Open Marine Systems Research Articles

Origin and role of non-skeletal carbonate in coralligenous build-ups: new geobiological perspectives in biomineralization processes

Abstract. The coralligenous build-ups located on the Mediterranean shelf in front of Marzamemi (SE Sicily, Italy) represent useful natural examples to use in studying the relationship between skeletal organisms and non-skeletal components in marine bioconstructions. Coralligenous build-ups are formed in open marine systems, and their comparison with coeval bioconstructions (biostalactites) of confined environments, like submarine caves, allows depicting the complex interactions between metazoans and microbial communities in the formations of recent bioconstructions in different Mediterranean settings. In this study, two coralligenous build-ups were characterized in terms of organisms and sediments involved in their formation. The framework mainly consists of coralline algae and subordinate bryozoans and serpulids. Sponges affect the general morphology of the bioconstructions both interacting with skeletonized organisms and through bioerosion activity. The micrite or microcrystalline calcite is present in minor amounts compared to other components that form the build-ups and consists of two types: autochthonous (in situ) and allochthonous (detrital). Fine autochthonous micrite mineralized directly inside the framework cavities and shows aphanitic or peloidal fabric, produced by organomineralization processes of soft sponge tissues and microbial metabolic activity, respectively. The detrital micrite occurring inside cavities derives from external sources or erosion processes of the bioconstructions themselves. This component has been classified as organic or inorganic based on the organic matter contents deduced by UV epifluorescence. A great quantity of sponges live in cavities of the coralligenous build-ups and compete with carbonatogenic bacteria for the same cryptic spaces, limiting the production of microbialites. The sharing of a similar relationship between sponges and microbial communities by coralligenous concretion and biotic crusts of particular submarine caves suggests that this competition is not habitat-specific. On the contrary, it may develop in a range of environmental settings, from open to cryptic systems, and could be used to clarify the role of metazoans vs. microbialites in palaeoecological reconstructions.

Benthic-pelagic coupling and isotopic fractionation of barium in Kiel Bight, SW Baltic Sea

Barium (Ba) isotopes are a promising new tracer for riverine freshwater input to the ocean and marine biogeochemical cycling. However, many processes that affect Ba cycling at continental margins have not yet been investigated with respect to Ba isotope fractionation. Here, we present a comprehensive data set of Ba concentration and isotope data for water column, pore water and sediment samples from Kiel Bight, a seasonally stratified and hypoxic fjord in the southwestern Baltic Sea. The surface water Ba concentration and Ba isotope inventory of the water column can generally be explained by mixing of riverine freshwater and Atlantic seawater. However, the deep-water below the seasonal pycnocline (10 - 15 m water depth) is characterized by a pronounced positive Ba concentration anomaly (up to 915 nM) that is accompanied by a δ138Ba of ~+0.25 ‰, which is lighter than expected from the seawater-freshwater mixing line (Ba: 77 nM, δ138Ba: +0.32 ‰ at a salinity of 18). Pore water profiles indicate a Ba flux across the sediment-water interface, which contributes to the enrichment in isotopically light Ba in the deep-water. Pore waters of surface sediments and deep-waters are oversaturated with respect to barite. Therefore, barite dissolution is unlikely to account for the benthic Ba flux. Water column Ba concentrations closely correlate with those of the nutrients phosphate and silica, which are removed from surface waters by biological processes and recycled from the sediment by diffusion across the sediment-water interface. As nutrient-to-Ba ratios differ among sites and from those observed in open-marine systems, we propose that Ba is removed from surface waters by adsorption onto biogenic particles (rather than assimilation) and regenerated within surface sediments upon organic matter degradation. Pore water data for subsurface sediments in Kiel Bight indicate preferential transfer of isotopically heavy Ba into an authigenic phase during early diagenesis. Quantifying the burial flux associated with this authigenic Ba phase along continental margins could potentially help to settle the isotopic imbalance between known Ba source and sink fluxes in the ocean.

