Effects of physical disturbance on nematode communities in sand and mud: a microcosm experiment

A microcosm experiment was carried out to evaluate the effects of continuous and spasmodic physical disturbance of differing frequency on the structure of nematode communities of intertidal sand and mud. There was a marked, characteristic change in abundance and diversity for both sediment types. In the sand microcosms, the majority of univariate measures of community structure, including species diversity, were lowest in the sediments subjected to a high frequency of disturbance. For the mud microcosms, most univariate measures reached their highest values in the treatments with an intermediate frequency of disturbance and were lower in treatments subjected to both higher and lower frequencies. Multivariate ordinations for both nematode assemblages showed a clear separation of undisturbed controls and disturbed treatments, but only for the muddy sediment was there a graded change in community composition with increasing frequency of disturbance. These results confirmed our a priori expectation that nematode assemblages from mobile sandy sediments would be more resilient to physical disturbance than those from sheltered muds, and these observations are considered in the context of Connell's intermediate disturbance hypothesis. Received: 7 May 1997 / Accepted: 10 October 1997     

The impact of seabed disturbance on nematode communities: linking field and laboratory observations

Physical disturbance is a key factor in controlling the spatial and temporal composition of shallow-water benthic communities. Like shallow waters, deeper waters are increasingly subject to a range of anthropogenic disturbances, which can lead to significant alterations in sedimentation patterns. These alterations often exceed naturally occurring changes. We used a combined analysis of six independent data sets arising from large-scale field surveys and small-scale laboratory experiments to investigate the effects of seabed disturbance on nematode communities. Disturbance response was documented as a function of disturbance type (coastal development, dredged material disposal, bottom trawling, glacial fjord) and intensity (low, medium, high). Natural and man-induced seabed disturbance exerted differential effects on exposed populations, generating changes in the taxonomic (genus) and functional (feeding type) attributes of their assemblages. The genus composition of nematode assemblages from geographically separate seas converged with increased level of various types of man-made disturbance. Assemblages present along a gradient of natural disturbance in a glacial fjord followed an opposite response vector, suggesting that community changes induced by anthropogenic activities, or experimental treatments simulating the principal impacts of these, inherently differ from disturbance of natural origin. Changes in trophic diversity and structure were primarily driven by factors confounded with physical disturbance, such as metal contamination. Coupling the results of analyses at multiple scales proved a useful means of providing deeper insights into the general response of ecological communities to environmental change.     

The role of habitat heterogeneity in structuring the community of intertidal free-living marine nematodes

The role of habitat complexity has been widely neglected in the study of meiofaunal community patterns. We studied the intertidal nematode community of a structurally complex macrotidal beach exhibiting contrasting microhabitats (sandbars and runnels) to understand the influence of environmental gradients and habitat heterogeneity in the community structure. We tested whether topographical complexity affected (1) the zonation pattern in terms of abundance and diversity, and (2) local diversity by promoting compartmentalization into distinct faunal groups. Our analyses revealed three major faunal assemblages along the exposure gradient associated to differences in mean grain size and chlorophyll a. Diversity patterns involved a mid-intertidal peak, consistent with the intermediate disturbance hypothesis, and another peak at the limit with the subtidal region, consistent with the transition zone. These results highlight the predominance of environmental gradients in establishing intertidal zonation. However, microhabitats differed in environmental conditions and possessed significantly distinct nematofaunal communities. Runnels featured higher levels of taxonomic and functional diversity, many unique genera, and the community differed from the assemblage at the limit to the subtidal, stressing their role as distinct microhabitats. The nematofauna of the structurally complex beach was more diverse than the one from a homogeneous beach nearby, supporting the hypothesis that structural heterogeneity promotes diversity by compartmentalization and highlighting the importance of microhabitats in the assessment of biodiversity. Contrary to previous predictions, our results indicate potentially high regional marine nematode diversity in the Upper Gulf of California.     

