Trophic relationships in the Changjiang River estuarine salt marshes: preliminary investigation from δ13C and δ15N analysis

To obtain information on food web structure in salt marshes of the Changjiang (Yangtze) River Estuary, the δ13C and δ15N values of primary producers and consumers were determined. The mean δ13C values of 31 dominant consumers ranged from -23.13‰ to -14.37‰. Except for several species (Eriocheir sinensis, Sinonovacula constricta and Potamocorbula ustulata), consumers had intermediate δ13C values between those of benthic microalgae and Spartina alterniflora. The mean δ15N values of 31 dominant consumers varie...     

Trophic relationships along a bathymetric gradient at the deep-sea observatory HAUSGARTEN

Deep-seafloor communities, especially those from the ice-covered Arctic, are subject to severe food limitation as the amount of particulate organic matter (POM) from the surface is attenuated with increasing depth. Here, we use naturally occurring stable isotope tracers (δ¹⁵N) to broaden our rudimentary knowledge of food web structure and the response of benthic organisms to decreasing food supplies along the bathymetric transect (～1300–5600 m water depth) of the deep-sea observatory HAUSGARTEN. Encompassing five trophic levels, the HAUSGARTEN food web is among the longest indicating continuous recycling of organic material typical of food-limited deep-sea ecosystems. The δ¹⁵N signatures ranged from 3.0‰ for Foraminifera to 21.4‰ (±0.4) for starfish (Poraniomorpha tumida). The majority of organisms occupied the second and third trophic level. Demersal fish fed at the third trophic level, consistent with results from stomach contents analysis. There were significant differences in the δ¹⁵N signatures of different functional groups with highest δ¹⁵N values in predators/scavengers (13.2±0.2‰) followed by suspension feeders (11.2±0.2‰) and deposit feeders (10.2±0.3‰). Depth (=increasing food limitation) affected functional groups in different ways. While the isotopic signatures of predators/scavengers did not change, those of suspension feeders increased with depth, and the reverse was found for deposit feeders. In contrast to the results of other studies, the δ¹⁵N signatures in POM samples obtained below 800 m did not vary significantly with depth indicating that changes in δ¹⁵N values are unlikely to be responsible for the depth-related δ¹⁵N signature changes observed for benthic consumers. However, the δ¹⁵N signatures of sediments decreased with increasing depth, which also explains the decrease found for deposit feeders. Suspension feeders may rely increasingly on particles trickling down the HAUSGARTEN slope and carrying higher δ¹⁵N signatures than the decreasing POM supplies, which elevates the δ¹⁵N value of their tissues. Our results imply that a depth-stratified approach should be taken to avoid a misinterpretation of data obtained at different depths.     

Trophic structure and functioning in a eutrophic and poorly flushed lagoon in southwestern Taiwan

Tapong Bay, a eutrophic and poorly flushed tropical lagoon, supports intensive oyster culture. Using the Ecopath approach and network analysis, a mass-balanced trophic model was constructed to analyze the structure and matter flows within the food web. The lagoon model is comprised of 18 compartments with the highest trophic level of 3.2 for piscivorous fish. The high pedigree index (0.82) reveals the model to be of high quality. The most-prominent living compartment in terms of matter flow and biomass in the lagoon is cultured oysters and bivalves, respectively. The mixed trophic impacts indicate that phytoplankton and periphyton are the most-influential living compartments in the lagoon. Comparative analyses with the eutrophic and well-flushed Chiku Lagoon and non-eutrophic tropical lagoons show that high nutrient loadings might stimulate the growth and accumulation of phytoplankton and periphyton and therefore support high fishery yields. However, net primary production, total biomass, fishery yields per unit area, and mean transfer efficiency of Tapong Bay were remarkably lower than those of Chiku Lagoon. The lower transfer efficiency likely results from the low mortality of cultured oysters and invasive bivalves from predation or the lower density of benthic feeders constrained by the hypoxic bottom water as a result of poor flushing. This might therefore result in a great proportion of flows to detritus. However, the hypoxic bottom water might further reduce the recycling of the entering detritus back into the food web. In contrast to many estuaries and tropical lagoons, poor flushing of this eutrophic tropical lagoon might induce a shift from detritivory to herbivory in the food web.     