Long-chain alkenones in the lake sediments of North-Minusinsk Valley (southern Siberia): implications for paleoclimate reconstructions

Long-chain alkenones (LCAs) are polyunsaturated methyl or ethyl ketones exclusively produced by Groups 1–3 Isochrysidales, an order of haptophyte algae. LCAs have great potential as temperature and salinity proxies for paleo-reconstructions. Alkenone producers occur extensively in lacustrine and in open marine systems. However, factors influencing LCAs occurrence, abundance, compositional pattern, and unsaturation indices are insufficiently understood in lakes of southern Siberia. Here we investigated alkenone distributions in freshwater to saline lakes of a semi-arid region of southern Siberia (n = 22). We find that total alkenone concentrations increase dramatically above a salinity of about 20 g L−1. Analysis of 18S rRNA and the alkenone composition suggest that all alkenone-containing lakes are inhabited by Group 2 haptophytes except one low-saline lake, where Group 1 representatives prevail. The C37/C38 ratio is negatively correlated with salinity, and, hence, this parameter can be used as a salinity indicator. At the same time, %C37:4 is not correlated with salinity, whereas the average chain length of alkenones increases with salinity. The taxonomic composition of haptophytes and the alkenone composition were comparable to those in the lakes in the Canadian Prairies, potentially because of the similarities in climatic factors and ionic composition of lake water in the two regions. This study shows for the first time that unsaturation indexes U40K and U40K′ are positively correlated with salinity and, thus, can be used as a salinity proxy for regional climate reconstruction.

Rapid flocculation and settling of positively buoyant microplastic and fine-grained sediment in natural seawater

Interactions between microplastic (MP) and fine-grained suspended sediment in natural waters are important for the environmental fate of plastic particles. Estuaries are transitional areas between freshwater and open marine systems and are recognized as important accumulation zones for MPs. However, there is a knowledge gap on the processes driving the sedimentation of MPs in estuaries, especially with regard to positively buoyant MPs. Here we show from settling tube experiments that positively buoyant and non-spherical MP HDPE particles in different size-fractions (63–500 μm) and concentrations (1 and 5 mg l−1) rapidly flocculate and settle with natural fine-grained sediment in natural seawater. Our results demonstrate that flocculation is a key process for the vertical transport of MP in estuaries. The implication is that land-based sources of positively buoyant HDPE MP transported by rivers will likely settle and accumulate in estuarine environments and thereby increase the concentration of MP in the benthic zone.

Understanding seasonal migration of Shishamo smelt in coastal regions using environmental DNA.

Migratory organisms have their own life histories that efficiently link multiple ecosystems. Therefore, comprehensive understanding of migration ecologies of these organisms is essential for both species conservation and ecosystem management. However, monitoring migration at fine spatiotemporal scales, especially in open marine systems, often requires huge costs and effort. Recently, environmental DNA (eDNA) techniques that utilize DNA released from living organisms into their environment became available for monitoring wild animals without direct handling. In this study, we conducted an eDNA survey for understanding marine migration of an endemic fish species, Shishamo smelt (Spirinchus lanceolatus). We examined 1) seasonal habitat changes in coastal regions and 2) environmental factors potentially driving the migration of this species. The eDNA concentrations along a 100 km-long coastline exhibited spatiotemporal variation, suggesting that this species shifts their habitat away from nearshore areas between spring and summer. We also found a significantly negative association between the eDNA concentration and sea surface temperature. That finding suggests that the offshore migration of this species is associated with increased sea surface temperature. This study reveals new aspects of S. lanceolatus life history in coastal regions. Together with our previous eDNA study on the freshwater migration of S. lanceolatus, this study illustrates the potential of eDNA techniques for understanding the whole life history of this migratory species.

The fate of a Neoproterozoic intracratonic marine basin: Trace elements, TOC and IRON speciation geochemistry of the Bambuí Basin, Brazil