Effects of physical disturbance on an estuarine intertidal community: field and mesocosm results compared

The response of an estuarine macrobenthic community to different intensities of physical disturbance was examined using comparable field and mesocosm experiments. In the field study, areas of sediment (2500 cm²) were disturbed by surface-raking at a range of disturbance intensities: no disturbance, once every 2 mo, once every month, once every 2 wk, once or three times every week. These disturbance regimes were also imposed on buckets of sediment (855 cm²) collected from the field study site and maintained within a mesocosm system. In the univariate data analysis, neither field nor mesocosm communities displayed any trends in relation to the frequency of disturbance. Diversity indices were generally reduced in the mesocosm communities compared with the field communities. Multivariate analysis indicated significant differences between the field communities subjected to the most intense disturbance and those communities disturbed every ±2 wk or less. These differences were attributed to reductions in the five numerically dominant species in the high- disturbance treatments. More statistically significant differences were observed between the different disturbance intensities in the mesocosm experiment than in the field study. The implications of differences between field and mesocosm experiments results are discussed. Received: 14 June 1999 / Accepted: 3 December 1999     

Spatio-temporal patterns in the diversity of demersal fish communities off the south coast of South Africa

The diversity of ecological communities has been the focus of many studies. Because biodiversity provides several indicators used in an Ecosystem Approach to Fisheries (EAF) to track changes in fish communities, we investigated the spatial and temporal patterns in the diversity of some demersal fish communities subjected to varying fishing pressure. Depth and catch rate were the most important predictors in explaining changes in diversity followed by longitude and survey year. Diversity, as measured by the various indices except for taxonomic distinctness (∆*), initially declined with increasing depth to about a depth of 80 m, then increased to about 150 m after which it declined. Taxonomic distinctness index (∆*) showed an increase in the taxonomic heterogeneity of the demersal community below the 300-m isobath. Diversity remained relatively constant with increase in longitude to around 24°E (which has the lowest diversity) after which it increased. The assessment of the temporal trend in diversity indicates that survey year has a significant effect on all diversity indices except for ∆*. Diversity increased and dominance declined with time. This may be result of a decline in the abundance of dominant species or an increase in the abundance less dominant species, or a combination of both effects. Multivariate analysis of the set of diversity indices showed three groups of indices: those reflecting species richness (S, Margalef’s d), those measuring mainly taxonomic relatedness (∆*), and those balancing the richness and evenness components of diversity (J′, H′, λ, ∆, Hill’s N1, and Hill’s N2). The relationship between evenness, catch rate, and size was also investigated. Size classes with highest evenness were found to have lowest catch rate and vice versa. This highlights the need to consider the size and trophic level of species when linking diversity to the functioning of ecosystems.     

An experimental study on recolonization and succession of marine macrobenthos in defaunated sediment

Hypoxia/anoxia in coastal waters is a world wide problem which often results in mass mortality and defaunation of benthos. In this study, field experiments were carried out to examine recolonization and succession of macrobenthic infauna in defaunated sediments, and the time required for recovery from complete defaunation to a stable community. Trays (33 cm length × 25.5 cm width × 11 cm depth) of defaunated sediment were exposed at the subtidal of a pristine site in subtropical Hong Kong. Temporal changes of macrobenthic communities in defaunated sediment were analyzed by univariate and multivariate statistics, and compared with those in undisturbed natural sediment at the same site. Initial colonization of macrobenthos occurred rapidly. A total of 42 species was found, with an average of 258 animals per tray and 24 species per tray recorded in the first month. Abundance showed a small peak (496 animals per tray) after 3 months, reached a sharp peak (1154 animals per tray) after 6 months, and declined thereafter. Species number increased gradually, reached a maximum (68 species per tray) after 9 months, and then decreased. Recolonization was predominantly contributed by larval settlement rather than adult migration. Temporal changes in abundance, species number and diversity of the macrobenthic community in defaunated sediment resemble the spatial changes along a decreasing pollution gradient previously defined by other authors. Results of this experiment suggest that newly available sediment may allow more species to colonize (or coexist) than sediment pre-occupied by an established community. This is probably due to less interspecific competition in the former habitat. No significant difference in abundance or species richness was observed between defaunated and natural sediments after 15 months, suggesting that a stable community had been achieved, although minor variations in species composition were still discernible between defaunated and natural sediments. Received: 24 May 1999 / Accepted: 20 October 1999     

Biogenic disturbance determines invasion success in a subtidal soft-sediment system

