The Influence of Dry-season Conditions on the Bottom Dwelling Fauna of an East Australian Sub-tropical Estuary

Spatial patterns in benthic infaunal community structure of Port Curtis estuary (north-eastern Australia) were determined from quantitative grab samples and examined in relation to environmental variables. A total of 149 riverine, estuarine and open coastal stations were sampled during the winter survey, and 5744 individuals from 466 species identified. Filter-feeding organisms (primarily polychaetes, molluscs and crustaceans) dominated the benthos, and accounted for 50% of the total species abundance and 30% of the total species richness. Most taxa were uncommon, and 98% of species individually represented less than 2% of the total abundance. Distributional patterns in total species richness and abundance were highly correlated, and both parameters varied significantly with sediment grain size. Numbers of species and individuals were typically highest in coarse-sand and gravel sediments, and were lowest in fine, well-sorted, sands. Cluster analysis of species abundance data revealed nine community groupings characterised by small species sub-sets with restricted distributions. These groupings were primarily related to sediment grain size, sediment organic carbon content, salinity and depth. Not all variation in community structure was explained by these environmental variables, and it is likely that other unmeasured factors play an important role in determining benthic faunal composition in the estuary.     

Assessment of Littoral Benthic Invertebrate Communities at the Land–Water Interface in Lakes Recovering from Severe Acid- and Metal-Damage

Benthic invertebrate communities within confluence sites, or areas of sediment deposition, are shaped by the input of catchment products including coarse woody debris, organic and inorganic particulates, and contaminants, but these sites also appear to be potential “hotspots” where recolonization of severely damaged ecosystems begins. Two species of leaf packs and a sweep netting technique were used to assess benthic invertebrate communities across a gradient of 14 confluence sites in 3 recovering lakes near the copper and nickel smelters in Sudbury, Canada. Environmental variables including delta habitat composition, delta area and length, and composition of deposited materials were used to detect spatial patterns in littoral benthic invertebrate communities. Benthic invertebrate community relationships with water chemistry were also assessed. Partial redundancy analysis (pRDA) showed that all sampling methods detected similar gradients of increasing invertebrate community richness and diversity as area and length of the sediment delta and the surface organic matter abundance increased. Two-way nested ANOVAs showed significant differences (p < .05) in taxa richness and diversity metrics among sites. Of the three methods, the benthic invertebrate community measurements from the birch leaf packs provided the strongest correlations with measures of organic matter inputs or habitat characteristics of the confluence zones. These correlations suggest that tree planting in riparian areas, or organic matter or macrophyte additions to littoral zones, may enhance littoral benthic invertebrate richness and diversity in acid and metal damaged lakes.     

Effects of hypoxia and organic enrichment on estuarine macrofauna in the inner part of Ariake Bay

In the inner part of Ariake Bay, Japan, hypoxia frequently occurs in summer at the organically enriched bottom with salinity stratification caused by flooding after the rainy season. Sediment organic enrichment can work as a stressor for macrobenthos. To investigate the effects of both hypoxia and sediment organic enrichment on macrobenthos, samples were collected at 20 stations by grab sampling in May and August, representing the situation before and after hypoxia, respectively. Although sediment grain size did not change, sediment TOC increased significantly in August. Multivariate analyses showed that the community structure changed significantly in August. The variation in the community structure among stations also increased, which indicated disturbance by stressors during the study period. Similarly, the species richness and total abundance of macrobenthos decreased significantly after hypoxia even after the TOC effect was removed. In addition, the amount of TOC change and the abundance of the main species did not correlate in any case. These results suggest that the community changes during the study period are not caused by stress from the increased sediment TOC but are mainly from the subsequent hypoxic stress.     

