Genotypic distribution of a specialist model microorganism, Methanosaeta, along an estuarine gradient: does metabolic restriction limit niche differentiation potential?

A reductionist ecological approach of using a model genus was adopted in order to understand how microbial community structure is driven by metabolic properties. The distribution along an estuarine gradient of the highly specialised genus Methanosaeta was investigated and compared to the previously determined distribution of the more metabolically flexible Desulfobulbus. Methanosaeta genotypic distribution along the Colne estuary (Essex, UK) was determined by DNA- and RNA-based denaturing gradient gel electrophoresis and 16S rRNA gene sequence analyses. Methanosaeta distribution was monotonic, with a consistently diverse community and no apparent niche partitioning either in DNA or RNA analyses. This distribution pattern contrasts markedly with the previously described niche partitioning and sympatric differentiation of the model generalist, Desulfobulbus. To explain this difference, it is hypothesised that Methanosaeta's strict metabolic needs limit its adaptation potential, thus populations do not partition into spatially distinct groups and so do not appear to be constrained by gross environmental factors such as salinity. Thus, at least for these two model genera, it appears that metabolic flexibility may be an important factor in spatial distribution and this may be applicable to other microbes.     

Contrasting patterns of niche partitioning between two anaerobic terminal oxidizers of organic matter

Understanding the ecological principles underlying the structure and function of microbial communities remains an important goal for microbial ecology. We examined two biogeochemically important taxa, the sulfate-reducing bacterial genus, Desulfobulbus, and the methanogenic archaeal genus, Methanosaeta, to compare and contrast niche partitioning by these two taxa that are ecologically linked as anaerobic terminal oxidizers of organic material. An observational approach utilizing functional gene pyrosequencing was combined with a community-based reciprocal incubation experiment and characterization of a novel Desulfobulbus isolate. To analyze the pyrosequencing data, we constructed a data analysis pipeline, which we validated with several control data sets. For both taxa, particular genotypes were clearly associated with certain portions of an estuarine gradient, consistent with habitat or niche partitioning. Methanosaeta genotypes were generally divided between those found almost exclusively in the marine habitat (∼30% of operational taxonomic units (OTUs)), and those which were ubiquitously distributed across all or most of the estuary (∼70% of OTUs). In contrast to this relatively monotonic distribution, for Desulfobulbus, there were many more genotypes, and their distributions represented a wide range of inferred niche widths from specialist genotypes found only at a single site, to ubiquitous or generalist genotypes found in all 10 sites examined along the full estuarine gradient. Incubation experiments clearly showed that, for both taxa, communities from opposite ends of the estuary did not come to resemble one another, regardless of the chemical environment. Growth of a Desulfobulbus isolated into pure culture indicated that the potential niche of this organism is significantly larger than the realized niche. We concluded that niche partitioning can be an important force structuring microbial populations, with biotic and abiotic components having very different effects depending on the physiology and ecology of each taxon.     

Spatial and temporal variability of early post-settlement survivorship and growth in the barnacle Balanus glandula along an estuarine gradient

The performance of a species can be significantly altered by subtle changes in the physical environmental. The intertidal barnacle Balanus glandula is predominantly an open coast species in the Northeast Pacific. However, B. glandula commonly inhabits estuaries where environmental conditions such as salinity and temperature drastically differ from the open coast. We used survivorship and growth rates as a measure of performance in recently metamorphosed laboratory reared juvenile B. glandula outplanted along an environmental gradient at the mouth, mid-estuarine, and riverine end of the South Slough Estuary, Oregon, USA. Juvenile performance was highly variable over spatial and temporal scales and dependent upon existing environmental conditions. Surprisingly, along this estuarine gradient, juveniles performed better at a mid-estuarine location than at the mouth of the estuary. Typically, the riverine end of the estuary was the least suitable habitat along the estuarine gradient due to high juvenile mortality and a low growth rate. Although seasonally variable, survivorship and growth decreased with height along a vertical intertidal gradient as well. In a reciprocal transplant experiment, populations from both ends of the estuarine gradient displayed similar survivorship and growth rates. Our results demonstrate that the interactions of environmental conditions that vary temporally and spatially along a gradient strongly affect the success of an individual surviving and prospering during the early juvenile period.     