Theoretically, disturbance and diversity can influence the success of invasive colonists if (1) resource limitation is a prime determinant of invasion success and (2) disturbance and diversity affect the availability of required resources. However, resource limitation is not of overriding importance in all systems, as exemplified by marine soft sediments, one of Earth's most widespread habitat types. Here, we tested the disturbance-invasion hypothesis in a marine soft-sediment system by altering rates of biogenic disturbance and tracking the natural colonization of plots by invasive species. Levels of sediment disturbance were controlled by manipulating densities of burrowing spatangoid urchins, the dominant biogenic sediment mixers in the system. Colonization success by two invasive species (a gobiid fish and a semelid bivalve) was greatest in plots with sediment disturbance rates < 500 cm(3) x m(-2) x d(-1), at the low end of the experimental disturbance gradient (0 to > 9000 cm(3) x m(-2) x d(-1)). Invasive colonization declined with increasing levels of sediment disturbance, counter to the disturbance-invasion hypothesis. Increased sediment disturbance by the urchins also reduced the richness and diversity of native macrofauna (particularly small, sedentary, surface feeders), though there was no evidence of increased availability of resources with increased disturbance that would have facilitated invasive colonization: sediment food resources (chlorophyll a and organic matter content) did not increase, and space and access to overlying water were not limited (low invertebrate abundance). Thus, our study revealed the importance of biogenic disturbance in promoting invasion resistance in a marine soft-sediment community, providing further evidence of the valuable role of bioturbation in soft-sediment systems (bioturbation also affects carbon processing, nutrient recycling, oxygen dynamics, benthic community structure, and so on.). Bioturbation rates are influenced by the presence and abundance of large burrowing species (like spatangoid urchins). Therefore, mass mortalities of large bioturbators could inflate invasion risk and alter other aspects of ecosystem performance in marine soft-sediment habitats.     

Distribution and diversity of macrobenthic communities in the Jiangsu Reach of the Yangtze River北大核心CSCD

Macrobenthic communities are an important part of aquatic ecological system in the Yangtze River. Studies on macrobenthic community structure and diversity have important significance for development and protection of water resources in the Yangtze River. In this research, macrobenthic communities and their habitat in the Jiangsu Reach of the Yangtze River were investigated at 17 sites in May and September of 2013. Shannon-wiener diversity index was used to assess the diversity of macrobenthic communities. The results showed that there were 36 species at these 17 sites; the abundance and diversity of macrobenthic community were the highest in the Nanjing Reach and the Changzhou Reach, the lowest in the Nantong Reach of the Yangtze River. Nephthys sp., Grandidierella chaohuensis sp., Gammarus sp., Tubifex sp. and Limnodrilus hoffmeisteri were the dominant taxa in the Jiangsu Reach of the Yangtze River. Cluster analysis based on species abundance yielded four groups among the 17 sites. One-way analysis of similarity (ANOSIM) suggested significant differences in macrobenthic community structure among the four groups (P < 0.05). The results indicated that factors including aquatic plants, water flow velocity, sediment types and channels can influence the distribution and diversity of macrobenthic communities in the Jiangsu Reach of the Yangtze River.     

Effects of long-term physical disturbance by commercial scallop fishing on subtidal epifaunal assemblages and habitats

 This paper examines spatial differences in the distribution of by-catch assemblages from the scallop [Pecten maximus (L.) and Aequipecten opercularis (L.)] fishing grounds in the North Irish Sea, during 1995. The sites examined have been exposed to differing known levels of fishing disturbance by scallop dredging, based on unusually high-resolution data extracted from fishermens' logbooks. Uni- and multi-variate techniques have been used on a production dataset (a value which incorporates both abundance and biomass figures), as well as abundance and biomass data individually. The original species list was reduced to higher taxonomic groupings in line with the theory that the latter is more appropriate for detecting anthropogenic change. Species diversity and richness, total number of species, and total number of individuals all decrease significantly with increasing fishing effort. Species dominance increases with effort. Total abundance, biomass and production, and the production of most of the major individual taxa investigated decrease significantly with increasing effort. Multivariate analysis reveals a significant relationship between fishing effort and by-catch assemblage structure. The taxa most responsible for the differences are the echinoids and cnidarians, but prosobranch molluscs and crustaceans also contribute to the differences. By-catch assemblage structure is more closely related to fishing effort than any other environmental parameter investigated, including depth and sediment type. We observed an approximately linear decrease in diversity with increasing fishing disturbance, and suggest this is primarily due to selective removal of sensitive species and, more importantly, habitat homogenisation. These results were interpreted in the light of ecological theories relating disturbance to community structure. The argument that invertebrate scavenger populations benefit from prolonged exposure to fishing disturbance was also examined, but no supporting evidence was found. Received: 29 January 2000 / Accepted: 31 May 2000     

