Top‐Down Control of Reed Detritus Processing in a Lake Littoral Zone: Experimental Evidence of a Seasonal Compensation between Fish and Invertebrate Predation

We investigated whether predatory fish exert a top‐down control on reed leaf packs processing in a lake littoral zone through a trophic cascade. Exclosure experiments were repeated in summer and winter, under high and low natural fish abundance, respectively. Fish exclusion effects on detritus processing and fungal conditioning were consistent with trophic cascade predictions only in summer. In winter, however, results indicated that a trophic cascade was induced by predatory invertebrates. In both seasons, variations in detritivores abundance generally supported a cascade scenario, whereas several taxon‐specific departures occurred during the experimental periods. We conclude suggesting that predators may continuously regulate leaf detritus processing in lake littoral zones, through a seasonal shift in the relative contribution of fish and invertebrate predation. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effects of a top invertebrate predator (<Emphasis Type="Italic">Dytiscus alaskanus</Emphasis>; Coleoptera: Dytiscidae) on fishless pond ecosystems

We investigated the predatory effects of Dytiscus alaskanus, a large predaceous diving beetle, on the biomass, species composition and diversity of fishless pond communities. The effects were tested using presence and absence treatments of D. alaskanus in 24 mesocosms distributed among six ponds. We sampled phytoplankton, zooplankton and macroinvertebrates every two weeks for a six week period. Periphyton was sampled from the mesocosm walls on the final day. Total macroinvertebrate biomass decreased in the presence of dytiscids while species richness was not affected. Macroinvertebrate predators, snails and Gammarus lacustris decreased in the dytiscid treatments. Laboratory feeding experiments confirmed feeding preferences consistent with the mesocosm results. Periphyton biomass was six times greater in the dytiscid enclosures, concomitant with the decreased grazing by gastropods and other invertebrate primary consumers indicating a benthic trophic cascade. Top–down effects of dytiscids on other predatory invertebrates led to increased total zooplankton biomass, largely due to increased abundances of large and small cladocerans. Zooplankton species richness increased in the dytiscid enclosures. Inconsistent with trophic cascade theory, phytoplankton did not respond to top–down effects of D. alaskanus within the study period. Overall, the results show D. alaskanus predation caused trophic effects via two distinct food chains, a dytiscid–snail–periphyton trophic cascade, and a dytiscid–predatory macroinvertebrates–zooplankton partial trophic cascade.     

Marine reserves indirectly affect fine‐scale habitat associations,but not overall densities,of small benthic fishes

Many large, fishery‐targeted predatory species have attained very high relative densities as a direct result of protection by no‐take marine reserves. Indirect effects, via interactions with targeted species, may also occur for species that are not themselves targeted by fishing. In some temperate rocky reef ecosystems, indirect effects have caused profound changes in community structure, notably the restoration of predator–urchin–macroalgae trophic cascades. Yet, indirect effects on small benthic reef fishes remain poorly understood, perhaps because of behavioral associations with complex, refuge‐providing habitats. Few, if any, studies have evaluated any potential effects of marine reserves on habitat associations in small benthic fishes. We surveyed densities of small benthic fishes, including some endemic species of triplefin (Tripterygiidae), along with fine‐scale habitat features in kelp forests on rocky reefs in and around multiple marine reserves in northern New Zealand over 3 years. Bayesian generalized linear mixed models were used to evaluate evidence for (1) main effects of marine reserve protection, (2) associations with habitat gradients, including complexity, and (3) differences in habitat associations inside versus outside reserves. No evidence of overall main effects of marine reserves on species richness or densities of fishes was found. Both richness and densities showed strong associations with gradients in habitat features, particularly habitat complexity. In addition, some species exhibited reserve‐by‐habitat interactions, having different associations with habitat gradients inside versus outside marine reserves. Two species (Ruanoho whero and Forsterygion flavonigrum) showed stronger positive associations with habitat complexity inside reserves. These results are consistent with the presence of a behavioral risk effect, whereby prey fishes are more strongly attracted to habitats that provide refuge from predation in areas where predators are more abundant. This work highlights the importance of habitat structure and the potential for fishing to affect behavioral interactions and the interspecific dynamic attributes of community structure beyond simple predator–prey consumption and archetypal trophic cascades.     

Marine reserves demonstrate top-down control of community structure on temperate reefs

Replicated ecological studies in marine reserves and associated unprotected areas are valuable in examining top-down impacts on communities and the ecosystem-level effects of fishing. We carried out experimental studies in two temperate marine reserves to examine these top-down influences on shallow subtidal reef communities in northeastern New Zealand. Both reserves examined are known to support high densities of predators and tethering experiments showed that the chance of predation on the dominant sea urchin, Evechinus chloroticus, within both reserves was approximately 7 times higher relative to outside. Predation was most intense on the smallest size class (30-40 mm) of tethered urchins, the size at which urchins cease to exhibit cryptic behaviour. A high proportion of predation on large urchins could be attributed to the spiny lobster, Jasus edwardsii. Predation on the smaller classes was probably by both lobsters and predatory fish, predominantly the sparid Pagrus auratus. The density of adult Evechinus actively grazing the substratum in the urchin barrens habitat was found to be significantly lower at marine reserve sites (2.2ǂ.3 m^-2) relative to non-reserve sites (5.5ǂ.4 m^-2). There was no difference in the density of cryptic juveniles between reserve and non-reserve sites. Reserve populations were more bimodal, with urchins between 40 and 55 mm occurring at very low numbers. Experimental removal of Evechinus from the urchin barrens habitat over 12 months lead to a change from a crustose coralline algal habitat to a macroalgal dominated habitat. Such macroalgal habitats were found to be more extensive in both reserves, where urchin densities were lower, relative to the adjacent unprotected areas that were dominated by urchin barrens. The patterns observed provide evidence for a top-down role of predators in structuring shallow reef communities in northeastern New Zealand and demonstrate how marine reserves can reverse the indirect effects of fishing and re-establish community-level trophic cascades.     

