Relationships among development, ecology, and morphology in the evolution of Echinoderm larvae and life cycles

The evolution of life cycles involves transitions between discrete states in one or more of the characters that comprise a developmental pattern. In this paper, we examine three of the major life cycle characters and the states for these characters. Using examples from echinoderms, we discuss the evolutionary transitions that have occurred in the type of morphogenesis, developmental habitat, and mode of nutrition during development. We evaluate the functional requirements associated with these transitions to infer the likelihood (frequency or rapidity) of change in a given character and of biases in the polarity of character state transitions. Using comparisons of closely related species, we evaluate the change between states in one character for dependence on the state of, or correlated changes in, other characters. Based on our analysis of congeneric species that differ in developmental habitat, we conclude that the transition between pelagic and benthic development is an ecological change that is independent of changes in morphogenesis and should be reversible. In contrast, the transition from feeding to nonfeeding development has been considered to be irreversible because it involves marked changes in larval morphology. We re-examine the transition between different modes of larval nutrition in light of recent studies that show that there exists a continuum of nutritional strategies between planktotrophy and lecithotrophy. This continuum is largely determined by variation in maternal investment and does not involve alterations in larval morphology. We suggest that the boundary between planktotrophy and lecithotrophy is frequently crossed and that this transition is reversible. Ecological changes represent the crossing of a functional threshold. Only after crossing the threshold, do larvae experience qualitatively different selective pressures that can lead to subsequent changes in morphology and development. Two different changes have occurred in the type of morphogenesis: the simplification of larval morphology that is associated with obligate (nonfeeding) lecithotrophy and the loss of the larval body plan in the evolution from indirect to direct development. It is the modification of morphology independent of the ecological changes that requires alterations in developmental processes, constrains evolutionary options, imposes irreversibility, and establishes the discrete nature of larval patterns in marine invertebrates.     

Community ecological modelling as an alternative to physiographic classifications for marine conservation planning

Accurate mapping of marine species and habitats is an important yet challenging component of establishing networks of representative marine protected areas. Due to limited biological data, marine classifications based on abiotic data are often used as surrogates to represent biological patterns. We tested the surrogacy of an existing physiographic marine classification using non-metric multidimensional scaling and permutational analysis of variance to determine whether species composition was significantly different among physiographic units. We also present an alternative ecological classification that incorporates biological and environmental data in a community modeling approach. We use data on 174 species of demersal fish and benthic invertebrates to identify mesoscale biological assemblages in a 100,000 km² study area in the northeast Pacific Ocean. We identified assemblages using cluster analysis then used a random forest model with 12 environmental variables to delineate mesoscale ecological units. Our community modelling approach resulted in five geographically coherent ecological units that were best explained by changes in depth, temperature and salinity. Our model showed high predictive performance (AUC = 0.93) and the resulting ecological units represent more distinct species assemblages than those delineated by physiographic variables alone. A strength of our analysis is the ability to map model uncertainty to identify transition zones at unit boundaries. The output of this study provides a biotic driven classification that can be used to better achieve representativity in the MPA planning process.     

From fronds to fish: the use of indicators for ecological monitoring in marine benthic ecosystems,with case studies from temperate Western Australia

Ecological indicators are used for monitoring in marine habitats the world over. With the advent of Ecosystem Based Fisheries Management (EBFM), the need for cost effective indicators of environmental impacts and ecosystem condition has intensified. Here, we review the development, utilisation and analysis of indicators for monitoring in marine benthic habitats, and outline important advances made in recent years. We use the unique, speciose benthic system of Western Australia (WA) as a detailed case study, as the development of indicators for EBFM in this region is presently ongoing, and major environmental drivers (e.g. climate change) and fishing practices are currently influencing WA marine systems. As such, the work is biased towards, but not restricted to, indicators that may be important tools for EBFM, such as biodiversity surrogates and indicators of fishing pressure. The review aimed to: (1) provide a concise, up-to-date account of the use of ecological indicators in marine systems; (2) discuss the current, and potential, applications of indicators for ecological monitoring in WA; and (3) highlight priority areas for research and pressing knowledge gaps. We examined indicators derived from benthic primary producers, benthic invertebrates and fish to achieve these goals.     