Towards end-to-end models for investigating the effects of climate and fishing in marine ecosystems

End-to-end models that represent ecosystem components from primary producers to top predators, linked through trophic interactions and affected by the abiotic environment, are expected to provide valuable tools for assessing the effects of climate change and fishing on ecosystem dynamics. Here, we review the main process-based approaches used for marine ecosystem modelling, focusing on the extent of the food web modelled, the forcing factors considered, the trophic processes represented, as well as the potential use and further development of the models. We consider models of a subset of the food web, models which represent the first attempts to couple low and high trophic levels, integrated models of the whole ecosystem, and size spectrum models. Comparisons within and among these groups of models highlight the preferential use of functional groups at low trophic levels and species at higher trophic levels and the different ways in which the models account for abiotic processes. The model comparisons also highlight the importance of choosing an appropriate spatial dimension for representing organism dynamics. Many of the reviewed models could be extended by adding components and by ensuring that the full life cycles of species components are represented, but end-to-end models should provide full coverage of ecosystem components, the integration of physical and biological processes at different scales and two-way interactions between ecosystem components. We suggest that this is best achieved by coupling models, but there are very few existing cases where the coupling supports true two-way interaction. The advantages of coupling models are that the extent of discretization and representation can be targeted to the part of the food web being considered, making their development time- and cost-effective. Processes such as predation can be coupled to allow the propagation of forcing factors effects up and down the food web. However, there needs to be a stronger focus on enabling two-way interaction, carefully selecting the key functional groups and species, reconciling different time and space scales and the methods of converting between energy, nutrients and mass.     

Abrupt transitions of the top-down controlled Black Sea pelagic ecosystem during 1960–2000: Evidence for regime-shifts under strong fishery exploitation and nutrient enrichment modulated by climate-induced variations

Functioning of the Black Sea ecosystem has profoundly changed since the early 1970s under cumulative effects of excessive nutrient enrichment, strong cooling/warming, over-exploitation of pelagic fish stocks, and population outbreak of gelatinous carnivores. Applying a set of criteria to the long-term (1960–2000) ecological time-series data, the present study demonstrates that the Black Sea ecosystem was reorganised during this transition phase in different forms of top-down controlled food web structure through successive regime-shifts of distinct ecological properties. The Secchi disc depth, oxic–anoxic interface zone, dissolved oxygen and hydrogen sulphide concentrations also exhibit abrupt transition between their alternate regimes, and indicate tight coupling between the lower trophic food web structure and the biogeochemical pump in terms of regime-shift events.The first shift, in 1973–1974, marks a switch from large predatory fish to small planktivore fish-controlled system, which persisted until 1989 in the form of increasing small pelagic and phytoplankton biomass and decreasing zooplankton biomass. The increase in phytoplankton biomass is further supported by a bottom-up contribution due to the cumulative response to high anthropogenic nutrient load and the concurrent shift of the physical system to the “cold climate regime” following its ∼20-year persistence in the “warm climate regime”. The end of the 1980s signifies the depletion of small planktivores and the transition to a gelatinous carnivore-controlled system. By the end of the 1990s, small planktivore populations take over control of the system again. Concomitantly, their top-down pressure when combined with diminishing anthropogenic nutrient load and more limited nutrient supply into the surface waters due to stabilizing effects of relatively warm winter conditions switched the “high production” regime of phytoplankton to its background “low production” regime.The Black Sea regime-shifts appear to be sporadic events forced by strong transient decadal perturbations, and therefore differ from the multi-decadal scale cyclical events observed in pelagic ocean ecosystems under low-frequency climatic forcing. The Black Sea observations illustrate that eutrophication and extreme fishery exploitation can indeed induce hysteresis in large marine ecosystems, when they can exert sufficiently strong forcing onto the system. They further illustrate the link between the disruption of the top predators, proliferation of new predator stocks, and regime-shift events. Examples of these features have been reported for some aquatic ecosystems, but are extremely limited for large marine ecosystems.     