Neoproterozoic marine systems are associated with major paleoecological changes that took place in the Ediacaran and during the Ediacaran-Cambrian transition. During this timespan, the Bambuí basin located on east Brazil held a peculiar paleoenvironmental scenario. Due to its intracratonic evolution, the basin was partially disconnected from neighboring open marine systems. This setting raises a very interesting opportunity to understand how an isolated Neoproterozoic marine system evolved in contrast with typical (globally connected) open marine systems. To understand the paleoenvironmental changes that took place in the Bambuí basin, we investigate the pre-glaciogenic deposits of the Carrancas Fm and the post-glaciogenic mixed (shale-carbonate) successions of the Bambuí Group. Through the analysis of iron speciation, TOC, trace element and C-O isotope systematics, our study suggests a very complex environmental evolution. Firstly, our samples are marked by strong contamination of detrital continental material that can be related to an increased bioproductivity on both the Carrancas Fm. and lower Bambuí group stratigraphic units, and provenance data show that all studied sediments probably shared common source areas. Iron speciation data, Ce anomalies and RSE enrichments shows that lower Bambuí Group stratigraphic units were likely deposited in an open marine scenario featuring high bioproductivity in shallow waters and euxinic incursions in predominant anoxic/ferruginous bottom waters. On the other hand, upper Bambuí stratigraphic units register a marine evolution in a restricted scenario, where anoxic ferruginous conditions probably reached surface waters. Finally, our data show that the lack of oceanic connection prevented the re-supply of marine sulfate, RSE, bionutrients and ultimately of dissolved oxygen which may have decreased biological activity and probably hindered biological evolution, preventing the rise of a typical modern-like Cambrian ecosystem. In this sense, our data suggest that oceanic connectivity and proper re-supply of inorganic marine input were important features in the development of complex life in the Ediacaran-Cambrian environment.

Quantifying the effect of diagenetic recrystallization on the Mg isotopic composition of marine carbonates

The Mg and Sr isotopic compositions (δ26Mg and 87Sr/86Sr) of pore fluids and bulk carbonates from Ocean Drilling Project Site 1171 (South Tasman Rise; 2148.2m water depth) are reported, in order to evaluate the potential of diagenesis to alter carbonate-based geochemical proxies in an open marine system. Given the trace amounts of Mg in marine carbonates relative to coexisting pore fluids, diagenesis can alter carbonate δ26Mg, a promising proxy for seawater δ26Mg that may help elucidate long-term changes in the global Mg cycle. Constraints on the effect of diagenetic recrystallization on carbonate δ26Mg are therefore critical for accurate proxy interpretations. This study provides context for assessing the fidelity of geochemical proxy-reconstructions using the primary components (i.e., foraminiferal tests and nannofossils) of bulk carbonate sediments.We find that pore fluid δ26Mg values (on the DSM3 scale) at Site 1171 increase systematically with depth (from −0.72‰ to −0.39‰ in the upper ∼260m), while the δ26Mg of bulk carbonates decrease systematically with depth (from −2.23‰ to −5.00‰ in the upper ∼260m). This variability is ascribed primarily to carbonate recrystallization, with a small proportion of the variability due to down-hole changes in nannofossil and foraminiferal species composition. The inferred effect of diagenesis on bulk carbonate δ26Mg correlates with down-core changes in Mg/Ca, Sr/Ca, Na/Ca, and 87Sr/86Sr. A depositional reactive-transport model is employed to validate the hypothesis that calcite recrystallization in this system can generate sizeable shifts in carbonate δ26Mg. Model fits to the data suggest a fractionation factor and a partition coefficient that are consistent with previous work, assuming calcite recrystallization rates of ⩽7%/Ma constrained by Sr geochemistry. In addition, either partial dissolution or a distinctly different previous diagenetic regime must be invoked in order to explain aspects of the elemental chemistry and 87Sr/86Sr of relatively deep sediments from Holes A and C. This study indicates that the dynamics of a given sedimentary system can significantly alter bulk carbonate geochemistry, and presents a framework for considering the potential impact of such alteration on picked archives such as foraminiferal tests and nannofossils. Ultimately, this study contributes to the development of δ26Mg as a proxy for seawater δ26Mg by quantifying the susceptibility of carbonate δ26Mg to diagenetic alteration, particularly in sediments in open marine systems. This study suggests that because of the sensitivity of carbonate δ26Mg to diagenetic recrystallization, it can, in certain systems, be used to quantify the impact of diagenesis on carbonate-based geochemical proxies.