Estuarine and scalar patterns of invasion in the soft-bottom benthic communities of the San Francisco Estuary

The spatial patterns of nonindigenous species in seven subtidal soft-bottom communities in the San Francisco Estuary were quantified. Sixty nonindigenous species were found out of the 533 taxa enumerated (11%). Patterns of invasion across the communities were evaluated using a suite of invasion metrics based on the abundance or species richness of nonindigenous species. Patterns of invasion along the estuarine gradient varied with the invasion metric used, and the ecological interpretation of the metrics is discussed. Overall, the estuarine transition community located in the estuarine turbidity maximum zone (mean 5 practical salinity unit (psu)), main estuarine community (mean 16 psu), and marine muddy community (mean 28 psu) were more invaded than two fresh-brackish communities (mean < 1 psu) and a marine sandy community (mean 27 psu). Nonindigenous species were numerically dominant over much of the Estuary, making up more than 90% of the individuals in two communities. The percentage of the total species composed of nonindigenous species increased at smaller spatial scales: 11% at the estuary (gamma) scale, 21% at the community (alpha) scale, and 42% at the grab (point) scale. Wider spatial distributions of nonindigenous species and a relatively greater percentage of rare native species may have resulted in this pattern. Because of this scale dependency, comparisons among sites need to be made at the same spatial scale. Native species were positively correlated with nonindigenous species in several of the communities, presumably due to similar responses to small-scale differences in habitat quality. The rate of invasion into the soft-bottom communities of the San Francisco Estuary appears to have increased over the last one to two decades and many of the new introductions have become numerically dominant.     

Latitudinal variation in spongivorous fishes and the effectiveness of sponge chemical defenses

It has been proposed that predation pressure declines with increasing latitude and a positive correlation exists between predation intensity and the investment into chemical defenses. However, little direct evidence supports the idea that tropical species are better defended chemically than their temperate counterparts. Temperate reefs of the South Atlantic Bight (SAB) off Georgia, USA, provide a unique opportunity to study tropical sponges in a temperate environment. We documented sponge species richness and abundance, sponge predator density, and examined the ability of eight sponge species to chemically deter predation by fishes on two reefs in the SAB. We used rarefaction analysis and ANOVA to compare our results for sponge species richness and density, respectively, with similar published studies conducted on reefs of the sub-tropical Atlantic (i.e., Florida Keys). These analyses were combined with similar statistical comparisons for spongivorous fish species richness and density. Results showed that sponge species richness was lower, but sponge density was higher, on the temperate SAB reefs than on the subtropical reefs. Both spongivorous fish diversity and density were lower on the SAB reefs. The greater abundance of sponges and lower density of predators on SAB reefs suggest a lower frequency of predation on sponges on SAB reefs. Of the eight sponge species assayed from the SAB reefs, five possessed chemical extracts that were significantly less deterrent to fish predators than their tropical/subtropical conspecifics. When the results were combined across all sponge species, the chemical deterrence of fish predators was significantly lower for extracts obtained from the temperate sponge community as compared to the tropical/subtropical assemblage. These results support the more general hypothesis that a lower density and diversity of sponge predators occurs at high as compared to low latitudes in the western Atlantic and may contribute to decreased investment in chemical defenses.     

Non-neutral community dynamics: empirical predictions for ecosystem function and diversity from linearized consumer–resource interactions

A general model of linearized species interactions, essentially Lotka–Volterra theory, applied to questions of biodiversity has previously been shown to be a powerful tool for understanding local species–abundance patterns and community responses to environmental change for a single trophic level. Here this approach is extended to predict community composition and responses to environmental changes in trophically structured systems. We show how resource and consumer species richness and their relative abundances vary with the means and variances in enrichment level and strengths of intra- and interspecific interactions. Also demonstrated are the responses of local resource and consumer species richness to the global species pools at both trophic levels, as well as the covariation with net resource productivity. These predictions for resource and consumer specific responses to changes in environmental enrichment and global biodiversity are directly testable.     

Acute toxicity testing with the European estuarine amphipod <Emphasis Type="Italic">Corophium multisetosum</Emphasis>

This study reports on the use of the estuarine amphipod Corophium multisetosum in acute toxicity testing. The species was successfully acclimated to the laboratory and was used in a water-only whole effluent 96 h acute bioassay and in a 10 days whole estuarine sediment test. C. multisetosum response was compared to other species in 96 h bioassays, testing boiling cork effluent and iron filings lixiviates. The amphipod showed high sensitivity and the results were similar to those obtained with others species namely, the freshwater cladoceran Daphnia magna, the estuarine amphipod Gammarus chevreuxi and the seawater annelid Sabellaria alveolata. In a 10 days static exposure to natural impacted estuarine sediments, the response of the species was compared to a fertilization bioassay with the sea-urchin Paracentrotus lividus. Both species indicated the same sediment samples as the most potentially toxic. These samples were collected in sites where the resident macrofauna benthic community is also the most affected, with strong reduction of the species richness, abundance and biomass. The results revealed that C. multisetosum presents high potential to be used in routine acute toxicity testing in the estuarine environment.     