Frequency of injury and the ecology of regeneration in marine benthic invertebrates

Many marine invertebrates are able to regenerate lost tissue following injury, but regeneration can come at a cost to individuals in terms of reproduction, behavior and physiological condition, and can have effects that reach beyond the individual to impact populations, communities, and ecosystems. For example, removal and subsequent regeneration of clams' siphons, polychaetes' segments, and brittlestars' arms can represent significant energetic input to higher trophic levels. In marine soft-sediment habitats, injury changes infaunal bioturbation rates and thus secondarily influences sediment-mediated competition, adult-larval interactions, and recruitment success. The importance of injury and regeneration as factors affecting the ecology of marine invertebrate communities depends on the frequency of injury, as well as on individual capacity for, and speed of, regeneration. A key question to answer is: "How frequently are marine benthic invertebrates injured?" Here, I review the sources and the frequencies of injury in a variety of marine invertebrates from different benthic habitats, discuss challenges, and approaches for accurately determining injury rates in the field, consider evidence for species-specific, temporal and geographic variation in injury rates, and present examples of indirect effects of injury on marine invertebrates to illustrate how injury and regeneration can modify larger-scale ecological patterns and processes.     

Overview of the chemical ecology of benthic marine invertebrates along the western Antarctic peninsula

Thirteen years ago in a review that appeared in the American Zoologist, we presented the first survey of the chemical and ecological bioactivity of Antarctic shallow-water marine invertebrates. In essence, we reported that despite theoretical predictions to the contrary the incidence of chemical defenses among sessile and sluggish Antarctic marine invertebrates was widespread. Since that time we and others have significantly expanded upon the base of knowledge of Antarctic marine invertebrates' chemical ecology, both from the perspective of examining marine invertebrates in new, distinct geographic provinces, as well as broadening the evaluation of the ecological significance of secondary metabolites. Importantly, many of these studies have been framed within established theoretical constructs, particularly the Optimal Defense Theory. In the present article, we review the current knowledge of chemical ecology of benthic marine invertebrates comprising communities along the Western Antarctic Peninsula (WAP), a region of Antarctica that is both physically and biologically distinct from the rest of the continent. Our overview indicates that, similar to other regions of Antarctica, anti-predator chemical defenses are widespread among species occurring along the WAP. In some groups, such as the sponges, the incidence of chemical defenses against predation is comparable to, or even slightly higher than, that found in tropical marine systems. While there is substantial knowledge of the chemical defenses of benthic marine invertebrates against predators, much less is known about chemical anti-foulants. The sole survey conducted to date suggests that secondary metabolites in benthic sponges are likely to be important in the prevention of fouling by benthic diatoms, yet generally lack activity against marine bacteria. Our understanding of the sensory ecology of Antarctic benthic marine invertebrates, despite its great potential, remains in its infancy. For example, along the WAP, community-level non-consumptive effects occur when amphipods chemically sense fish predators and respond by seeking refuge in chemically-defended macroalgae. Such interactions may be important in releasing amphipods from predation pressure and facilitating their unusually high abundances along the WAP. Moreover, recent studies on the sensory biology of the Antarctic keystone sea star Odontaster validus indicate that chemotactile-mediated interactions between conspecifics and other sympatric predatory sea stars may have significant ramifications in structuring community dynamics. Finally, from a global environmental perspective, understanding how chemical ecology structures marine benthic communities along the WAP must increasingly be viewed in the context of the dramatic impacts of rapid climatic change now occurring in this biogeographic region.     

A preliminary investigation of diversity,abundance, and distributional patterns of chitons in intertidal boulder fields of differing rock type in South Australia

The rock type of hard substrata marine habitats can affect numerous benthic invertebrates, but little is known of the effects on molluscan assemblages, for example, the chitons often found under intertidal boulders. We compared chiton assemblage composition, abundance, species richness, and patterns of frequency distribution in 10 boulder fields containing either hard metamorphic/igneous boulders or soft limestone boulders in two geographical areas in South Australia. Similar species richness occurred in both types of boulder fields, but hard rock boulder fields had greater overall abundances, because of particularly large abundances of some common species. Differences in abundances of common species also resulted in significantly different assemblages occurring between the boulder field types. Some species appeared aggregated among boulders, but this pattern was variable between boulders in differing areas and of differing rock type. In one area, a common species had variable aggregation that caused frequency distributions to differ significantly between boulders of different rock types. These results indicate that rock type needs to be considered when investigating ecological patterns and processes involving specialist molluscs such as under-boulder chitons and to ensure comprehensive marine reserve planning for protecting rare invertebrates in rocky intertidal reefs.     