Distribution patterns of the fish assemblage in the Mamanguape River Estuary,North-eastern Brazil

The Mamanguape River Estuary was studied along a continuum ranging from shallow sandstone reefs adjacent to the river mouth up to the limit of influence of seawater, the upper portions of the estuary. Fish samples were gathered using three types of nets along 17 sample sites, grouped in four regions according to salinity range (reefs and low, middle and upper estuaries), to seasonality (dry and rainy seasons) and to habitat usage (marine visitor, marine estuarine-opportunist, marine estuarine-dependent, estuarine resident and estuarine &; marine). Differences were found in the fish assemblages along the estuarine-reef gradient, with most species (n?=?30) being considered marine, estuarine-opportunists or estuarine-dependent, according to its abundance and distribution. A low number of species (n?=?11) were considered estuarine residents. Some species exhibited significant differences in spatial distribution pattern, in which juveniles and adults predominated in different portions of the estuary, suggesting an ontogenetic migration both in relation to the adjacent reef area and across estuarine regions. Several species were newly recorded in the Mamanguape Estuary: Anchoa spinifer, Halichoeres poeyi, Hyporhamphus roberti, Scomberomorus cavalla, Sphyraena barracuda and Ocgocephalus vesperilio.     

Diversity and spatio-temporal distribution of ammonia-oxidizing Archaea and Bacteria in sediments of the Westerschelde estuary

The diversity and spatio-temporal distribution of ammonia-oxidizing Archaea (AOA) and Bacteria (AOB) were investigated along a salinity gradient in sediments of the Westerschelde estuary. Sediment samples were collected from three sites with different salinities, and at six time points over the year. Denaturing gradient gel electrophoresis of PCR-amplified 16S rRNA and amoA gene fragments was used to identify the AOA and AOB present. Members of the AOA were mainly belonging to the Crenarchaeota Group 1, which were found at all sites, while members of the genus Nitrosomonas, which were abundant at the brackish sites, and of the genus Nitrosospira, which were present in early spring at the marine sites, were found to be the dominant AOB. Statistical analysis indicated that salinity and temperature were the main factors controlling the diversity and distribution of both AOA and AOB. Variability in net primary production rates was also correlated with species composition of both groups, but changes in the nitrite concentration only to the distribution of the AOA.     

Zonation of intertidal macrobenthos in the estuaries of Schelde and Ems

Based on data, collected in 1980–1990, the intertidal benthic macrofauna of the Schelde and Ems estuaries was compared. The spatial occurrence of the benthic macrofauna along the salinity gradient, including the freshwater tidal area was emphasized. Both estuaries appeared to have a very similar species composition, especially at genus level. The higher number of species observed in the Schelde estuary was probably due to a greater habitat diversity. In both estuaries species diversity decreased with distance upstream. The total density did not vary along the estuarine gradient, whereas biomass is highest in the polyhaline zone.In both estuaries distinct intertidal benthic communities were observed along the salinity gradient: a marine community in the polyhaline zone, a brackish community in the mesohaline zone, and a third community in the oligohaline and freshwater tidal zones of the estuary. These three communities were very similar between both estuaries. Their main characteristics were discussed together with the occurrence and distribution of the dominant species.For the Schelde estuary and to a lesser extent also for the Ems estuary, there was evidence that anthropogenic stress had a negative effect on the intertidal macrobenthic communities of the oligohaline/freshwater tidal zone. Only Oligochaeta were dominating, whereas the very euryhaline and/or true limnetic species were missing. In the mesohaline zone, the Schelde estuary was dominated by large numbers of short-living, opportunistic species, whereas in the Ems estuary relatively more stable macrobenthic communities were observed. A comparison with some other European estuaries showed in general similar trends as those observed for the Schelde and Ems estuaries.     