海洋浮游微食物网结构及其影响因素

微食物环是海洋生态系统中重要的物质和能量过程,是传统食物链的有效补充。微食物环研究是当前海洋生态学研究的热点之一,但对其结构的系统研究较少,海洋微食物网结构在2000年才被Garrison提出。尽管微食物网各个类群的丰度在不同海洋环境中有相对变化,但是这些变化都处于一定的范围之内,其丰度结构约为纤毛虫10 cell ml-1、鞭毛虫103 cell ml-1、微微型真核浮游生物104 cell ml-1、蓝细菌104-5 cell ml-1、异养细菌106 cell ml-1、病毒107 particle ml-1。海洋浮游食物链中捕食者和饵料生物粒径的最佳比值为10:1,实际研究中该比值会略低,例如纤毛虫与其饵料的粒径比值为8:1,鞭毛虫为3:1。Pico和Nano浮游植物的丰度比（Pico:Nano）是研究微食物网结构的指数之一,该指数具有不受研究尺度影响的优点,可用于研究区域性和全球性微食物网结构。近年来,学者们从多角度对海洋微食物网的结构开展了研究,不同海区微食物网各类群丰度、生物量的时间和空间变化研究有很多报道,微食物网的结构可受空间、季节、摄食、营养盐等多种因素影响。在对不同空间微食物网的研究中,学者往往研究不同物理性质的水团中各类群生物丰度的不同,以此来表征微食物网结构的不同;同一海区微食物网结构的季节变化也是使用各个类群丰度和生物量的变化来表示,该变化主要受水文环境因素影响。摄食者对微食物网各类生物的影响通过三种途径：1. 中型浮游动物摄食;2. 中型浮游动物摄食微型浮游动物,通过营养级级联效应影响低营养级生物;3. 中型浮游动物通过释放溶解有机物、营养盐影响细菌和低营养级生物。浮游植物通过产生化感物质和溶解有机物影响微食物网结构,而营养盐的浓度及变化则可以对微食物网产生直接或间接影响。     

Trophic functioning of the St. Lucia estuarine lake during a drought phase assessed using stable isotopes

The St. Lucia Estuary is Africa’s largest estuarine system and is currently experiencing the stress of prolonged freshwater deprivation, manifested by extremely low water levels and hypersalinity. These unprecedented conditions have raised questions regarding the trophic functioning of the ecosystem. Despite the substantial amount of research previously undertaken within this system, no studies of food web structure and function have yet been documented. This study therefore aimed to examine the food web structure of the St. Lucia estuary system through the use of carbon and nitrogen stable isotope analysis. Analysis of carbon isotope ratios indicates that benthic carbon sources are most utilised at sites with low water levels and generally higher salinity (Catalina Bay, Charter’s Creek). Conversely, the estuarine region of the mouth and Narrows, with its elevated water levels and lower salinity, still sustains a viable pelagic food web. Analysis of δ¹⁵N ratios indicates that the number of trophic transfers (food chain length) might be related to water levels. Overall, the study provides a greater understanding of the ecological processes of this complex estuarine lake, which may allow for future comparisons of trophic functioning under drought and normal/wet conditions to be made.     

2020年冬季大亚湾西南海域主要渔业生物碳氮稳定同位素研究

为了解大亚湾西南海域食物网的营养结构特征, 本研究于2020年1月份使用底拖网采集了该海域的渔业生物, 并分析了35种主要渔业生物的碳氮稳定同位素值。根据δ¹³C和δ¹⁵N值, 计算出该海域食物网6种营养结构的生态指标和主要渔业生物的营养级, 并绘制了连续营养谱。本次调查渔业生物主要为鱼类和虾蟹类, 鱼类的δ¹³C和δ¹⁵N值范围分别为-17.63‰ ~ -14.85‰和12.92‰~15.46‰, 平均值分别为-16.47‰和13.80‰; 虾蟹类的δ¹³C和δ¹⁵N值范围分别为-17.67‰ ~ -15.51‰和11.05‰~12.62‰, 平均值分别为-16.30‰和11.85‰。根据δ¹⁵N值, 用相加模型(trophic position by the additive model, TPA)和缩比模型(trophic position by the scaled model, TPS)分别计算了主要渔业生物的营养级, 结果显示两个模型计算的结果无显著性差异(P>0.1), 呈现鱼类平均营养级>虾蟹类的趋势。本研究发现大亚湾西南海域食物网初始食物来源较为单一, 存在食物链营养层级较少和长度不足, 食物网营养级多样性较低和营养结构冗余程度高的现象。与30多年前相比, 大亚湾近年高营养级生物量减少, 食物网结构由复杂趋向简单化, 生态系统稳定性较差。本研究结果不仅为了解大亚湾食物网结构组成提供了基础资料, 也为保护大亚湾渔业资源, 维持生态系统结构的稳定性提供参考依据。     

Pathways of organic matter through food webs of diverse habitats in the regulated Nakdong River estuary (Korea)