Evolutionary innovations in the fossil record: the intersection of ecology, development, and macroevolution

The origins of evolutionary innovations have been intensively studied, but relatively little is known about their large-scale ecological patterns. For post-Paleozoic benthic marine invertebrates, which have the richest and most densely sampled fossil record, order-level taxa tend to appear first in onshore, disturbed habitats, even in groups that are now exclusively deep-water (so that present-day distributions are not reliable indicators of original environments). New results presented here show that the onshore-origination pattern is robust to shifts in taxonomic methods and to new paleontological discoveries, and the few available studies suggest that this pattern can also be seen in terms of excursions in morphospace or the acquisition of derived character states, without reference to taxonomic categories. The environmental pattern at high levels contrasts significantly with the origin of low-level novelties (such as defined genera and families) in crinoids, echinoids, and bryozoans, where first appearances tend to conform to their clade-specific bathymetric diversity gradients. This discordance seems to eliminate potential driving mechanisms that simply scale up within-population genetic or ecological processes. Little is known about the factors that promote the onshore-offshore expansion of orders across the continental shelf, or that drive some clades to abandon ancestral habitats for an exclusively deep-water distribution. The origin of evolutionary innovation must ultimately reside in developmental changes, but the onshore-origination bias could emerge from two different dynamics: the pattern could be primarily genetic and developmental, i.e., innovations truly arise onshore; or primarily ecological, i.e., innovations arise randomly but preferentially survive onshore. Whatever the ultimate driving mechanisms, these macroevolutionary patterns show that theories of large-scale evolutionary novelty must include an ecological dimension.     

Ecological regulation of development: induction of marine invertebrate metamorphosis

In the marine environment a wide range of invertebrates have a pelagobenthic lifecycle that includes planktonic larval and benthic adult phases. Transition between these morphologically and ecologically distinct phases typically occurs when the developmentally competent larva comes into contact with a species-specific environmental cue. This cue acts as a morphogenetic signal that induces the completion of the postlarval/juvenile/adult developmental program at metamorphosis. The development of competence often occurs hours to days after the larva is morphologically mature. In the non-feeding--lecithotrophic--larvae of the ascidian Herdmania curvata and the gastropod mollusc Haliotis asinina, gene expression patterns in pre-competent and competent stages are markedly different, reflecting the different developmental states of these larval stages. For example, the expression of Hemps, an EGF-like signalling peptide required for the induction of Herdmania metamorphosis, increases in competent larvae. Induction of settlement and metamorphosis results in further changes in developmental gene expression, which apparently is necessary for the complete transformation of the larval body plan into the adult form.     

A comparative analysis of metabarcoding and morphology‐based identification of benthic communities across different regional seas

In a world of declining biodiversity, monitoring is becoming crucial. Molecular methods, such as metabarcoding, have the potential to rapidly expand our knowledge of biodiversity, supporting assessment, management, and conservation. In the marine environment, where hard substrata are more difficult to access than soft bottoms for quantitative ecological studies, Artificial Substrate Units (ASUs) allow for standardized sampling. We deployed ASUs within five regional seas (Baltic Sea, Northeast Atlantic Ocean, Mediterranean Sea, Black Sea, and Red Sea) for 12–26 months to measure the diversity and community composition of macroinvertebrates. We identified invertebrates using a traditional approach based on morphological characters, and by metabarcoding of the mitochondrial cytochrome oxidase I (COI) gene. We compared community composition and diversity metrics obtained using the two methods. Diversity was significantly correlated between data types. Metabarcoding of ASUs allowed for robust comparisons of community composition and diversity, but not all groups were successfully sequenced. All locations were significantly different in taxonomic composition as measured with both kinds of data. We recovered previously known regional biogeographical patterns in both datasets (e.g., low species diversity in the Black and Baltic Seas, affinity between the Bay of Biscay and the Mediterranean). We conclude that the two approaches provide complementary information and that metabarcoding shows great promise for marine monitoring. However, until its pitfalls are addressed, the use of metabarcoding in monitoring of rocky benthic assemblages should be used in addition to classical approaches rather than instead of them.     