Short and long-term movement and site fidelity of juvenile Haemulidae in back-reef habitats of a Caribbean embayment

In the context of a main project that aims to recover modern data on diatom distribution applicable to paleosalinity reconstructions in coastal areas of Southern South America, the composition and distribution of dead diatom assemblages in the littoral zone of the Quequén Salado estuary (Argentina) were studied. Diatom zones were defined along the estuarine gradient by cluster analysis and related to the salinity range and sediment composition by Canonical Correspondence Analysis. Four diatom zones were identified. A mixture of marine, brackish and freshwater diatoms, probably allochthonous, characterized the inlet (zone I). Marine/brackish taxa, represented mainly by Paralia sulcata dominated zone II, characterized by polyhaline conditions and sandy sediments. Zone III was characterized by mesohaline conditions, muddy sediments and the dominance of the estuarine diatom Amphora helenensis. Brackish/freshwater and freshwater diatoms dominated the headwaters (zone IV), where salinity was always below 5‰. The comparison of Quequén Salado diatom assemblages with previous results from the Quequén Grande estuary showed a similar taxonomic composition between both estuaries. However, differences in the salinity ranges of the estuaries (related to differences in the degree of human impact and tidal range) lead to a displacement in their spatial distribution along the longitudinal estuarine axis. This paper contributes to the knowledge of the ecological requirements of South American estuarine diatoms and provides useful data for paleosalinity reconstructions in the region. Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users Handling editor: J. Padisak     

Estuarine recruitment of a marine goby reconstructed with an isotopic clock

Information on movement patterns of marine fishes between estuarine populations and stocks at sea is fundamental to understanding their population dynamics, life history tactics and behavior. Furthermore, understanding estuarine habitat use by marine fishes is crucial for their effective conservation and integrated estuarine management. Although large numbers of young marine fish make use of temperate estuaries in highly predictable abundance patterns, very little is known about how estuarine populations interact with the populations at sea. Recruitment of sand goby Pomatoschistus minutus (Pallas, 1770) into the low salinity zone of the Scheldt estuary (Belgium) was reconstructed over an entire year by means of an isotopic clock. These results were combined with a growth model to yield age and length at immigration. Sand gobies entered the upper Scheldt estuary almost continuously from May onwards, except in July when they appeared to avoid the estuary due to warm summer temperatures. About 70% of the fish caught in the upper estuary resided there for less than 1 month, which indicates a strong temporal overlap of immigration and emigration. This complex migration pattern suggests that estuarine residence is caused by trade-offs made at the individual level, whereby migration is probably triggered by temperature. The high turnover of individuals in the estuarine population leads us to question the functional role of the estuary for marine fishes. Sand gobies entering the upper estuary had a wide range of ages and body sizes, although they were at least 2 months old and had a minimum standard length of approximately 20 mm. This study shows that the use of an isotopic clock strongly complements catch data and is useful to describe the connectivity between populations.     

Distribution of thermophilic endospores in a temperate estuary indicate that dispersal history structures sediment microbial communities

Endospores of thermophilic bacteria are found in cold and temperate sediments where they persist in a dormant state. As inactive endospores that cannot grow at the low ambient temperatures, they are akin to tracer particles in cold sediments, unaffected by factors normally governing microbial biogeography (e.g., selection, drift, mutation). This makes thermophilic endospores ideal model organisms for studying microbial biogeography since their spatial distribution can be directly related to their dispersal history. To assess dispersal histories of estuarine bacteria, thermophilic endospores were enriched from sediments along a freshwater‐to‐marine transect of the River Tyne in high temperature incubations (50°C). Dispersal histories for 75 different taxa indicated that the majority of estuarine endospores were of terrestrial origin; most closely related to bacteria from warm habitats associated with industrial activity. A subset of the taxa detected were marine derived, with close relatives from hot deep marine biosphere habitats. These patterns are consistent with the sources of sediment in the River Tyne being predominantly terrestrial in origin. The results point to microbial communities in estuarine and marine sediments being structured by bi‐directional currents, terrestrial run‐off and industrial effluent as vectors of passive dispersal and immigration.     