The benthic macroinvertebrates of the Nakdong River estuary were sampled at three different habitats: two salt marsh (Scirpus triqueter and Phragmites australis) beds and a bare intertidal flat. Fishes were sampled in the main channel. The trophic importance of marsh vascular plants, microphytobenthos, and riverine and channel particulate organic matter to macroinvertebrate and fish production was studied using stable carbon and nitrogen isotope tracers. There was a dramatic change in coverage of macrophytes (salt marshes and seagrass) after the construction of an estuarine barrage in 1987 in the Nakdong River estuary, with the S. triqueter bed increasing, the P. australis bed decreasing, and Zostera marina habitats being nearly lost. Although the invertebrate δ¹³C were within a narrower range than those of the primary producers, the values varied considerably among consumers in these habitats. However, the isotope signatures of consumers showed similarities among different habitats. Cluster analysis based on their isotopic similarity suggested that the isotope variability among species was related more to functional feeding groups than to habitats or taxonomic groups. While δ¹³C values of suspension feeders were close to that of the channel POM (mainly phytoplankton), other benthic feeders and predators had δ¹³C similar to that of microphytobenthos. Isotopic mixing model estimates suggest that algal sources, including microphytobenthos and phytoplankton, play an important role in supporting the benthic food web. Despite the huge productivity of emergent salt marshes, the contribution of the marsh-derived organic matter to the estuarine food webs appears to be limited to some nutrition for some invertebrates just within marsh habitats, with little on the bare intertidal flats or in the channel fish communities. Isotope signatures of the channel fishes also confirm that algal sources are important in supporting fish nutrition. Our findings suggest that benthic and pelagic microalgae made a large contribution to consumer diets, while marsh plants may not have a large role in supporting food webs in this estuarine system.     

应用稳定同位素技术研究江苏近海食物网营养结构的季节性变化

本研究基于稳定同位素技术对江苏近海春、夏、秋季采集的不同生物类群(鱼类、甲壳类、头足类和底栖动物等)进行稳定同位素的分析,利用SIBER模型计算不同季节消费者的营养生态位,并计算出6种营养结构的量化指标.结果表明,3个季节的δ13C值、δ15N值均差异不显著(p>0.05).春季δ13C值范围为?24.47‰～?14....     

Environmental controls on food web regimes: A fluvial perspective

Because food web regimes control the biomass of primary producers (e.g., plants or algae), intermediate consumers (e.g., invertebrates), and large top predators (tuna, killer whales), they are of societal as well as academic interest. Some controls over food web regimes may be internal, but many are mediated by conditions or fluxes over large spatial scales. To understand locally observed changes in food webs, we must learn more about how environmental gradients and boundaries affect the fluxes of energy, materials, or organisms through landscapes or seascapes that influence local species interactions. Marine biologists and oceanographers have overcome formidable challenges of fieldwork on the high seas to make remarkable progress towards this goal. In river drainage networks, we have opportunities to address similar questions at smaller spatial scales, in ecosystems with clear physical structure and organization. Despite these advantages, we still have much to learn about linkages between fluxes from watershed landscapes and local food webs in river networks. Longitudinal (downstream) gradients in productivity, disturbance regimes, and habitat structure exert strong effects on the organisms and energy sources of river food webs, but their effects on species interactions are just beginning to be explored. In fluid ecosystems with less obvious physical structure, like the open ocean, discerning features that control the movement of organisms and affect food web dynamics is even more challenging. In both habitats, new sensing, tracing and mapping technologies have revealed how landscape or seascape features (e.g., watershed divides, ocean fronts or circulation cells) channel, contain or concentrate organisms, energy and materials. Field experiments and direct in situ observations of basic natural history, however, remain as vital as ever in interpreting the responses of biota to these features. We need field data that quantify the many spatial and temporal scales of functional relationships that link environments, fluxes and food web interactions to understand how they will respond to intensifying anthropogenic forcing over the coming decades.     

Functional changes due to invasive species: Food web shifts at shallow Posidonia oceanica seagrass beds colonized by the alien macroalga Caulerpa racemosa