Adaptive evolution of larvae and life cycles

The larval patterns of marine invertebrates pose intriguing questions for both evolutionary and developmental biologists. However, combined investigations have been rare. Quantitative models analyze the selective factors that drive evolutionary change in larval nutrition and timing of metamorphosis. Developmental studies describe the morphogenesis characterizing ancestral and derived larval patterns. Rigorous evolutionary analysis of the transition to derived modes of development is lacking and detailed developmental and ecological data are needed to test and refine theoretical models. A major challenge facing studies of life cycle evolution is the elucidation of the genetic structure and covariance of important developmental and larval traits.     

Effects of capability for dispersal on the evolution of diversity in antarctic benthos

The likelihood of marine invertebrates to maintain large geographic ranges is widely dependent on the ability of their early ontogenetic stages to disperse over long distances. Marine benthic invertebrates inhabiting the cold-stenothermal environment of the Southern Ocean are known for their overall reduced number of pelagic larvae, or drifting stages of any kind, when compared with organisms elsewhere in the sea. The diversity of organisms thriving in Antarctic waters is the result of evolution in situ and of the intrusion of species from surrounding seas. The reasons for a high level of endemism and a stunning diversity of benthic invertebrates found today are frequently discussed in the literature, but the mechanisms whereby diversity has been controlled over time remain largely theoretical. Here, I suggest that, indeed, early life-history patterns play a key role in defining the radiation and the speciation potential of Antarctic benthic invertebrates. In arguing this case, I synthesize the growing body of molecular studies on population connectivity in Antarctic benthic invertebrates, and compare this information with knowledge of their life histories and biogeography. I conclude that differences in early life-history patterns are key to the resilience potential of species in response to late Cenozoic glacial periods and propose that there is a direct relationship between rate of speciation and the ability of taxa to disperse.     

Beyond corals and fish: the effects of climate change on noncoral benthic invertebrates of tropical reefs

Climate change is threatening tropical reefs across the world, with most scientists agreeing that the current changes in climate conditions are occurring at a much faster rate than in the past and are potentially beyond the capacity of reefs to adapt and recover. Current research in tropical ecosystems focuses largely on corals and fishes, although other benthic marine invertebrates provide crucial services to reef systems, with roles in nutrient cycling, water quality regulation, and herbivory. We review available information on the effects of environmental conditions associated with climate change on noncoral tropical benthic invertebrates, including inferences from modern and fossil records. Increasing sea surface temperatures may decrease survivorship and increase the developmental rate, as well as alter the timing of gonad development, spawning, and food availability. The broad latitudinal distribution and associated temperature ranges of several pantropical taxa suggest that some reef communities may have an in‐built adaptive capacity. Tropical benthic invertebrates will also show species‐specific sublethal and lethal responses to sea‐level rise, ocean acidification, physical disturbance, runoff, turbidity, sedimentation, and changes in ocean circulation. In order to accurately predict a species' response to these stressors, we must consider the magnitude and duration of exposure to each stressor, as well as the physiology, mobility, and habitat requirements of the species. Stressors will not act independently, and many organisms will be exposed to multiple stressors concurrently, including anthropogenic stressors. Environmental changes associated with climate change are linked to larger ecological processes, including changes in larval dispersal and recruitment success, shifts in community structure and range extensions, and the establishment and spread of invasive species. Loss of some species will trigger economic losses and negative effects on ecosystem function. Our review is intended to create a framework with which to predict the vulnerability of benthic invertebrates to the stressors associated with climate change, as well as their adaptive capacity. We anticipate that this review will assist scientists, managers, and policy‐makers to better develop and implement regional research and management strategies, based on observed and predicted changes in environmental conditions.     

Biodiversity gradient in the Baltic Sea: a comprehensive inventory of macrozoobenthos data

In the Helsinki Commission Red List project 2009–2012, taxonomic and distributional data of benthic (macro) invertebrates were compiled by the present authors in a comprehensive checklist of the Baltic Sea fauna. Based on the most recent and comprehensive data, this paper presents the diversity patterns observed among benthic invertebrates in the Baltic Sea. As expected, the total number of species per sub-region generally declined along the salinity gradient from the Danish Straits to the northern Baltic Sea. This relationship is well known from the Baltic Sea and has resulted in a general assumption of an exponentially positive relationship between species richness and salinity for marine species, and a negative relationship for freshwater species. In 1934, Remane produced a diagram to describe the hypothetical distribution of benthic invertebrate diversity along a marine–freshwater salinity gradient. Our results clearly indicated the validity of this theory for the macrozoobenthic diversity pattern within the Baltic Sea. Categorisation of sub-regions according to species composition showed both separation and grouping of some sub-regions and a strong alignment of similarity patterns of zoobenthic species composition along the salinity gradient.     