Effects of organic enrichment on meiofaunal abundance and community structure in sublittoral soft sediments

In experimental mesocosms established at Solbergstrand, Oslofjord, Norway, organic enrichment was effected by the addition of powdered Ascophyllum nodosum (L.) Le Jol., in quantities equivalent to 50gC·m^?2 and 200gC·m^?2, to boxes of sublittoral soft sediment. After 56 days, the structure of the meiofaunal communities in these treatments was compared with that of the control boxes. At this time the meiofaunal communites at each level of organic enrichment were markedly different from each other and from that in the control sediment. The responses of the two major components of the meiofauna, however, were different. Although the abundance of nematodes was slightly reduced in the high dose treatment this was not accompanied by detectable changes in community structure. Harpacticoid copepods, on the other hand, increased significantly in abundance in the treatment boxes and showed a general trend towards increased dominance and decreased diversity with increasing levels of organic enrichment, although in the low dose treatment there was also an increase in the number of species present. It is also shown that the nematode/copepod ratio is unreliable as a biomonitoring tool and it is suggested that the differential responses in community structure between the nematode and copepod components of the meiofauna might be a better indication of stress at the community level.     

Parasites of flounder (Platichthys flesus L.) from the German Bight, North Sea, and their potential use in biological effects monitoring

In the frame of an integrated biological effect monitoring programme, the parasite community of flounder (Platichthys flesus) was investigated at different locations in the German Bight from 1995 to 2000. In order to assess the impact of environmental contamination caused by anthropogenic activities on the parasite community, selected parasitological parameters that displayed significant differences between the sampling sites were subjected to correlation analyses with site-specific contamination and individual pollution loads of their fish hosts. In addition, correlation analyses were conducted with the responses of selected genetic, biochemical, histopathological, physiological and immunological parameters of fish, used as potential biomarkers. In total, 802 flounder were analysed for these parameters. Information on the chemical background at the sampling sites was derived from sediment samples and from 120 samples of blue mussel (Mytilus edulis) tissue, collected at each of the sampling sites. Based on chemical data available from the sediment and blue mussel samples, a pollution gradient could be established between the sampling sites for individual contaminants. The relative abundance of Acanthochondria cornuta, Cucullanus heterochrous and Zoogonoides viviparus, and the community measures species richness and number of heteroxenous species decreased with increasing concentrations of individual heavy metals or hydrocarbons in sediment and blue mussel samples. Most of the parasitological parameters significantly reflected the established site-specific contamination gradient, when data were pooled over all sampling campaigns. Significant correlations were also found with the contamination level of individual flounder. The parasitological parameters included the parasite species Lepeophtheirus pectoralis and Lernaeocera branchialis, which were not correlated to site-specific contamination. Several biomarkers were significantly correlated to the abundance of parasitic copepods A. cornuta, Lernaeocera branchialis and Lepeophtheirus pectoralis and to parasite community parameters. The results showed that the abundance of several metazoan parasite species, species richness and parasite diversity were reduced in contaminated habitats, and that differences between sites were not only related to natural factors, such as salinity, but also to pollution-induced stress. Thus, it can be concluded that the parasite community of fish responds to the level of pollution at a specific site as well as to residues of xenobiotics in individual fish. These findings give indications that the parasite community of fish is a valuable parameter for the assessment of ecological consequences of chemical contamination in aquatic habitats.     