Three routes to crypsis: Stasis, convergence, and parallelism in the Mastigias species complex (Scyphozoa, Rhizostomeae)

Evolutionary inference can be complicated by morphological crypsis, particularly in open marine systems that may rapidly dissipate signals of evolutionary processes. These complications may be alleviated by studying systems with simpler histories and clearer boundaries, such as marine lakes—small bodies of seawater entirely surrounded by land. As an example, we consider the jellyfish Mastigias spp. which occurs in two ecotypes, one in marine lakes and one in coastal oceanic habitats, throughout the Indo-West Pacific (IWP). We tested three evolutionary hypotheses to explain the current distribution of the ecotypes: (H1) the ecotypes originated from an ancient divergence; (H2) the lake ecotype was derived recently from the ocean ecotype during a single divergence event; and (H3) the lake ecotype was derived from multiple, recent, independent, divergences. We collected specimens from 21 locations throughout the IWP, reconstructed multilocus phylogenetic and intraspecific relationships, and measured variation in up to 40 morphological characters. The species tree reveals three reciprocally monophyletic regional clades, two of which contain ocean and lake ecotypes, suggesting repeated, independent evolution of coastal ancestors into marine lake ecotypes, consistent with H3; hypothesis testing and an intraspecific haplotype network analysis of samples from Palau reaffirms this result. Phylogenetic character mapping strongly correlates morphology to environment rather than lineage (r=0.7512, p<0.00001). Considering also the deeper relationships among regional clades, morphological similarity in Mastigias spp. clearly results from three separate patterns of evolution: morphological stasis in ocean medusae, convergence of lake morphology across distinct species and parallelism between lake morphologies within species. That three evolutionary routes each result in crypsis illustrates the challenges of interpreting evolutionary processes from patterns of biogeography and diversity in the seas. Identifying cryptic species is only the first step in understanding these processes; an equally important second step is exploring and understanding the processes and patterns that create crypsis.

Textures and In Situ Chemical and Isotopic Analyses of Pyrite, Huijiabao Trend, Youjiang Basin, China: Implications for Paragenesis and Source of Sulfur

Many Carlin-like Au deposits occur within the late Paleozoic and Triassic Youjiang basin of southwest China. The Huijiabao trend in Guizhou Province contains over 300 metric tons (t; 10.6 Moz) of Au at an average grade of 7 to 18 g/t in a narrow corridor that is about 20 km long and 5 km wide. Petrographic and SEM studies of pyrite in barren host rocks and high-grade orebodies led to the recognition of four stages of pyrite. Py1 consists of fine-grained framboidal crystals in black mudstone. Py2 is comprised of coarser grained euhedralsubhedral clusters that are spatially related to organic matter. Py3 is coarse grained, euhedral, and occurs as overgrowths on Py1 and Py2. Py3’s porous texture, inclusion of randomly oriented detrital minerals, and association with quartz recrystallization suggest it was deformed during Late Triassic orogenesis with Py1 and Py2. Py4 generally occurs as rims on Py1 to Py3 and is intergrown with arsenopyrite. Sensitive high-resolution ion microprobe (SHRIMP) δ 34S analyses of each pyrite type and arsenopyrite show that Py1 is related to Py2 and that Py3 is related to Py4 and arsenopyrite. The S isotope compositions of Py1 (−7.5 to +5.9‰) and Py2 (−5.3 to +7.9‰) are bimodal, which suggests that H2S was generated by biogenic sulfate reduction in open marine and sulfate limited systems during sedimentation and/or diagenesis. The compositions of Py3 (−2.6 to +1.5‰), Py4 (−1.2 to +1.5‰), and arsenopyrite (−0.8 to +0.9‰) are homogeneous and have an intermediate range of values near 0‰ that suggest that H2S was derived either from average pyrite (0.2‰) in sedimentary rocks or from a concealed magmatic source. Laser ablation-inductively coupled plasma-mass spectrometer (LA–ICP–MS) trace element analyses (As, Ni, Co, Cu, Ag, Se, V) support different origins and show that Py3 and Py4 are ore related. The lower w (Co)/ w (Ni) and w (S)/ w (Se) ratios of Py1 and Py2 are consistent with formation during sedimentation or diagenesis, whereas the higher ratios of Py3, Py4, and arsenopyrite are consistent with a hydrothermal origin. The lower concentrations of Au in Py1 (0.23–2.5 ppm) and Py2 (0.06–12 ppm) show that little Au was added during sedimentation or diagenesis. The higher concentrations of Au in hydrothermal Py3 (1.1–110 ppm) and Py4 (0.34–810 ppm) indicate that most of the Au was introduced during subsequent hydrothermal fluid flow. The low Au contents of arsenopyrite (0.09–0.52 ppm) suggests they formed from Au-depleted fluids. The Au/As ratios of Py1 and Py2 are typical of diagenetic pyrite whereas Py3 and Py4 have ratios that approach those of ore-stage pyrite in Nevada Carlin-type deposits. The fracturing of Py3 and its cementation by Py4 suggests that ore fluid movement was associated with deformation. Published isochron ages on arsenopyrite (Re-Os ~200 Ma) and late calcite-realgar veinlets (Sm-Nd ~135 Ma) in the Huijiabao trend are older than mafic dikes (84 Ma) exposed ~20 km to the east. If the 200 and 135 Ma ages are valid, H2S and Au may be derived from a sedimentary source because igneous intrusions of this age have not been found. If these ages are not valid and the gold deposits are actually Late Cretaceous in age, then H2S and Au may be derived from a magmatic source. Additional geochronology and isotopic tracer studies are needed to resolve this uncertainty.

