystem engineering species make manifold contributions to ecosystem services. Although further discussion and research leading to a more complete understanding is required, oysters and other molluscs (e.g. mussels) in estuarine ecosystems provide services far beyond the mere top-down control of phytoplankton blooms, such as (1) seston filtration, (2) benthic–pelagic coupling, (3) creation of refugia from predation, (4) creation of feeding habitat for juveniles and adults of mobile species, and for sessile stages of species that attach to molluscan shells, and (5) provision of nesting habitat.

Polychaetes are common in most marine habitats and dominate many infaunal communities. Functional guild classification based on taxonomic identity and morphology has linked community structure to ecological function. The functional guilds now include osmotrophic siboglinids as well as sipunculans, echiurans, and myzostomes, which molecular genetic analyses have placed within Annelida. Advances in understanding of encounter mechanisms explicitly relate motility to feeding mode. New analyses of burrowing mechanics explain the prevalence of bilateral symmetry and blur the boundary between surface and subsurface feeding. The dichotomy between microphagous deposit and suspension feeders and macrophagous carnivores, herbivores, and omnivores is further supported by divergent digestive strategies. Deposit feeding appears to be limited largely to worms longer than 1 cm, with juveniles and small worms in general restricted to ingesting highly digestible organic material and larger, rich food items, blurring the macrophage-microphage dichotomy that applies well to larger worms.

A perceived recent increase in global jellyfish abundance has been portrayed as a symptom of degraded oceans. This perception is based primarily on a few case studies and anecdotal evidence, but a formal analysis of global temporal trends in jellyfish populations has been missing. Here, we analyze all available long-term datasets on changes in jellyfish abundance across multiple coastal stations, using linear and logistic mixed models and effect-size analysis to show that there is no robust evidence for a global increase in jellyfish. Although there has been a small linear increase in jellyfish since the 1970s, this trend was unsubstantiated by effect-size analysis that showed no difference in the proportion of increasing vs. decreasing jellyfish populations over all time periods examined. Rather, the strongest nonrandom trend indicated jellyfish populations undergo larger, worldwide oscillations with an approximate 20-y periodicity, including a rising phase during the 1990s that contributed to the perception of a global increase in jellyfish abundance. Sustained monitoring is required over the next decade to elucidate with statistical confidence whether the weak increasing linear trend in jellyfish after 1970 is an actual shift in the baseline or part of an oscillation. Irrespective of the nature of increase, given the potential damage posed by jellyfish blooms to fisheries, tourism, and other human industries, our findings foretell recurrent phases of rise and fall in jellyfish populations that society should be prepared to face.

BACKGROUND: Vibrio parahaemolyticus, the leading cause of seafood-associated gastroenteritis in the United States, typically is associated with the consumption of raw oysters gathered from warm-water estuaries. We describe a recognized outbreak of V. parahaemolyticus infection associated with the consumption of seafood from Alaska. METHODS: After we received reports of the occurrence of gastroenteritis on a cruise ship, we conducted a retrospective cohort study among passengers, as well as active surveillance throughout Alaska to identify additional cases, and an environmental study to identify sources of V. parahaemolyticus and contributors to the outbreak. RESULTS: Of 189 passengers, 132 (70 percent) were interviewed; 22 of the interviewees (17 percent) met our case definition of gastroenteritis. In our multiple logistic-regression analysis, consumption of raw oysters was the only significant predictor of illness; the attack rate among people who consumed oysters was 29 percent. Active surveillance identified a total of 62 patients with gastroenteritis. V. parahaemolyticus serotype O6:K18 was isolated from the majority of patients tested and from environmental samples of oysters. Patterns on pulsed-field gel electrophoresis were highly related across clinical and oyster isolates. All oysters associated with the outbreak were harvested when mean daily water temperatures exceeded 15.0 degrees C (the theorized threshold for the risk of V. parahaemolyticus illness from the consumption of raw oysters). Since 1997, mean water temperatures in July and August at the implicated oyster farm increased 0.21 degrees C per year (P<0.001 by linear regression); 2004 was the only year during which mean daily temperatures in July and August at the shellfish farm did not drop below 15.0 degrees C. CONCLUSIONS: This investigation extends by 1000 km the northernmost documented source of oysters that caused illness due to V. parahaemolyticus. Rising temperatures of ocean water seem to have contributed to one of the largest known outbreaks of V. parahaemolyticus in the United States.