草原不同沙化地段蝗虫与植物群落多样性的变化及相互关系的数值分析

通过对植物和蝗虫种群生物量的调查，研究了宁夏盐池县干草原沙化过程中植物和蝗虫群落结构、多样性与丰富度的变化，以及其相互关系．结果表明，植物和蝗虫群落随着草地的沙化程度表现出序列性演替过程，蝗虫群落多样性直接受植物群密结构变化的影响，但二者的变化趋势并不完全平行．草地沙化对蝗虫群落多样性和丰富度是一种灾变性影响：沙化导致植被结构与土壤硬度的变化，造成蝗虫产卵地条件的改变．     

Limited trophodynamics effects of trawling on three Mediterranean fishes

Trawling has a significant effect on the structure of marine communities, yet the ubiquity of trawling impacts makes testing such effects difficult. This study examines trawling impacts on trophodynamics of three fishes among the Gulfs of Castellammare and Termini Imerese (northern Sicily), the first of which has been subject to a trawling ban since 1990 that initially resulted in an eightfold increase in total fish biomass. The Gulf of Termini Imerese remains heavily fished and was treated as a control site. Nitrogen and carbon stable isotope data were used to assess fishing induced changes in trophic levels or source of production supporting three demersal fish species; Mullus barbatus, Merluccius merluccius and Lophius budegassa following a control-impact approach. The exclusion of trawling resulted in only small alteration of δ¹⁵N in two of the three-three species. There were no systematic changes in the δ¹³C of any species sampled. Thus, a large influence of trawling on the trophodynamics of the studied species at sampled size was discounted. Although stable isotopes do not have spatial or temporal resolution to identify detailed shifts in diet composition, their integrative nature highlights that the trophic role these species play is robust to fishing impacts at scales over which the fishery operates. This is despite a significant increase in the abundance of these species and of total fish biomass within the protected area.

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Disturbance, Diversity, and Invasion: Implications for Conservation

Disturbance is an important component of many ecosystems, and variations in disturbance regime can affect ecosystem and community structure and functioning. The "intermediate disturbance hypothesis" suggests that species diversity should be highest at moderate levels of disturbance. However, disturbance is also known to increase the invasibility of communities. Disturbance therefore poses an important problem for conservation management, Here, we review the effects of disturbances such as fire grazing, soil disturbance and nutrient addition on plant species diversity and invasion with particular emphasis on grassland vegetation. Individual components of the disturbance regime can have marked effects on species diversity, but it is often modifications of the existing regime that have the largest influence. Similarly, disturbance can enhance invasion of natural communities, but frequently it is the interaction between different disturbances that has the largest effect. The natural disturbance regime is now unlikely to persist within conservation areas since fragmentation and human intervention have usually modified physical and biotic conditions. Active management decisions must now be made on what disturbance regime is required and this requires decisions on what species are to be encouraged or discouraged.     

Mesozooplankton community structure in offshore and coastal waters of the Ionian Sea (eastern Mediterranean) during mixed and stratified conditions

Depth-stratified samples, collected during a period where the water column was vertically mixed (March 2000) and a period of thermal stratification (September 2000), were analyzed in order to investigate the horizontal and vertical distribution patterns and composition of mesozooplankton, especially copepod species assemblages, in a pelagic (Ionian Sea) and a coastal area (Patraikos Gulf) of the eastern Mediterranean. Total mesozooplankton abundance and biomass were significantly lower in the highly oligotrophic offshore waters of the Ionian Sea when compared to the semi-enclosed Patraikos Gulf during both seasons. Small-sized copepods dominated the mesozooplankton community. An ‘offshore’ and a ‘coastal’ copepod assemblage were defined in the surface layer (0–50 m) only during March when differences in environmental conditions (i.e., temperature, salinity and fluorescence) were strong between the two areas. Copepod vertical community structure in offshore waters differed between sampling months. In March one assemblage (0–200 m) was mainly identified, while in September three distinct assemblages (0–50, 50–100 and 100–200 m) were observed, related to different vertical distribution patterns of the various copepod species. A pronounced seasonal change of the dominant copepods was evident in the surface layer, where strong differences in hydrological properties were observed from March to September. Below this layer, the copepod community was relatively stable showing decreasing seasonal differences with increasing depth.     