From regionally predictable to locally complex population structure in a freshwater top predator: river systems are not always the unit of connectivity in Northern Pike Esox lucius

AbstractContemporary genetic diversity is the product of both historical and contemporary forces, such as climatic and geological processes affecting range distribution and continuously moulded by evolutionary forces selection, gene flow and genetic drift. Predatory freshwater fishes, such as Northern Pike Esox lucius, commonly exhibit small population sizes, and several local populations are considered endangered. Pike inhabit diverse habitat types, including lakes, rivers and brackish marine waters, thus spanning from small isolated patches to large open marine systems. However, pike population structure from local to regional scales is relatively poorly described, in spite of its significance to developing conservation measures. We analysed microsatellite variation in a total of 1185 North European pike from 46 samples collected across both local and regional scales, as well as over time, to address two overarching questions: Is pike population structure associated with local and/or regional connectivity patterns, and which factors likely have the main influence on the contemporary distribution of genetic diversity? To answer this, we combined estimators of population diversity and structure to assess evidence of whether populations within (i) habitats, (ii) drainage systems and (iii) geographical regions are closer related than among these ranges, and whether patterns are temporally stable. Contrasting previous predictions that genetic drift obscures signals of postglacial colonisation history, we identified clear regional differences in population genetic signatures, suggesting a major effect of drainage divides on colonisation history and connectivity. However, several populations deviated from the general pattern, showing that local processes may be complex and need to be examined case‐by‐case.

Plastic as a Carrier of POPs to Aquatic Organisms: A Model Analysis

It has been hypothesized that persistent organic pollutants (POPs) in microplastic may pose a risk to aquatic organisms. Here we develop and analyze a conceptual model that simulates the effects of plastic on bioaccumulation of POPs. The model accounts for dilution of exposure concentration by sorption of POPs to plastic (POP "dilution"), increased bioaccumulation by ingestion of plastic-containing POPs ("carrier"), and decreased bioaccumulation by ingestion of clean plastic ("cleaning"). The model is parametrized for the lugworm Arenicola marina and evaluated against recently published bioaccumulation data for this species from laboratory bioassays with polystyrene microplastic. Further scenarios include polyethylene microplastic, nanosized plastic, and open marine systems. Model analysis shows that plastic with low affinity for POPs such as polystyrene will have a marginal decreasing effect on bioaccumulation, governed by dilution. For stronger sorbents such as polyethylene, the dilution, carrier, and cleaning mechanism are more substantial. In closed laboratory bioassay systems, dilution and cleaning dominate, leading to decreased bioaccumulation. Also in open marine systems a decrease is predicted due to a cleaning mechanism that counteracts biomagnification. However, the differences are considered too small to be relevant from a risk assessment perspective.