We reviewed studies providing quantitative measurements of abundance of fishes and large mobile crustaceans on oyster reefs and on nearby sedimentary habitat in the southeast United States. For each species, we compared density by size (age) class on oyster reefs and sedimentary bottom as a means of estimating the degree to which restoration of oyster reef on sedimentary bottom could augment abundances. By applying published information on growth rates of each species and a combination of empirical data and published information on age-specific survivorship, we calculated the per-unit-area enhancement of production of fishes and large mobile crustaceans expected from the addition of oyster reef habitat. For this calculation, we gave the reef habitat full credit for the expected lifetime production of species whose recruitment was judged to be limited by the area of oyster reefs based on nearly exclusive association of recruits to reefs. For species that were only modestly enhanced in abundance by oyster reefs, we gave the reef credit for the fraction of production that is derived from consumption of reef-associated prey, using a combination of gut content data and natural history information. This combination of analyses and calculations revealed that 10 m 2 of restored oyster reef in the southeast United States is expected to yield an additional 2.6 kg yr -1 of production of fish and large mobile crustaceans for the functional lifetime of the reef. Because the reef is biogenic and self-sustaining, the lifetime of a reef protected from bottom-disturbing fishing gear is limited by intense storms or sedimentation. A reef lasting 20 to 30 yr would be expected to augment fish and large mobile crustacean production by a cumulative amount of 38 to 50 kg 10 m -2 , discounted to present-day value. This set of calculations assumes that oyster reef habitat now limits production of reef-associated fish and crustaceans in the southeast United States. This assumption seems reasonable based on the tight associations of so many fishes with reef-dependent prey, and the depletion of reef habitat over the past century.

Since the recognition of prokaryotes as essential components of the oceanic food web, bacterioplankton have been acknowledged as catalysts of most major biogeochemical processes in the sea. Studying heterotrophic bacterioplankton has been challenging, however, as most major clades have never been cultured or have only been grown to low densities in sea water. Here we describe the genome sequence of Silicibacter pomeroyi, a member of the marine Roseobacter clade (Fig. 1), the relatives of which comprise approximately 10-20% of coastal and oceanic mixed-layer bacterioplankton. This first genome sequence from any major heterotrophic clade consists of a chromosome (4,109,442 base pairs) and megaplasmid (491,611 base pairs). Genome analysis indicates that this organism relies upon a lithoheterotrophic strategy that uses inorganic compounds (carbon monoxide and sulphide) to supplement heterotrophy. Silicibacter pomeroyi also has genes advantageous for associations with plankton and suspended particles, including genes for uptake of algal-derived compounds, use of metabolites from reducing microzones, rapid growth and cell-density-dependent regulation. This bacterium has a physiology distinct from that of marine oligotrophs, adding a new strategy to the recognized repertoire for coping with a nutrient-poor ocean.

&amp;lt;p&amp;gt;Biomass is defined as organic matter from living organisms represented in all kingdoms. It is recognized to be an excellent source of proteins, polysaccharides and lipids and, as such, embodies a tailored feedstock for new products and processes to apply in green industries. The industrial processes focused on the valorization of terrestrial biomass are well established, but marine sources still represent an untapped resource. Oceans and seas occupy over 70% of the Earth&amp;rsquo;s surface and are used intensively in worldwide economies through the fishery industry, as logistical routes, for mining ores and exploitation of fossil fuels, among others. All these activities produce waste. The other source of unused biomass derives from the beach wrack or washed-ashore organic material, especially in highly eutrophicated marine ecosystems. The development of high-added-value products from these side streams has been given priority in recent years due to the detection of a broad range of biopolymers, multiple nutrients and functional compounds that could find applications for human consumption or use in livestock/pet food, pharmaceutical and other industries. This review comprises a broad thematic approach in marine waste valorization, addressing the main achievements in marine biotechnology for advancing the circular economy, ranging from bioremediation applications for pollution treatment to energy and valorization for biomedical applications. It also includes a broad overview of the valorization of side streams in three selected case study areas: Norway, Scotland, and the Baltic Sea.&amp;lt;/p&amp;gt;

Shorelines at the interface of marine, estuarine and terrestrial biomes are among the most degraded and threatened habitats in the coastal zone because of their sensitivity to sea level rise, storms and increased human utilization. Previous efforts to protect shorelines have largely involved constructing bulkheads and seawalls which can detrimentally affect nearshore habitats. Recently, efforts have shifted towards "living shoreline" approaches that include biogenic breakwater reefs. Our study experimentally tested the efficacy of breakwater reefs constructed of oyster shell for protecting eroding coastal shorelines and their effect on nearshore fish and shellfish communities. Along two different stretches of eroding shoreline, we created replicated pairs of subtidal breakwater reefs and established unaltered reference areas as controls. At both sites we measured shoreline and bathymetric change and quantified oyster recruitment, fish and mobile macro-invertebrate abundances. Breakwater reef treatments mitigated shoreline retreat by more than 40% at one site, but overall vegetation retreat and erosion rates were high across all treatments and at both sites. Oyster settlement and subsequent survival were observed at both sites, with mean adult densities reaching more than eighty oysters m(-2) at one site. We found the corridor between intertidal marsh and oyster reef breakwaters supported higher abundances and different communities of fishes than control plots without oyster reef habitat. Among the fishes and mobile invertebrates that appeared to be strongly enhanced were several economically-important species. Blue crabs (Callinectes sapidus) were the most clearly enhanced (+297%) by the presence of breakwater reefs, while red drum (Sciaenops ocellatus) (+108%), spotted seatrout (Cynoscion nebulosus) (+88%) and flounder (Paralichthys sp.) (+79%) also benefited. Although the vertical relief of the breakwater reefs was reduced over the course of our study and this compromised the shoreline protection capacity, the observed habitat value demonstrates ecological justification for future, more robust shoreline protection projects.