Microbial biogeography along an estuarine salinity gradient: combined influences of bacterial growth and residence time

Shifts in bacterioplankton community composition along the salinity gradient of the Parker River estuary and Plum Island Sound, in northeastern Massachusetts, were related to residence time and bacterial community doubling time in spring, summer, and fall seasons. Bacterial community composition was characterized with denaturing gradient gel electrophoresis (DGGE) of PCR-amplified 16S ribosomal DNA. Average community doubling time was calculated from bacterial production ([(14)C]leucine incorporation) and bacterial abundance (direct counts). Freshwater and marine populations advected into the estuary represented a large fraction of the bacterioplankton community in all seasons. However, a unique estuarine community formed at intermediate salinities in summer and fall, when average doubling time was much shorter than water residence time, but not in spring, when doubling time was similar to residence time. Sequencing of DNA in DGGE bands demonstrated that most bands represented single phylotypes and that matching bands from different samples represented identical phylotypes. Most river and coastal ocean bacterioplankton were members of common freshwater and marine phylogenetic clusters within the phyla Proteobacteria, Bacteroidetes, and ACTINOBACTERIA: Estuarine bacterioplankton also belonged to these phyla but were related to clones and isolates from several different environments, including marine water columns, freshwater sediments, and soil.     

Benthic meiofauna as indicator of ecological changes in estuarine ecosystems: The use of nematodes in ecological quality assessment

Estuarine meiofauna communities have been only recently considered to be good indicators of ecological quality, exhibiting several advantages over macrofauna, such as their small size, high abundance, rapid generation times and absence of a planktonic phase. In estuaries we must account not only for a great natural variability along the estuarine gradient (e.g. sediment type and dynamics, oxygen availability, temperature and flow speed) but also for the existence of anthropogenic pressures (e.g. high local population density, presence of harbors and dredging activities).Spatial and temporal biodiversity patterns of meiofauna and free-living marine nematodes were studied in the Mondego estuary (Portugal). Both taxonomic and functional approaches were applied to nematode communities in order to describe the community structure and to relate it with the environmental parameters along the estuary. At all sampling events, nematode assemblages reflected the estuarine gradient, and salinity and grain size composition were confirmed to be the main abiotic factors controlling the distribution of the assemblages.Moreover, the low temporal variability may indicate that natural variability is superimposed by the anthropogenic pressures present in some areas of the estuary. The characterization of both meiofauna and nematode assemblages highlighted the usefulness of the integration of both taxonomic and functional attributes, which must be taken into consideration when assessing the ecological status of estuaries.     

Spatial and seasonal variations in fish assemblages of the Yangtze River estuary

This study aimed to test the hypothesis that biotic influences play a role in determining estuarine fish assemblages. Thus the mechanisms that regulate the observed spatial niche segregation between fish species with morphological and ecological similarities within estuaries were investigated. Fishes were sampled seasonally at 30 stations along an entire salinity gradient of the Yangtze River estuary from spring 2010 to winter 2011, using bottom trawling (10 mm mesh size in the cod end). A total of 62 species belonging to 28 families were collected. Marine migrants (21) and estuarine species (19) dominated the assemblage and accounted for the highest abundances (marine migrants = 42.5%, estuarine species = 38.3%), with the marine migrants having the highest biomass (57.2%). Canonical correspondence analyses indicated that chlorophyll a, salinity, temperature, and geographic distance were the four main variables influencing the occurrence of fishes within the system. A stable fish assemblage in the upper zone during the wet season (spring and summer) was more obvious than in the dry season. The Pianka index showed a clear spatial segregation in four pairs of tonguesoles species (Cygonobius), and in most pairs of gobiids (Gobiidae), and a high degree of overlapping (>0.60) in only four pairs of gobiid species during the wet season. However, according to the results of the null model the observed segregating or pattern sharing was not caused by interspecific competition.     

Fish community response to increased river flow in the Kariega Estuary,a freshwater-deprived,permanently open southern African system

The littoral and demersal ichthyofaunal community structure in the freshwater-deprived, permanently open Kariega Estuary was investigated following heavy rain in November 2006 and was compared to low-flow condition data from 1991 and 1996. All surveys took place during the spring months and allowed for a comparison of a wet and a dry spring period. The 2006 freshwater pulse generated a strong horizontal salinity gradient within the estuary. In the absence of freshwater inflow, the ichthyofaunal community in the littoral zone was numerically dominated by estuarine resident species, whilst after the freshwater pulse an increased contribution of marine migrant species was observed. Within the demersal zone, marine straggler species dominated during the dry spring period and estuarine residents during the wet spring period. Numerical analyses of the littoral and demersal fish assemblages indicated the presence of three distinct groupings — corresponding to the upper and middle reaches during separate wet and dry periods, and a community associated with the lower reaches of the estuary. It is suggested that the shift in community structure between the dry and wet spring periods could be related to altered physico-chemical and trophic conditions within the estuary, as well as the increased presence of freshwater and estuarine olfactory cues within the coastal zone, which would have resulted in the recruitment of 0+ estuary-associated marine species into the Kariega system.     