Applicability of ecological evaluation tools in estuarine ecosystems: the case of the lower Mondego estuary (Portugal)

In accordance with the Water Framework Directive guidelines (WFD, 2000, European Communities Official Journal L327 2000/60/EC), classification schemes and ecological evaluation tools (based on benthic invertebrate fauna data sets from 1990 to 2002) were applied in the lower Mondego estuary. Two distinct scenarios could be tested due to the implementation of mitigation practices in 1999, following a long eutrophication process, which started by the early 1980s. Some discrepancies in the results were found by the application of the different indices. The AMBI index (accounting for taxonomic composition) and the ABC method (accounting for abundance and biomass k-dominance patterns) classifications often disagreed with those based on species diversity (Margalef and Shannon-Wiener). The ambiguous results made the classification a complex task to achieve, contrary to the Directive’s objective of maintaining it simple and clear. Our results suggest the necessity of adjusting some of the indices and their ranges to estuarine characteristics, namely to account the typical dominance and abundance of some particular species. These aspects are not taken into consideration by some of the indices proposed, which are more adapted to typical marine conditions. Based on our results, these widely applied indices might still improve their efficiency in estuarine systems allowing their use in the resembling types already established within the new Directive agenda.     

Differences in heat shock protein 70 expression during larval and early spat development in the Eastern oyster, Crassostrea virginica (Gmelin, 1791)

For a variety of species, changes in the expression of heat shock proteins (HSP) have been linked to key developmental changes, i.e., gametogenesis, embryogenesis, and metamorphosis. Many marine invertebrates are known to have a biphasic life cycle where pelagic larvae go through settlement and metamorphosis as they transition to the benthic life stage. A series of experiments were run to examine the expression of heat shock protein 70 (HSP 70) during larval and early spat (initial benthic phase) development in the Eastern oyster, Crassostrea virginica. In addition, the impact of thermal stress on HSP 70 expression during these early stages was studied. C. virginica larvae and spat expressed three HSP 70 isoforms, two constitutive, HSC 77 and HSC 72, and one inducible, HSP 69. We found differences in the expression of both the constitutive and inducible forms of HSP 70 among larval and early juvenile stages and in response to thermal stress. Low expression of HSP 69 during early larval and spat development may be associated with the susceptibility of these stages to environmental stress. Although developmental regulation of HSP 70 expression has been widely recognized, changes in its expression during settlement and metamorphosis of marine invertebrates are still unknown. The results of the current study demonstrated a reduction of HSP 70 expression during settlement and metamorphosis in the Eastern oyster, C. virginica.     

Large-scale species-richness gradients in the Atlantic Ocean

The increase in species richness from the poles to the Equator has been observed in numerous terrestrial and aquatic taxa. A number of different hypotheses have been put forward as explanations for this trend, e.g. area and energy availability. However, whether these hypotheses apply to large spatial scales in marine environments remains unclear. The present study shows a clear latitudinal gradient from high to low latitude (from 80 degrees N to 70 degrees S) in marine species richness for 6643 species (fishes and invertebrates) in 10 different taxa dwelling in benthic and pelagic habitats on both sides of the Atlantic. The patterns in benthic taxa are strongly influenced by coastal hydrographic processes, with marked peaks and troughs, and consequently the gradients are not symmetric along both Atlantic sides. Pelagic taxa show a plateau-shaped distribution and the influence from coastal events on gradients could not be demonstrated. The relationships between species richness and different environmental factors indicate that area size does not explain the latitudinal pattern in benthic species richness on a large spatial scale. Sea-surface temperature (positive relationship) is the best predictor of this pattern for benthic species, and nitrate concentration (negative relationship) is the best predictor for pelagic species. The results call into question the existence of a single primary cause that would explain the pattern in marine species richness on a large spatial scale.     