Temporal and spatial changes in plant dune diversity in urban resorts

We investigated the temporal and spatial changes in the floristic composition and abundance in sand dunes along a coastal strip in the province of Buenos Aires (Argentina). Firstly, we analysed changes over a span of 70 years, comparing 41 of our own inventories carried out in 14 beach resorts in 2005–2006 with 18 inventories made in the first half of the XXth century (1930–1940) in the same study area. We grouped sampled plots into four categories for comparison, taking into account the time period (1930–1940 or 2005–2006) and location at each end of the surveyed coastal strip in the northern or southern ‘tourist’ sectors. Secondly, we analysed the effect of afforestation with exotic trees on extant sand dune vegetation for the period 2005–2006. A total of 74 plant inventories were carried out in afforested and natural dunes at the same resorts. In both comparisons we contrasted plant richness, percentage plant cover, similarity and diversity. A cluster analysis was used to classify categories taking into account plant abundance and composition. Significant differences in total diversity were observed between sampling years, suggesting an increase in diversity at the present time due to urbanization and recreation related activities. Nevertheless, no single exotic plant species was clearly dominant across the sampled sites, suggesting that these coastal areas are, up to now, resistant to alien invasion. Diversity indexes discriminated by plant groups indicating different habitat conditions and exotics were shown to be more sensitive to existing changes than to total richness and cover.     

Effects of biological disturbance by<Emphasis Type="Italic"> Monoporeia affinis</Emphasis> (Amphipoda) on small-scale migration of marine nematodes in low-energy soft sediments

Free-living marine, benthic nematodes quickly colonise sediments where physical forces are strong enough to suspend them into the water column. In the absence of such forces colonisation is much slower and is more likely to be affected by biological factors. The aim of the study was to investigate if nematodes disperse more readily in the presence of biological disturbance where physical disturbance is rare or non-existent. Amphipods are able to greatly rework sediments, and thereby induce disturbance to the infauna. A laboratory experiment with the amphipod Monoporeia affinis and nematodes from a low-energy, 30-m-deep location was conducted in mesocosms where the nematodes were given the choice to colonise azoic sediment at three amphipod densities, zero, low and high. Each area of azoic sediment in the mesocosms was divided into three equilateral sections from the nematode source, i.e. 10, 23 and 36 cm. At termination, after 7 weeks, there were no significant differences in nematode abundance and assemblage structure between treatments despite considerable biological disturbance created by the amphipods. The number of nematodes was 16%, 15% and 11% of the total numbers in the source at the three sections 10, 23 and 36 cm, respectively. There were distinct differences in the nematode community composition between distances, with the small surface-dwelling taxon Leptolaimus spp. being a rapid and the numerically dominant coloniser of the azoic sediments. Migration of nematodes over short distances is likely to be slow in the absence of strong physical forces. To our knowledge, this is the first paper ever that investigates the influence of macrofauna on nematode short-range migration.Communicated by L. Hagerman, Helsingør     

Type of disturbance and ecological history determine structural stability.

This study aims to reveal whether complexity, namely, community and trophic structure, of chronically stressed soil systems is at increased risk or remains stable when confronted with a subsequent disturbance. Therefore, we focused on a grassland with a history of four centuries of patchy contamination. Nematodes were used as model organisms because they are an abundant and trophically diverse group and representative of the soil food web and ecosystem complexity. In a field survey, a relationship between contaminants and community structures was established. Following, two groups of soil mesocosms from the field that differed in contamination level were exposed to different disturbance regimes, namely, to the contaminant zinc and a heat shock. The zinc treatment revealed that community structure is stable, irrespective of soil contamination levels. This implies that centuries of exposure to contamination led to adaptation of the soil nematode community irrespective of the patchy distribution of contaminants. In contrast, the heat shock had adverse effects on species richness in the highly contaminated soils only. The total nematode biomass was lower in the highly contaminated field samples; however, the biomass was not affected by zinc and heat treatments of the mesocosms. This means that density compensation occurred rapidly, i.e., tolerant species quickly replaced sensitive species. Our results support the hypothesis that the history of contamination and the type of disturbance determine the response of communities. Despite that ecosystems may be exposed for centuries to contamination and communities show adaptation, biodiversity in highly contaminated sites is at increased risk when exposed to a different disturbance regime. We discuss how the loss of higher trophic levels from the entire system, such as represented by carnivorous nematodes after the heat shock, accompanied by local biodiversity loss at highly contaminated sites, may result in detrimental effects on ecosystem functions.     