Multiple stable isotope analyses were used to examine the trophic shifts at faunal assemblages within the invading macroalga Caulerpa racemosa in comparison to established communities of Posidonia oceanica seagrass meadows. Sampling of macrobenthic invertebrates and their potential food sources of algal mats and seagrass meadows in Mallorca (NW Mediterranean) showed differences in species composition of faunal and primary producers among seagrass and C. racemosa. Accordingly, changes in food web structure and trophic guilds were observed, not only at species level but also at community level. The carbon and nitrogen isotope signatures of herbivores, detritivores and deposit feeders confirmed that the seagrass provided a small contribution to the macrofaunal organisms. δ¹³C at the P. oceanica seagrass and at the C. racemosa assemblages differed, ranging from −6.19 to −21.20‰ and −2.67 to −31.41‰, respectively. δ¹⁵N at the Caulerpa mats was lower (ranging from 2.64 to 10.45‰) than that at the seagrass meadows (3.51–12. 94‰). Significant differences in isotopic signatures and trophic level among trophic guilds at P. oceanica and C. racemosa were found. N fractionation at trophic guild level considerable differed between seagrass and macroalgae mats, especially for detritivores, deposit feeders, and herbivores. Filter feeders slightly differed with a relatively lower N signal at the seagrass and CR values at community level and at trophic guild level were higher in the C. racemosa invaded habitats indicating an increase in diversity of basal resource pools. C. racemosa did seem to broaden the niche diversity of the P. oceanica meadows it colonised at the base of the food web, may be due to the establishment of a new basal resource. The extent of the effects of invasive species on ecosystem functioning is a fundamental issue in conservation ecology. The observed changes in invertebrate and macrophytic composition, stable isotope signatures of concomitant species and consequent trophic guild and niche breadth shifts at invaded Caulerpa beds increase our understanding of the seagrass systems.     

Food web structure of the coastal area adjacent to the Tagus estuary revealed by stable isotope analysis

The identification of energy sources, pathways and trophic linkages among organisms is crucial for the understanding of food web dynamics. Stable isotopes were used to identify the trophic level of food web components and track the incorporation of organic matter of different origins in the coastal ecosystem adjacent to the Tagus estuary. It was shown that the river Tagus is a major source of organic carbon to this system. Also, the wide difference in δ¹³C among the primary consumers allowed the identification of the pelagic and the benthic energy pathways. The maximum trophic level observed was 2.4 for Sepia officinalis. This value is indicative of a short food web. It was concluded that the diet of the upper trophic level species relies directly on the lower food web levels to a considerable extent, instead of relying mostly on intermediate trophic level species. Moreover, the δ¹⁵N values of primary consumers were very close to that of particulate organic matter, probably due to poorly known processes occurring at the basis of the food web. This lowers the trophic length of the whole food web. Reliance on benthic affinity prey was high for all upper trophic level secondary consumers.     

Temporal variability in the isotopic niches of rocky shore grazers and suspension‐feeders

Carbon and nitrogen stable isotope ratios were measured in rocky inter‐tidal suspension‐feeders (brown mussels and cape reef worms) and grazers (goat's eye limpets and cape sea urchins) to determine the influences of lifestyle and time on the diets of consumers. Niche partitioning between consumer species within the same feeding guild was assessed using isotopic niche area (a proxy for trophic niche). Specimens were collected monthly at a single site in Southeastern South Africa from July 2010 to June 2011. Temporal variations in isotopic signatures were generally greater in the suspension‐feeders compared with the grazers, isotopic niche widths were smaller in the suspension‐feeders and intra‐population variations in isotope signatures were larger in the grazers. No inter‐specific niche overlap (according to standard ellipse areas) was observed within either feeding guild unless standardization calculations were used. Temporal variations in the diets of all the consumers appeared uncoupled from temporal variability in the isotopic signatures of basal resources in the region; as such, shifts in the consumer diets most likely pertained to feeding behaviour and food preferences. Our data provide new insights into how syntopic rocky shore consumers coexist by partitioning their temporally variable food environment.     

Trophic ecology of bottom fishes assemblage along coastal areas of Thailand

Food preferences, feeding attributes, trophic guilds, and ontogenetic changes in diet compositions of 45 fish species collected along coastal waters in the southern areas of the South China Sea, Thailand were examined. Most species had high food intake and fed on specific ranges of food types. Shrimp was the most important food (31.7%), followed by calanoid copepod (16.8%), fish (12.7%) and gammarid amphipod (8.3%). These fishes can be categorized into six different trophic guilds and further divided into four categories namely; piscivore, zooplanktivore, zoobenthivore and miscellaneous/opportunist. Numbers of feeding guilds at each depth vary between four and five. Four of these are consistently found throughout the year at all zones; shrimp predator, piscivore, calanoid copepod feeder and a combination of polychaete and other food feeders. Ontogenetic studies indicate that fishes are more likely to group according to species rather than size. This scientific information is important when examining the complex association between fishes and identifying groups of species using similar resources.     