Marine fisheries as ecological experiments

There are many examples of ecological theory informing fishery management. Yet fisheries also provide tremendous opportunities to test ecological theory through large-scale, repeated, and well-documented perturbations of natural systems. Although treating fisheries as experiments presents several challenges, few comparable tests exist at the ecosystem scale. Experimental manipulations of fish populations in lakes have been widely used to develop and test ecological theory. Controlled manipulation of fish populations in open marine systems is rarely possible, but fisheries data provide a valuable substitute for such manipulations. To highlight the value of marine fisheries data, we review leading ecological theories that have been empirically tested using such data. For example, density dependence has been examined through meta-analysis of spawning stock and recruitment data to show that compensation (higher population growth) occurs commonly when populations are reduced to low levels, while depensation (the Allee effect) is rare. As populations decline, spatial changes typically involve populations contracting into high-density core habitats while abandoning less productive habitats. Fishing down predators may result in trophic cascades, possibly shifting entire ecosystems into alternate stable states, although alternate states can be maintained by both ecological processes and continued fishing pressure. Conversely, depleting low trophic level groups may affect central-place foragers, although these bottom–up effects rarely appear to impact fish—perhaps because many fish populations have been reduced to the point that they are no longer prey limited. Fisheries provide empirical tests for diversity–stability relations: catch data suggest that more diverse systems recover faster and provide more stable returns than less diverse systems. Fisheries have also provided examples of the tragedy of the commons, as well as counter-examples where common property resources have been managed successfully. We also address two barriers to use of fisheries data to answer ecological questions: differences in terminology for similar concepts and misuse of records of fishery landings (catch data) as a proxy for biomass trends.

Fish as major carbonate mud producers and missing components of the tropical carbonate factory

Carbonate mud is a major constituent of recent marine carbonate sediments and of ancient limestones, which contain unique records of changes in ocean chemistry and climate shifts in the geological past. However, the origin of carbonate mud is controversial and often problematic to resolve. Here we show that tropical marine fish produce and excrete various forms of precipitated (nonskeletal) calcium carbonate from their guts ("low" and "high" Mg-calcite and aragonite), but that very fine-grained (mostly < 2 μm) high Mg-calcite crystallites (i.e., > 4 mole % MgCO(3)) are their dominant excretory product. Crystallites from fish are morphologically diverse and species-specific, but all are unique relative to previously known biogenic and abiotic sources of carbonate within open marine systems. Using site specific fish biomass and carbonate excretion rate data we estimate that fish produce ∼6.1 × 10(6) kg CaCO(3)/year across the Bahamian archipelago, all as mud-grade (the < 63 μm fraction) carbonate and thus as a potential sediment constituent. Estimated contributions from fish to total carbonate mud production average ∼14% overall, and exceed 70% in specific habitats. Critically, we also document the widespread presence of these distinctive fish-derived carbonates in the finest sediment fractions from all habitat types in the Bahamas, demonstrating that these carbonates have direct relevance to contemporary carbonate sediment budgets. Fish thus represent a hitherto unrecognized but significant source of fine-grained carbonate sediment, the discovery of which has direct application to the conceptual ideas of how marine carbonate factories function both today and in the past.

Composition and seasonal variability of picoeukaryote communities at two subtropical coastal sites with contrasting trophic conditions

Phylogenetic diversity (18S rRNA gene) of picoeukaryotes in Hong Kong coastalwaters is dominated by heterotrophic forms and influenced by the seasonallydriven combination of freshwater discharge from the Pearl River, oceanic water ofthe South China Sea and coastal water from the China Coastal Current.KEYWORDS: picoeukaryote; subtropical coastal waters; 18S-rDNA; DGGE;seasonal variationPicoeukaryotes, small eukaryotic plankton with diam-eter ,3 mm, have ubiquitous distribution and areimportant contributors to global primary productionand carbon and mineral cycles (Li, 1994). They arephylogenetically, morphologically and metabolicallydiverse and are capable of phototrophy, phagotrophy,osmotrophy and mixotrophy (Moreira and Lo´pez-Garci´a, 2002).Traditionally, morphological features have been usedfor picoeukaryote identification and classification.However, those features alone are not sufficient to dis-tinguish clearly among species or groups of picoeukar-yotes due to their tiny sizes and cryptic morphologicalcharacteristics (Sims et al., 2002), leading to underesti-mation of the diversity at phylum level (Massana et al.,2002). In contrast, molecular approaches based on the18S rRNA gene provide new insights into picoeukaryotecommunity structure in various ecosystems. Over thepast several years, studies have demonstrated tremen-dous molecular phylogenetic diversity among theseorganisms in open marine systems, including the equa-torial Pacific Ocean (Moon-van der Staay et al., 2001),the Antarctic Polar Front (Lo´pez-Garci´a et al., 2001),the Mediterranean and Scotia Sea as well as the NorthAtlantic Ocean (Di´ez et al., 2001b). A large number ofnovel picoeukaryote phylogenetic groups, some without