Sterilized seawater was used to assess the effects of temperature and salinity on the survival of Vibrio vulnificus. In the temperature range of 13 to 22 degrees C, numbers of V. vulnificus increased during the 6-day incubation. Temperatures outside this range reduced the time of V. vulnificus survival in sterile 10-ppt seawater. At these restrictive temperatures, V. vulnificus numbers were reduced by 90% after 6 days of incubation. Incubation between 0.5 and 10.5 degrees C demonstrated that V. vulnificus survives poorly below 8.5 degrees C. At salinities between 5 and 25 ppt and at 14 degrees C, V. vulnificus numbers actually increased or remained unchanged after 6 days of incubation. At salinities of 30, 35, and 38 ppt, numbers of V. vulnificus decreased 58, 88, and 83%, respectively. V. vulnificus could not be recovered from deionized water, indicating lysis. When a rifampin-resistant strain of V. vulnificus was used to inoculate sterilized and unsterilized seawater (20 ppt, 20 degrees C), numbers increased in sterile seawater but decreased to undetectable levels in 14 days in the unsterilized seawater, indicating that biological factors may play a role in the survival of V. vulnificus in the environment. Since our studies demonstrated sensitivity to low temperatures, the survival of V. vulnificus in naturally contaminated oysters at temperatures of 0, 2, and 4 degrees C was also determined. Numbers of endogenous V. vulnificus in oyster shellstock increased by more than 100-fold in shellstock stored at 30 degrees C but were reduced approximately 10- and 100-fold after 14 days at 2 to 4 degrees C and 0 degrees C, respectively. We conclude that both biological and physicochemical factors are important to the survival of V. vulnificus in the environment and that temperature is critical to controlling its growth in oyster shellstock.

Historic baselines are important in developing our understanding of ecosystems in the face of rapid global change. While a number of studies have sought to determine changes in extent of exploited habitats over historic timescales, few have quantified such changes prior to late twentieth century baselines. Here, we present, to our knowledge, the first ever large-scale quantitative assessment of the extent and biomass of marine habitat-forming species over a 100-year time frame. We examined records of wild native oyster abundance in the United States from a historic, yet already exploited, baseline between 1878 and 1935 (predominantly 1885-1915), and a current baseline between 1968 and 2010 (predominantly 2000-2010). We quantified the extent of oyster grounds in 39 estuaries historically and 51 estuaries from recent times. Data from 24 estuaries allowed comparison of historic to present extent and biomass. We found evidence for a 64 per cent decline in the spatial extent of oyster habitat and an 88 per cent decline in oyster biomass over time. The difference between these two numbers illustrates that current areal extent measures may be masking significant loss of habitat through degradation.

Jellyfish (Cnidaria, Scyphozoa) blooms appear to be increasing in both intensity and frequency in many coastal areas worldwide, due to multiple hypothesized anthropogenic stressors. Here, we propose that the proliferation of artificial structures – associated with (1) the exponential growth in shipping, aquaculture, and other coastal industries, and (2) coastal protection (collectively, “ocean sprawl”) – provides habitat for jellyfish polyps and may be an important driver of the global increase in jellyfish blooms. However, the habitat of the benthic polyps that commonly result in coastal jellyfish blooms has remained elusive, limiting our understanding of the drivers of these blooms. Support for the hypothesized role of ocean sprawl in promoting jellyfish blooms is provided by observations and experimental evidence demonstrating that jellyfish larvae settle in large numbers on artificial structures in coastal waters and develop into dense concentrations of jellyfish‐producing polyps.

Seagrasses are important marine foundation species that are reported to be declining worldwide, with almost 15% of species considered threatened. Seagrasses are highly productive plants that reconfigure water flow and influence nutrient cycling, as well as provide critical habitat for a wide array of fish and invertebrate species. Yet, many of these seagrass‐dependent species, including economically important fishes and invertebrates, are themselves in danger of overexploitation or extinction. In fact, there is on average more than one threatened associated species for every seagrass species across the globe. Links between threatened seagrasses and their dependent communities illustrate the importance of an ecosystem‐based management approach that incorporates interdependencies and facilitation among species.