RESPONSES OF THE PHYTOPLANKTON COMMUNITY GROWTH RATE TO NUTRIENT PULSES IN VARIABLE ESTUARINE ENVIRONMENTS

In estuaries, phytoplankton are exposed to rapidly changing conditions that may have profound effects on community structure and function. In these experiments, we evaluated the growth, productivity, and compositional responses of natural phytoplankton communities exposed to limiting nutrient additions and incubation conditions typical of estuarine habitats. Mesocosm bioassays were used to measure the short-term (2-day) growth rate, primary productivity, and group-specific biomass responses of the phytoplankton community in the Neuse River Estuary, North Carolina. A three-factor (mixing, sediment addition, and nutrient addition) experimental design was applied using 55-L mesocosm tanks. Growth rates were determined using the ¹⁴C photopigment radiolabeling method, and the abundance of algal groups was based on quantification of chemosystematic photopigments by HPLC. For Neuse River Estuary phytoplankton communities, stratified (nonmixed), turbid, and low-nitrate conditions favored increases in cryptomonad biomass. Mixed, turbid, high-nitrate conditions were favorable for increased primary productivity and chlorophytes, diatoms, and cyanobacteria. The highest community growth rates occurred under calm, high-nitrate conditions. This approach provided an assessment of the community-level phytoplankton responses and insights into the mechanisms driving blooms and bloom species in estuarine waters. The ability to rapidly alter growth rates to capitalize on conditions conducive for growth may play an important role in the timing, extent, and species involved with blooms in estuarine waters. Adaptive growth rate responses of individual species, as well as the community as a whole, further illustrate the sensitivity of estuarine ecosystems to excessive N inputs.     

Sediment Composition Influences Spatial Variation in the Abundance of Human Pathogen Indicator Bacteria within an Estuarine Environment

Faecal contamination of estuarine and coastal waters can pose a risk to human health, particularly in areas used for shellfish production or recreation. Routine microbiological water quality testing highlights areas of faecal indicator bacteria (FIB) contamination within the water column, but fails to consider the abundance of FIB in sediments, which under certain hydrodynamic conditions can become resuspended. Sediments can enhance the survival of FIB in estuarine environments, but the influence of sediment composition on the ecology and abundance of FIB is poorly understood. To determine the relationship between sediment composition (grain size and organic matter) and the abundance of pathogen indicator bacteria (PIB), sediments were collected from four transverse transects of the Conwy estuary, UK. The abundance of culturable Escherichia coli, total coliforms, enterococci, Campylobacter, Salmonella and Vibrio spp. in sediments was determined in relation to sediment grain size, organic matter content, salinity, depth and temperature. Sediments that contained higher proportions of silt and/or clay and associated organic matter content showed significant positive correlations with the abundance of PIB. Furthermore, the abundance of each bacterial group was positively correlated with the presence of all other groups enumerated. Campylobacter spp. were not isolated from estuarine sediments. Comparisons of the number of culturable E. coli, total coliforms and Vibrio spp. in sediments and the water column revealed that their abundance was 281, 433 and 58-fold greater in sediments (colony forming units (CFU)/100g) when compared with the water column (CFU/100ml), respectively. These data provide important insights into sediment compositions that promote the abundance of PIB in estuarine environments, with important implications for the modelling and prediction of public health risk based on sediment resuspension and transport.     

Modelling the responses of Australian subtropical rainforest birds to changes in environmental conditions along elevational gradients

Montane birds face significant threats from a warming climate, so determining the environmental factors that most strongly influence the composition of such assemblages is of critical conservation importance. Changes in temperature and other environmental conditions along elevational gradients are known to influence the species richness and abundance of bird assemblages occupying mountains. However, the role of species‐specific traits in mediating the responses of bird species to changing conditions remains poorly understood. We aimed to determine whether different bird species responded differently to changing environmental conditions in a relatively understudied biodiversity hotspot in subtropical rainforest on the east coast of Australia. We examined patterns in avian species richness and abundance along two rainforest elevational gradients using monthly point counts between September 2015 and October 2016. Environmental data on temperature, wetness, canopy cover and canopy height were collected simultaneously, and trait information on body size and feeding guild membership for each bird species was obtained from the Handbook of Australian, New Zealand and Antarctic Birds. We used a generalized linear mixed modelling (GLMM) framework to determine the drivers of species richness and abundance and to quantify species’ trait–environment interactions. GLMMs indicated that temperature alone was significantly positively correlated with species richness and abundance. Species richness declined with increasing elevation. When modelling abundance, we found that feeding guild membership did not significantly affect species’ responses to environmental conditions. In contrast, the predicted abundance of a species was found to depend on its body size, due to significant positive interactions between this trait, temperature and canopy cover. Our findings indicate that large‐bodied birds are likely to increase in abundance more rapidly than small‐bodied birds with continued climatic warming. These results underline the importance of temperature as a driving factor of avian community assembly along environmental gradients.     