Order of arrival affects competition in two reef fishes

Many communities experience repeated periods of colonization due to seasonally regenerating habitats or pulsed arrival of young-of-year. When an individual's persistence in a community depends upon the strength of competitive interactions, changes in the timing of arrival relative to the arrival of a competitor can modify competitive strength and, ultimately, establishment in the community. We investigated whether the strength of intracohort competitive interactions between recent settlers of the reef fishes Thalassoma hardwicke and T. quinquevittatum are dependent on the sequence and temporal separation of their arrival into communities. To achieve this, we manipulated the sequence and timing of arrival of each species onto experimental patch reefs by simulating settlement pulses and monitoring survival and aggressive interactions. Both species survived best in the absence of competitors, but when competitors were present, they did best when they arrived at the same time. Survival declined as each species entered the community progressively later than its competitor and as aggression by its competitor increased. Intraspecific effects of resident T. hardwicke were similar to interspecific effects. This study shows that the strength of competition depends not only on the identity of competitors, but also on the sequence and timing of their interactions, suggesting that when examining interaction strengths, it is important to identify temporal variability in the direction and magnitude of their effects. Furthermore, our findings provide empirical evidence for the importance of competitive lotteries in the maintenance of species diversity in demographically open marine systems.

Connectivity, regime shifts and the resilience of coral reefs

Connectivity of larvae among metapopulations in open marine systems can be a double-edged sword, allowing for the colonization and replenishment of both desirable and undesirable elements of interacting species-rich assemblages. This article studies the effect of recruitment by coral and macroalgae on the resilience of grazed reef ecosystems. In particular, we focus on how larval connectivity affects regime shifts between alternative assemblages that are dominated either by corals or by macroalgae. Using a model with bistability dynamics, we show that recruitment of coral larvae erodes the resilience of a macroalgae-dominated ecosystem when grazing is high, but has negligible effect when grazing is low. Conversely, recruitment by macroalgae erodes the resilience of a coral-dominated ecosystem when grazing is low, leading to a regime shift to macroalgae. Thus, spillover of coral recruits from highly protected areas will not restore coral cover or prevent flips to macroalgae in the surrounding seascape if grazing levels in these areas are depleted, but may be pivotal for re-building coral populations if grazing is high. Fishing restrictions and the re-introduction of herbivores should therefore be a prime conservation objective for preventing undesirable regime shifts. Connectivity by some components of coral reef assemblages (e.g., macroalgae, pathogens, crown-of-thorns starfish) may be detrimental to sustaining reefs, especially where overfishing and other drivers have eroded their resilience, making them more vulnerable to a regime shift.

Temporal changes in the strength of density‐dependent mortality and growth in intertidal barnacles

1. In demographically open marine systems, the extent to which density-dependent processes in the benthic adult phase are required for population persistence is unclear. At one extreme, represented by the recruitment limitation hypothesis, larval supply may be insufficient for the total population size to reach a carrying capacity and density-independent mortality predominates. At the opposite extreme, populations are saturated and density-dependent mortality is sufficiently strong to reshape patterns established at settlement. 2. We examined temporal variation in the way density-independent and density-dependent mortality interact in a typical sessile marine benthic invertebrate, the acorn barnacle Semibalanus balanoides (L.), over a 2-year period. 3. Recruitment was manipulated at two high recruitment sites in north Wales, UK to produce recruit densities covering the range naturally found in this species. Following manipulation, fixed quadrats were monitored using digital photography and temporal changes in mortality and growth rate were examined. 4. Over a 2-year period there was a clear, spatially consistent, over-compensatory relationship between the density of recruits and adult abundance indicating strong density-dependent mortality. The strength of density dependence intensified with increasing recruitment. 5. Density-dependent mortality did not operate consistently over the study period. It only operated in the early part of the benthic phase, but the pattern of adult abundance generated was maintained throughout the whole 2-year period. Thus, early life-history processes dictated adult population abundance and dynamics. 6. Examination of the natural recruitment regime in the area of study indicated that both positive and negative effects of recruitment will occur over scales varying from kilometres to metres.