Functional morphogenesis from embryos to adults: Late development shapes trophic niche in coral reef damselfishes

The damselfishes are one of the dominant coral reef fish lineages. Their ecological diversification has involved repeated transitions between pelagic feeding using fast bites and benthic feeding using forceful bites. A highly‐integrative approach that combined gene expression assays, shape analyses, and high‐speed video analyses was used to examine the development of trophic morphology in embryonic, larval, juvenile, and adult damselfishes. The anatomical characters that distinguish pelagic‐feeding and benthic‐feeding species do not appear until after larval development. Neither patterns of embryonic jaw morphogenesis, larval skull shapes nor larval bite mechanics significantly distinguished damselfishes from different adult trophic guilds. Analyses of skull shape and feeding performance identified two important transitions in the trophic development of a single species (the orange clownfish; Amphiprion percula): (a) a pronounced transformation in feeding mechanics during metamorphosis; and (b) more protracted cranial remodeling over the course of juvenile development. The results of this study indicate that changes in postlarval morphogenesis have played an important role in damselfish evolution. This is likely to be true for other fish lineages, particularly if they consist of marine species, the majority of which have planktonic larvae with different functional requirements for feeding in comparison to their adult forms.     

福建平潭岛海域底栖贝类物种多样性及其地理分布

海洋底栖贝类是海洋底栖无脊椎动物的重要代表。福建平潭岛地处台湾海峡西岸北部, 是中国大陆距台湾岛最近之处, 其海域内的底栖贝类区系对了解台湾海峡及其附近海域的生态环境和底栖生物地理格局具有重要意义。本文基于对2015-2018年间中山大学国土资源与环境系在平潭岛海域采集的3,346号底栖贝类标本的鉴定, 结合已发表文献整理出了平潭岛海域底栖贝类名录, 并梳理了各物种在中国近海的地理分布类型。结果显示, 本研究所采标本分属58科122属161种, 其中47种为平潭岛海域新记录。结合历史文献记录, 平潭岛海域共有底栖贝类98科244属395种。其中93.7%的种类在南海亦有分布, 82.0%的种类在东海有分布, 31.1%的种类在黄渤海有分布, 42.3%的种类在台湾岛东部海域有分布。南海-东海分布型种类占区系总种数的49.9%; 其次为南海-黄渤海分布型(27.6%)和南海-台湾海峡分布型(16.2%)。分布限于台湾海峡及其以北的种类仅占总种数的6.3%。研究结果表明, 平潭岛海域底栖贝类区系具典型的亚热带性质, 暖水性种类和广温性种类占主要优势, 与南海北部和东海大陆沿岸地区的关系密切, 与台湾岛周边海域的联系相对较弱。     

Deconstructing latitudinal species richness patterns in the ocean: does larval development hold the clue?

The consistent decrease in species richness with latitude shows several exceptions among marine organisms. We hypothesize that contrasting latitudinal diversity gradients can be explained by differences in critical life-history attributes, such as mode of larval development (MLD). We deconstructed latitudinal species richness patterns of marine benthic invertebrates according to MLD to elucidate differences in patterns of species richness and to reveal underlying processes. The patterns of species richness were remarkably similar across taxa within MLD but differed between MLD. Species richness decreased polewards in planktotrophic species and increased in direct developers. Temperature explained most of the variation in species richness. Low temperature at high latitudes may generally favour direct developing species, but, together with low chlorophyll- a concentration, limit the distribution of planktotrophic species. The contrasting influence of temperature on different MLDs might be explained by its effect on the length of planktonic life and on brooding costs.     

Sulphur stable isotopes can distinguish trophic dependence on sediments and plankton in boreal lakes

1. Stable isotopes of carbon are useful for differentiating between freshwater food chains based on planktonic algae or benthic algae, but are reported to be of limited use for identifying food chains based on sedimentary detritus. Because data from marine systems suggest that stable isotopes of sulphur (δ³⁴S values) have potential in this regard, we tested their utility in freshwater lakes.
2. We found that sulphate in the water column of four boreal lakes was enriched in ³⁴S compared to the sulphur in bulk sediments from these lakes. Furthermore, within a given lake, insects known to feed on sediment (directly or via predation) had δ³⁴S values similar to those of sediment, whereas planktonic and benthic invertebrates known to feed on suspended particles had δ³⁴S values similar to those of sulphate in the water column.
3. Using the stable S isotope values of invertebrates that obtain their S from either the sediment or the water column as end members in a two-source mixing model, we show that two fish species obtain their food from both planktonic and sedimentary sources. Furthermore, model results suggest that, as expected, the more benthic-feeding fish species obtains more of its S from the sediment compartment than does the species that feeds in the water-column.
4. Our results suggest that measurements of stable sulphur isotopes provide a means of distinguishing between members of food chains that are based in the water column from those based on sedimentary detritus. As such, they would be a useful complement to stable C isotopes that are used to distinguish between food chains based on planktonic or benthic algae.