Relative abundance and diversity of ammonia-oxidizing archaea and bacteria in the San Francisco Bay estuary

Ammonia oxidation in marine and estuarine sediments plays a pivotal role in the cycling and removal of nitrogen. Recent reports have shown that the newly discovered ammonia-oxidizing archaea can be both abundant and diverse in aquatic and terrestrial ecosystems. In this study, we examined the abundance and diversity of ammonia-oxidizing archaea (AOA) and betaproteobacteria (beta-AOB) across physicochemical gradients in San Francisco Bay--the largest estuary on the west coast of the USA. In contrast to reports that AOA are far more abundant than beta-AOB in both terrestrial and marine systems, our quantitative PCR estimates indicated that beta-AOB amoA (encoding ammonia monooxygenase subunit A) copy numbers were greater than AOA amoA in most of the estuary. Ammonia-oxidizing archaea were only more pervasive than beta-AOB in the low-salinity region of the estuary. Both AOA and beta-AOB communities exhibited distinct spatial structure within the estuary. AOA amoA sequences from the north part of the estuary formed a large and distinct low-salinity phylogenetic group. The majority of the beta-AOB sequences were closely related to other marine/estuarine Nitrosomonas-like and Nitrosospira-like sequences. Both ammonia-oxidizer community composition and abundance were strongly correlated with salinity. Ammonia-oxidizing enrichment cultures contained AOA and beta-AOB amoA sequences with high similarity to environmental sequences. Overall, this study significantly enhances our understanding of estuarine ammonia-oxidizing microbial communities and highlights the environmental conditions and niches under which different AOA and beta-AOB phylotypes may thrive.     

Phylogenetic analysis of particle-attached and free-living bacterial communities in the Columbia river, its estuary, and the adjacent coastal ocean.

The Columbia River estuary is a dynamic system in which estuarine turbidity maxima trap and extend the residence time of particles and particle-attached bacteria over those of the water and free-living bacteria. Particle-attached bacteria dominate bacterial activity in the estuary and are an important part of the estuarine food web. PCR-amplified 16S rRNA genes from particle-attached and free-living bacteria in the Columbia River, its estuary, and the adjacent coastal ocean were cloned, and 239 partial sequences were determined. A wide diversity was observed at the species level within at least six different bacterial phyla, including most subphyla of the class Proteobacteria. In the estuary, most particle-attached bacterial clones (75%) were related to members of the genus Cytophaga or of the alpha, gamma, or delta subclass of the class Proteobacteria. These same clones, however, were rare in or absent from either the particle-attached or the free-living bacterial communities of the river and the coastal ocean. In contrast, about half (48%) of the free-living estuarine bacterial clones were similar to clones from the river or the coastal ocean. These free-living bacteria were related to groups of cosmopolitan freshwater bacteria (beta-proteobacteria, gram-positive bacteria, and Verrucomicrobium spp.) and groups of marine organisms (gram-positive bacteria and alpha-proteobacteria [SAR11 and Rhodobacter spp.]). These results suggest that rapidly growing particle-attached bacteria develop into a uniquely adapted estuarine community and that free-living estuarine bacteria are similar to members of the river and the coastal ocean microbial communities. The high degree of diversity in the estuary is the result of the mixing of bacterial communities from the river, estuary, and coastal ocean.     