Nutrient conditions in the Yellow Sea and the East China Sea

This paper discusses the distributions of micronutrients in the Yellow Sea (YS) and the East China Sea (ECS), based on the data obtained during field observations. It is pointed out that there are abundant nutrients in the area of the east and northeast of the Changjiang River estuary throughout the year. It is believed that the nutrients come from the extension of the Changjiang River diluted water and the transportation of the Subei coastal water. The maximum extension range of the nutrients of the Changjiang River diluted water was observed during the catastrophic flooding period of the Changjiang River in the summer of 1998. Based on the Redfield ratio (Si:N:P = 16:16:1) at which three essential nutrients are utilized by marine phytoplankton, the Si/N/P ratios are calculated and studied for the upper water of the YS and the ECS. The results indicate that the Si/N ratios are very high, which suggests that silicate is not the limiting factor for the growth of phytoplankton in the YS and the ECS. Under the influence of terrestrial runoff, especially the Changjiang River runoff which is rich in combined nitrogen and has very high N/P ratios, high N/P ratios occur in the Changjiang estuary and its adjacent areas such as the south and southwest of the YS, the inshore area of the ECS and the area east of the Changjiang estuary in spring and summer. As a result, in contrast to general open marine systems, the systems in these areas resemble estuarine ones rather than typical marine ones. The primary production in a considerable portion of the YS and the ECS may be limited by phosphate rather than nitrogen.

Food webs and the transmission of parasites to marine fish.

Helminth parasites of fish in marine systems are often considered to be generalists, lacking host specificity for both intermediate and definitive hosts. In addition, many parasites in marine waters possess life cycles consisting of long-lived larval stages residing in intermediate and paratenic hosts. These properties are believed to be adaptations to the long food chains and the low densities of organisms distributed over broad spatial scales that are characteristic of open marine systems. Moreover, such properties are predicted to lead to the homogenization of parasite communities among fish species. Yet, these communities can be relatively distinct among marine fishes. For benthos, the heterogeneous horizontal distribution of invertebrates and fish with respect to sediment quality and water depth contributes to the formation of distinct parasite communities. Similarly, for the pelagic realm, vertical partitioning of animals with depth will lead to the segregation of parasites among fish hosts. Within each habitat, resource partitioning in terms of dietary preferences of fish further contributes to the establishment of distinct parasite assemblages. Parasite distributions are predicted to be superimposed on distributional patterns of free-living animals that participate as hosts in parasite life cycles. The purpose of this review is first, to summarize distribution patterns of invertebrates and fish in the marine environment and relate these patterns to helminth transmission. Second, patterns of transmission in marine systems are interpreted in the context of food web structure. Consideration of the structure and dynamics of food webs permits predictions about the distribution and abundance of parasites. Lastly, parasites that influence food web structure by regulating the abundance of dominant host species are briefly considered in addition to the effects of pollution and exploitation on food webs and parasite transmission.

DISEASE TRANSMISSION OF ASPERGILLOSIS IN SEA FANS: INFERRING PROCESS FROM SPATIAL PATTERN

Despite recent high impacts of disease in the ocean, quantitative studies of diseases in natural marine populations lag far behind terrestrial systems. Transmission processes are poorly known in relatively open marine systems. We studied infection of sea fans Gorgonia ventalina by the fungus Aspergillus sydowii. To assess detectability and mechanisms of secondary transmission of the fungus, we analyzed the spatial distribution of disease among the fans, using Ripley's K as a measure of disease aggregation. Coral populations and disease were mapped at three reefs in the Florida Keys. Where disease prevalence was low, the disease was distributed randomly among sea fans. This is consistent with disease transmission by input of infectious fungal material from terrestrial sources only. However, where disease prevalence was high, the disease was significantly aggregated among sea fans at the 2-8 m scale, consistent with secondary disease trans- mission. Such transmission could take place by physical contact between neighboring fans, or infected fans might shed fungal material into the water column, and other fans become infected by these fomites. Our results suggest that water-borne transmission occurs, but secondary transmission by physical contact is also likely. We cannot falsify the hypothesis that small-scale local environmental conditions also contribute to disease aggregation.