Fish assemblage–environment relationships suggest differential trophic responses to heavy metal contamination

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Macrobenthic–mud relations strengthen the foundation for benthic index development: A case study from shallow,temperate New Zealand estuaries

Globally the input of sediment to coastal systems, in particular to estuaries, is predicted to increase due to anthropogenic activities. Sediment mud content is a powerful driver of ecologically important, macrobenthic taxa in estuarine intertidal flats. Accordingly, forecasting species responses to increased sedimentation is fundamental for effective ecosystem management, particularly for productive, geologically young, and sand-dominated estuaries that characterise many countries, including New Zealand (NZ). Modelling studies have highlighted the non-linear, highly variable responses of taxa to mud concentration. However, existing taxon-specific models have not adequately accounted for the full mud gradient, the influence of potentially confounding variables (e.g. organic enrichment, heavy metal concentrations), or regional differences in species responses. Furthermore, such models are often based on qualitative expert consensus of the membership of taxa in ecological groups that characterise their sensitivity to mud content. In this study, data from 25 unmodified to highly disturbed, shallow NZ estuaries, were used to develop an ecologically relevant model to relate the responses of 39 taxa to sediment mud content for use in the intertidal flats of shallow, temperate estuaries. Preliminary analyses indicated that sediment mud content was the dominant driver of macroinvertebrate community composition among sites, total organic carbon was of secondary importance and heavy metals did not explain significant variation in composition. Regression analysis revealed a significant linear relationship between sediment mud and total organic carbon content, which permitted subsequent analyses to be based on mud content alone. Generalised additive models were used to develop taxon-specific models that, according to k-fold cross validation, accurately predicted both probability of presence (up to 79% deviance explained) and maximum density (up to 96% deviance explained) along the sediment mud gradient (0.1–92.3%). Estimates of “optimal mud range” and “distribution mud range” were quantitatively-derived for each taxon and used to categorise taxa into one of five ecological groups (identical to those used in existing biotic indices), based on their individual sensitivities to increasing mud content. By removing expert consensus from the grouping process, the classification methods established herein provide strong support for the use of quantitative indices for the assessment and management of estuarine condition in response to increasing sediment mud content. The findings indicate that NZ estuarine sediments (2–25% mud) support a more diverse and abundant macroinvertebrate assemblage and exhibit low organic enrichment (<1% total organic carbon) compared to estuaries with >25% mud content.     

Amphibian richness patterns in Atlantic Forest areas invaded by American bullfrogs

The relationship between invasion success and native biodiversity is central to biological invasion research. New theoretical and analytical approaches have revealed that spatial scale, land‐use factors and community assemblages are important predictors of the relationship between community diversity and invasibility and the negative effects of invasive species on community diversity. In this study we assess if the abundance of Lithobates catesbeianus, the American bullfrog, negatively affects the richness of native amphibian species in Atlantic Forest waterbodies in Brazil. Although this species has been invading Atlantic Forest areas since the 1930s, studies that estimate the invasion effects upon native species diversity are lacking. We developed a model to understand the impact of environmental, spatial and species composition gradients on the relationships between bullfrogs and native species richness. We found a weak positive relationship between bullfrog abundance and species richness in invaded areas. The path model revealed that this is an indirect relationship mediated by community composition gradients. Our results indicate that bullfrogs are more abundant in certain amphibian communities, which can be species‐rich. Local factors describing habitat heterogeneity were the main predictors of amphibian species richness and composition and bullfrog abundance. Our results reinforce the important role of habitats in determining both native species diversity and potential invasibility.     

Does functional redundancy of communities provide insurance against human disturbances? An analysis using regional-scale stream invertebrate data

Human-induced reductions in species richness might alter the quality of ecosystem services when the remaining species are not able to substitute the functions provided by extirpated species. We examined how human disturbances (nutrient enrichment, land use intensification, instream habitat degradation and the presence of alien species) influence the species richness of stream invertebrates. Stream invertebrates (425 native species) were collected by kick and sweep sampling technique at 274 stream sites covering the entire area of Hungary. We measured the species richness, functional richness (i.e. number of unique functional roles provided by community members) and functional redundancy (i.e. the functional insurance of the community) using information on the feeding habits of each species. To remove the effect of natural variability, we tested the effect of stressors on the residuals of models relating species richness, functional richness and functional redundancy with natural environmental gradients. Our results showed that species richness was negatively influenced by instream habitat degradation and nutrient enrichment. Independent of the way of quantifying functional richness and functional redundancy, we found that functional richness is more sensitive to human impact than functional redundancy of stream invertebrates. The finding that a reduction of species richness is associated with a loss of unique functional roles (functional richness) is important for conservation issues, because the number of unique functional roles is usually regarded as driver of ecosystem functioning.     