Tracing Mississippi River influences in estuarine food webs of coastal Louisiana

The Breton Sound estuary in southern Louisiana receives large amounts of Mississippi River water via a controlled diversion structure at the upstream end of the estuary. We used stable isotopes to trace spatial and seasonal responses of the downstream food web to winter and spring introductions of river water. Analysis of δ¹³C, δ¹⁵N, and δ³⁴S in the common local consumers such as grass shrimp (Palaemonetes sp.), barnacles (Balanus sp.), and small plankton-feeding fish (bay anchovies, Anchoa mitchilli) showed that the diversion was associated with two of the five major source regimes that were supporting food webs: a river regime near the diversion and a river-influenced productive marsh regime farther away from the diversion. Mixing models identified a third river-influenced source regime at the marine end of the estuary where major natural discharge from the Bird’s Foot Delta wraps around into estuarine waters. The remaining two source regimes represented typical estuarine conditions: local freshwater sources especially from precipitation and a brackish source regime representing higher salinity marine influences. Overall, the Mississippi River diversion accounted for 75% of food web support in the upper estuary and 25% in the middle estuary, with influence strongest along known flow pathways and closest to the diversion. Isotopes also traced seasonal changes in river contributions, and indicated increased plant community productivity along the major flow path of diversion water. In the Breton Sound estuary, bottom–up forcing of food webs is strongly linked to river introductions and discharge, occurring in spatial and temporal patterns predictable from known river input regimes and known hydrologic circulation patterns.     

Effects of the river discharge management on the nursery function of the Guadalquivir river estuary (SW Spain)

Within the Guadalquivir estuary, young recruits of marine species seem to respond to changes in freshwater flow by moving with the mass of estuarine water that is most “suitable” for them. The control of the river flow, from a dam 110 km upstream from the river mouth, has an immediate effect on the estuarine salinity gradient, displacing it either seaward or upstream. Consequently, there is a reduction or enlargement of the estuarine area that is used as nursery grounds. The analysis of the temporal estuarine recruitment and spatial distribution of young stages of marine species, during six annual cycles, provides evidence that the estuarine zone used as nursery grounds is mainly that part situated seaward from an isohaline value of 5. The relationship between the position of that isohaline (D₅) and the freshwater discharges from the dam was also examined during high and low tides. It was found that a high percentage of the isohaline position variation (75% and 73% at high and low tides, respectively) can be explained by the freshwater volume discharged from the dam during the previous week. These preliminary results suggest that an accurate model of the relationship between these two variables may be a useful tool for future management strategies of freshwater discharges to the estuary.     

Geographical and typological changes in fish guilds of South African estuaries

A comparative analysis of fish estuary association guilds was undertaken on some 190 South African estuaries. This pioneering study spanned three zoogeographic regions and included three broad estuarine types. The guild compositions of the estuaries were compared based on an importance value, incorporating taxonomic composition, numerical abundance and relative biomass. Multivariate analyses included both inter‐regional (zoogeographic) and intra‐regional (estuarine typology) comparisons. The major estuary‐associated guilds (estuarine species and marine migrant species) were important in all estuary types within all biogeographic regions. Significant differences both between regions and between estuary types within regions, however, were recorded. Cool–temperate estuaries were generally dominated by migratory species (estuarine migrants and marine migrant opportunists) while the importance of species dependent on estuaries (estuarine residents and estuarine‐dependent marine migrants) was higher in warm–temperate and subtropical regions. The significance of estuarine nursery areas, particularly in regions where estuaries are few in number, is highlighted. In terms of typology, migratory species assumed a greater importance in predominantly open systems, while freshwater and estuarine‐resident species were more important in predominantly closed systems. Predominantly closed estuaries, however, were also important for marine migrant species, which further highlights the significance of these systems as nursery areas for fishes.     

Influence of spatial and temporal variations on organic pollutant biodegradation rates in an estuarine environment.

The influence of spatial and temporal environmental variations on rates of organic pollutant biodegradation were assessed by using heterotrophic uptake kinetics. These studies were conducted at three sites, representing the gradient from freshwater to estuarine to marine systems. Of the compounds tested, total uptake Vmax rates decreased in the order of nitrilotriacetic acid, m-cresol, chlorobenzene, and 1,2,4-trichlorobenzene. In general, the freshwater site exhibited the highest uptake rates, with somewhat lower rates at the estuarine site. Rates at the marine site were much lower than at the other sites, except during the winter. Metabolic rates at both the freshwater and estuarine areas were significantly decreased during periods of low water temperature. Rates at the marine site were relatively uniform throughout the year. Linear regression analysis was used to compare m-cresol biodegradation rates to characteristics of the microbial community, which included direct microscopic counts, CFU counts, and cellular incorporation of amino acids. The observed rates did not consistently correlate well with any of the measured characteristics of the microbial community.