Effect of the presence of the shore crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, on burrowing behaviour and clearance rate of the common cockle, <Emphasis Type="Italic">Cerastoderma edule</Emphasis>

Bivalves demonstrate various morphological and behavioural adaptations to reduce the risk of being attacked by predators. This paper examines how the presence of the crab Carcinus maenas (L.), a natural predator of the cockle Cerastoderma edule (L.), affects its burrowing depth and clearance or feeding rate. Cockles were placed in experimental tanks and treated with three levels of predatory disturbance: (1) unfed crab loose inside the tank, (2) unfed crab inside a cage suspended in the water column and (3) no crab present. Cockles’ burrowing depth was measured in two sediment types: mud and sand. Cockles burrowed more deeply in treatments with no crabs. Burrowing depth in sand was significantly greater than in mud. Two factors may contribute to the reduction in burial depth of C. edule in the presence of C. maenas: the change in the vertical orientation of the cockle and the ‘cough response’. No significant difference was found in the cockles’ clearance rate among the different levels of predator threat.     

Ecological effects of cultivation on the machair sand dune systems of the Outer Hebrides, Scotland

The machair sand dune systems of the Outer Hebrides of Scotland are a unique habitat, which is rare within both a global and European context. Unusually, the machair habitat also represents an agricultural resource that is very important to the Hebridean people, having been subject to both grazing and cultivation throughout the historical period. Following designation as an Environmentally Sensitive Area (ESA) in 1988, the machairs of South Uist have been studied with the aim of understanding the links between agricultural practice and their plant community and ecosystem dynamics. This research focused primarily on the effects of cultivation practices and their role in maintaining plant species richness and community and ecosystem stability. Within two carefully selected areas, the successional plant communities of machair at different stages of recovery following ploughing and cultivation of cereals and potato patches or ‘lazy beds’ were identified at both a macro- and micro-level. Investigations of the vegetation recovery processes on turves taken from newly ploughed land indicated that the initial stages of recolonization are characterized primarily by rapid vegetative reproduction and growth, although re-vegetation by seeds is also an important factor. The implications of these findings for the long-term management of machair plant communities are discussed and in particular the need to maintain old cultivation practices such as shallow ploughing. The need for more detailed research into both seed banks and seed rain and into processes of vegetative reproduction is stressed.     

Short-term effects of an oil spill on the West coast of the Cape Peninsula, South Africa

In 1994, the sinking of the ‘Apollo Sea’ off the West coast of South Africa led to the deposition of ca. 2 500 tons of heavy fuel oil over 150 km of coastline. The impact of the spill on rocky shore invertebrates, rock-pool fish fauna and rock lobsters was assessed by conducting surveys shortly after the spill, and again two months later. Where possible, results of these surveys were compared with existing data from before the sinking of the Apollo Sea. Among the fish fauna of rock-pools, changes in total density of fishes were largely due to changes in the abundance ofClinus superciliosus and were within the range of natural variation for the species and the community as a whole. Community structure of the rock-pool fish fauna also remained unaffected. At three of four impacted rocky shore sites no changes could be detected in overall benthic community structure, although the winkleNodilittorina africana was affected. At the fourth site, a boulder beach, statistical analysis showed distinct differences in community structure between heavily and lightly contaminated areas, as well as between all areas compared with previously existing data. There were also significant changes in the lightly oiled areas between the first and second surveys after the spill. Oil-fouled lobster were found at one of three sites investigated. Ca. 7 % of the seabed in this particular area was polluted. Antennae and forelegs of almost all lobsters in the vicinity of the oil were fouled. Examination of the gut-contents confirmed that oil inhibits the ability of lobster to feed. Although there was no evidence that mortalities of lobster were caused by the spill, growth rates may be reduced by the decreased feeding rate. Overall, the impact of the spill was strikingly less than might have been predicted from the effects of other oil spills. Nomenclature: Names of fish species follow Smith & Heemstra (1986).