Describing the relative magnitude and controls of herbivory and decomposition is important in understanding the trophic transference, recycling, and storage of carbon and nutrients in diverse ecosystems. We examine the variability in herbivory and decomposition between and within a wide range of aquatic and terrestrial ecosystems. We also analyze how that variability is associated with differences in net primary production and producer nutritional quality. Net primary production and producer nutritional quality are uncorrelated between the two types of system or within either type. Producer nutritional quality is correlated to the percentage of primary production consumed by herbivores or percentage of detrital production decomposed annually, regardless of whether the comparison is made between the two types of systems or within either type of system. Thus, producer nutritional quality stands out as a consistent indicator of the importance of consumers as top‐down controls of producer biomass and detritus accumulation and nutrient recycling. However, absolute consumption by herbivores and absolute decomposition (both in g C·m −2 ·yr −1 ) are often associated with absolute primary production and independent of producer nutritional quality, because the variability in net primary production across systems largely exceeds that in the percentage consumed or decomposed. Thus, primary production often stands out as an indicator of the absolute flux of producer carbon transferred to consumers and of the potential levels of secondary production maintained in the system. These patterns contribute to our understanding of the variability and control of herbivory and decomposition, and implications on carbon and nutrient cycling, in aquatic and terrestrial ecosystems. Furthermore, in view of their robustness, they may offer a template for global change models seeking to predict anthropogenic effects on carbon and nutrient fluxes.

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 247:59-73 (2003) - doi:10.3354/meps247059 Eelgrass Zostera marina loss in temperate estuaries: relationship to land-derived nitrogen loads and effect of light limitation imposed by algae Jennifer Hauxwell1,2,4,*, Just Cebrián1,3,5, Ivan Valiela1 1Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts 02543, USA 2Wisconsin Department of Natural Resources, DNR Research Center, 1350 Femrite Drive, Monona, Wisconsin 53716, USA 3Dauphin Island Sea Lab, 101 Bienville Boulevard, PO Box 369-370, Dauphin Island, Alabama 36528, USA 4Present address: Wisconsin Department of Natural Resources, DNR Research Center, 1350 Femrite Drive, Monona, Wisconsin 53716, USA 5Present address: Dauphin Island Sea Lab, 101 Bienville Boulevard, PO Box 369-370, Dauphin Island, Alabama 36528, USA *Email: jennifer.hauxwell@dnr.state.wi.us ABSTRACT: In this paper, we explicitly link changes in community structure of estuarine primary producers to measured nitrogen loading rates from watersheds to estuaries, and quantify the relationship between nitrogen load, annual dynamics of algal growth and Zostera marina L. productivity, and overall eelgrass decline at the watershed-estuarine scale in estuaries of Waquoit Bay, Massachusetts, USA. Substantial eelgrass loss (80 to 96% of bed area lost in the last decade) was found at loads of ~30 kgN ha-1 yr-1, and total disappearance at loads ≥60 kg N ha-1 yr-1. Rather than decreased eelgrass growth rates, we observed an exponential decrease in shoot densities and bed area (and subsequently areal production) as nitrogen loads increased, suggesting that eelgrass decline in higher-nitrogen estuaries of the Waquoit system occurred largely via lack of recruitment or enhanced mortality of established shoots. Similar to the patterns observed in many other systems and the experimental results obtained in laboratories or mesocosms, the relationship we observed between nitrogen loads and eelgrass health within the Waquoit system was indirect: increased nitrogen stimulated growth and standing stocks of algal producers, that may have caused severe light limitation of eelgrass. From light budgets that considered water column, epiphyte, and macroalgal shading, we estimated chronic, severe light limitation to newly recruiting shoots in higher-nitrogen estuaries, due mainly to shading by a coexisting ≤15 cm macroalgal canopy. Two management recommendations aimed at eelgrass preservation emerge from this work. First, development and management of watersheds must be conducted such that land-derived nitrogen loading to estuaries is restricted. In the Waquoit Bay estuaries, for example, eelgrass is absent or rapidly disappearing from all but those receiving the lowest (≤15th percentile) loads. Second, shoot density and meadow area, rather than g rates per shoot, seem to be adequate variables for routine monitoring of eelgrass health. We also show that the shift from eelgrass- to algae-dominated communities has important consequences for total system primary production and carbon and nitrogen cycling. Estimated total primary production by coastal assemblages in the Waquoit Bay system was 135% higher in estuaries receiving relatively high versus low loads of land-derived nitrogen, suggesting important trophic and biogeochemical alterations to temperate estuarine ecosystems as a result of eutrophication. KEY WORDS: Seagrass · Macroalgae · Epiphytes · Phytoplankton · Irradiance · Waquoit Bay · Eutrophication · Estuary Full text in pdf format PreviousNextExport citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 247. Online publication date: February 04, 2003 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2003 Inter-Research.