Spatial analysis of the trophic interactions between two juvenile fish species and their preys along a coastal–estuarine gradient

Coastal and estuarine systems provide nursery grounds for many marine fish species. Their productivity has been correlated with terrigeneous inputs entering the coastal–estuarine benthic food web, thereby favouring the establishment of fish juveniles. Studies in these ecosystems often describe the nursery as a single large habitat without verifying nor considering the presence of contiguous habitats. Our study aimed at identifying different habitats based on macrozoobenthic communities and morpho-sedimentary characteristics and assessing the trophic interactions between fish juveniles and their benthic preys within these habitats. It included 43 sampling sites covering 5 habitats in which we described taxonomically and quantitatively the invertebrates and fish communities with stable isotopes and gut contents. It suggested that the benthic common sole Solea solea displayed feeding plasticity at the population level, separating the juveniles (G0) from the older fish (G1) into different “feeding sub-populations”. Size-based feeding plasticity was also observable in the spatial occupancy of that species in the studied bay. The demersal pouting, Trisopterus luscus, equally used the different habitats but displayed low feeding plasticity across and inside each habitat. Stable isotopes proved to be powerful tools to study the spatial distribution of trophic interactions in complex ecosystems like the bay of Vilaine and to define optimal habitats for fish that use the coastal–estuarine ecosystem as nursery grounds.     

Food web structure in a near-pristine mangrove area of the Australian Wet Tropics

Carbon and nitrogen isotopic composition was used to identify the main sources of carbon and describe the main trophic pathways in Deluge Inlet, a near-pristine mangrove estuary in tropical north Queensland, Australia. Producers' δ¹³C varied from −28.9‰ for mangroves to −18.6‰ for seagrass. Animals were also well separated in δ¹³C (−25.4‰ to −16.3‰ for invertebrates and −25.2‰ to −17.2‰ for fish), suggesting considerable differences in ultimate sources of carbon, from a substantial reliance on mangrove carbon to an almost exclusive reliance on seagrass. In general, invertebrates had lower δ¹⁵N than fish, indicating lower trophic levels. Among fish, δ¹⁵N values reflected well the assumed trophic levels, as species from lower trophic levels had lower δ¹⁵N than species from higher trophic levels. Trophic levels and trophic length were estimated based on δ¹⁵N of invertebrate primary consumers (6.1‰), with results suggesting a food web with four trophic levels. There was also evidence of a high level of diet overlap between fish species, as indicated by similarities in δ¹³C for fish species of higher trophic levels. Stable isotope data was also useful to construct a general model for this food web, where five main trophic pathways were identified: one based on both mangrove and microphytobenthos, one on plankton, two on both microphytobenthos and seagrass, and one based mainly on seagrass. This model again suggested the presence of four trophic levels, in agreement with the value calculated based on the difference in δ¹⁵N between invertebrate primary consumers and top piscivores.     

Study of the fauna inhabiting the holdfasts of Laminaria hyperborea (gunn.) fosl. along some environmental and geographical gradients

A survey of the holdfast fauna of Laminaria hyperborea was carried out at thirty-five sites around the UK, mostly along the North Sea and south coasts.Data from different age classes of holdfast from a polluted and unpolluted site indicate that volume of holdfast space is a most important measurement of sample size, a minimum size of sample being one litre and, preferably, three.Gradients of increased heavy metals were defined down the North Sea and west coasts and, by utilising measurements of kelp penetration, water clarity gradients were determined as increasing northwards on the west coast and both north and southwards on the North Sea coast from a point near Tyne and Wear county. Analysis of holdfast fauna showed numerous gradients of total numbers, species and diversity which correlate closely with the heavy metal and water clarity gradients, or with undetermined parameters analogous to latitude and longitude. Where there were increases of heavy metals or decreases of water clarity, various reciprocal relationships between suspension feeders and all other trophic groups were commonly observed. In the North Sea numbers of suspension feeders increased while numbers of all other trophic groups declined. On the west coast a heavy metal increase correlated with a decreasing species richness of all trophic categories except suspension feeders, while with total numbers only suspension feeders showed an increase. On the south coast transect neither gradient existed, and only here did all trophic categories covary, except at one very polluted site where the same reciprocal relationship was again indicated.In terms of species similarity the North Sea showed a greater pool of common species than occurred between sites of other regions and between North Sea sites with other regions, despite the widest range of heavy metal levels and wate clarity that existed there. Between-site similarity appeared to be much more dependent on geographical proximity than on the two measured environmental factors. It is suggested that effects of clarity-reduction can be measured by trophic analysis more successfully than by species presence or absence, and that changes of holdfast fauna along the North Sea coast are due largely to pollution.