Declines in littoral species richness across both spatial and temporal nutrient gradients: a palaeolimnological study of two taxonomic groups

1. Using a palaeolimnological approach in shallow lakes, we quantified the species richness responses of diatoms and Cladocera to phosphorus enrichment. We also examined differences in species richness responses between littoral and pelagic assemblages of our focal communities. To address both spatial and temporal relationships, our study includes an analysis of both surface sediments from 40 lakes and of a lake sediment record spanning c. 120 years. The objective of our study was to determine whether similar species richness patterns occurred across trophic levels, as well as along spatial and temporal gradients. 2. We found that both diatom and Cladocera species richness estimates significantly declined with increasing phosphorus across space and through time. When the assemblages were subdivided according to known habitat preferences, littoral biodiversity maintained a negative trend, whereas pelagic species richness tended to show no relationship with phosphorus. 3. Negative productivity–diversity patterns have been observed across almost all palaeolimnological studies that span large productivity gradients. This congruence in patterns is most likely due to the similarity in data collection methods and in focal communities studied. The contrasting responses between littoral and pelagic assemblages may be explained by the differences in physical habitat and the pool of taxa in each of these environments. Consistent with the literature, we found statistical support for the idea that littoral diversity declines could be explained by an interaction between macrophytes and nutrients along strong trophic gradients. The general lack of a diversity response in our pelagic assemblages could be attributable to the limited pool of subfossil taxa. The response of the pelagic diatom could also be related to their broad range of nutrient tolerances. 4. The observed negative response of species richness to phosphorus enrichment, particularly in the littoral assemblages, has implications for ecosystems functioning because communities with reduced biodiversity often are less resilient to anthropogenic change.     

The Scaling of Host Density with Richness Affects the Direction,Shape, and Detectability of Diversity-Disease Relationships

Pathogen transmission responds differently to host richness and abundance, two unique components of host diversity. However, the heated debate around whether biodiversity generally increases or decreases disease has not considered the relationships between host richness and abundance that may exist in natural systems. Here we use a multi-species model to study how the scaling of total host community abundance with species richness mediates diversity-disease relationships. For pathogens with density-dependent transmission, non-monotonic trends emerge between pathogen transmission and host richness when host community abundance saturates with richness. Further, host species identity drives high variability in pathogen transmission in depauperate communities, but this effect diminishes as host richness accumulates. Using simulation we show that high variability in low richness communities and the non-monotonic relationship observed with host community saturation may reduce the detectability of trends in empirical data. Our study emphasizes that understanding the patterns and predictability of host community composition and pathogen transmission mode will be crucial for predicting where and when specific diversity-disease relationships should occur in natural systems.     

High levels of sediment contamination have little influence on estuarine beach fish communities

While contaminants are predicted to have measurable impacts on fish assemblages, studies have rarely assessed this potential in the context of natural variability in physico-chemical conditions within and between estuaries. We investigated links between the distribution of sediment contamination (metals and PAHs), physico-chemical variables (pH, salinity, temperature, turbidity) and beach fish assemblages in estuarine environments. Fish communities were sampled using a beach seine within the inner and outer zones of six estuaries that were either heavily modified or relatively unmodified by urbanization and industrial activity. All sampling was replicated over two years with two periods sampled each year. Shannon diversity, biomass and abundance were all significantly higher in the inner zone of estuaries while fish were larger on average in the outer zone. Strong differences in community composition were also detected between the inner and outer zones. Few differences were detected between fish assemblages in heavily modified versus relatively unmodified estuaries despite high concentrations of sediment contaminants in the inner zones of modified estuaries that exceeded recognized sediment quality guidelines. Trends in species distributions, community composition, abundance, Shannon diversity, and average fish weight were strongly correlated to physico-chemical variables and showed a weaker relationship to sediment metal contamination. Sediment PAH concentrations were not significantly related to the fish assemblage. These findings suggest that variation in some physico-chemical factors (salinity, temperature, pH) or variables that co-vary with these factors (e.g., wave activity or grain size) have a much greater influence on this fish assemblage than anthropogenic stressors such as contamination.