Trophic interactions and community structure in the upwelling system off Central Chile (33-39°S)

Trophic interactions and community structure in the upwelling system off Central Chile (USCCh) (33-39°S) are analyzed using biological and ecological data concerning the main trophic groups and the Ecopath with Ecosim software version 5.0 (EwE). The model encompasses the fisheries, cetaceans, sea lion, marine birds, cephalopods, large-sized pelagic fish (sword fish), medium-sized pelagic fish (horse mackerel, hoki), small-sized pelagic fish (anchovy, common sardine), demersal fish (e.g. Chilean hake, black conger-eel), benthic invertebrates (red squat lobster, yellow squat lobster) and other groups such as zooplankton, phytoplankton and detritus. Input data was gathered from published and unpublished reports and our own estimates. Trophic interactions, system indicators and food web attributes are calculated using network analysis routines included in EwE. Results indicate that trophic groups are aligned around four trophic levels (TL) with phytoplankton and detritus at the TL=1, while large-sized pelagic fish and cetaceans are top predators (TL>4.0). The fishery is located at an intermediate to low trophic level (TL=2.97), removing about 15% of the calculated system primary production. The pelagic realm dominates the system, with medium-sized pelagic fish as the main fish component in biomass, while small-sized pelagic fish dominate total landings. Chilean hake is by far the main demersal fish component in both, biomass and yield. Predators consume the greater part of the production of the most important fishery resources, particularly juvenile stages of Chilean hake. Consequently, mortality by predation is an important component of total mortality. However, fishery also removes a large fraction of common sardine, anchovy, horse mackerel, and Chilean hake. The analysis of direct and indirect trophic impacts reveals that Chilean hake is a highly cannibalistic species. Chilean hake is also an important predator on anchovy, common sardine, benthic invertebrates, and demersal fish. The fisheries heavily impact on Chilean hake, common sardine, anchovy, and horse mackerel. Total system biomass (B=476 t km⁻² year⁻¹) and throughput (T=89454 t km⁻² year⁻¹) estimated in the USCCh model are in accordance with models of comparable systems. Considering system attributes derived from network analysis, the USCCh can be characterized as an immature system, with short trophic chains and low trophic transfer efficiency. Finally, we suggest that trophic interactions should be considered in stock assessment and management programs in USCCh. In addition, future research programs should be carried out in order to understand the ecosystem effects of fishing and trophic control in this highly productive food web.     

Testing the generality of the trophic cascade paradigm for sea otters: a case study with kelp forests in northern Washington,USA

Trophic cascade hypotheses for biological communities, linking predation by upper trophic levels to major features of ecological structure and dynamics at lower trophic levels, are widely subscribed and may influence conservation policy. Few such hypotheses have been evaluated for temporal or spatial generality. Previous studies of sea otter (Enhydra lutris) predation along the outer coast of North America suggest a pattern, often elevated to the status of paradigm, in which sea otter presence leads to reduced sea urchin (Strongylocentrotus spp.) biomass and rapid increases in abundance and diversity of annual algal species, followed by a decline in diversity as one or a few perennial algal species become dominant. Both sea otter predation and commercial sea urchin harvest are ecologically and economically important sources of urchin mortality in nearshore benthic systems in northern Washington marine waters. We recorded changes in density of macroalgae in San Juan Channel, a marine reserve in the physically protected inland waters of northern Washington, resulting from three levels of experimental urchin harvest: (1) simulated sea otter predation (monthly complete harvest of sea urchins), (2) simulated commercial urchin harvest (annual size-selective harvest of sea urchins), and (3) no harvest (control). The two experimental urchin removal treatments did not significantly increase the density of perennial (Agarum and Laminaria) or annual (Desmarestia, Costaria, Alaria and Nereocystis) species of macroalgae after 2 years, despite significant and persistent decreases in urchin densities. Our results suggest that other factors such as grazing by other invertebrates, the presence of dense Agarum stands, and recruitment frequency of macroalgae and macroinvertebrates may play a large role in influencing community structure in San Juan Channel and other physically protected marine waters within the range of sea otters. Handling editor: J. Trexler     

The dual importance of competition and predation as regulatory forces in terrestrial ecosystems: evidence from decomposer food-webs

The relative importance of predation and competition (resource limitation) in influencing the components of a below-ground food-web consisting of three trophic levels (bacteria and fungi; bacterial-feeding and fungal-feeding nematodes; and top predatory nematodes) was estimated using microbial biomass and nematode frequency data collected throughout a 1-year period in two agro-ecosystems. The study suggested that bacterial and fungal biomass were likely to be regulated by grazing and competition respectively, and that these differences were likely to be attributed to the biological (probably morphological) differences between bacteria and fungi, in contrast to the predictions of the hypothesis of Hairston et al. (1960). Top predatory nematodes were sometimes strongly related to the microbial but not microbial-feeding trophic levels, indicating that microbial biomass may directly influence top predator numbers, and that the intermediate level may simply serve as a conduit by which resources pass from the bottom to top trophic levels. This study also suggests that the detritus food-web acts as two distinct (bacterial-and fungal-based) compartments.     

The role and contribution of the seagrass Posidonia oceanica (L.) Delile organic matter for secondary consumers as revealed by carbon and nitrogen stable isotope analysis

The δ¹³C and δ¹⁵N values of primary producers and consumers were studied to obtain information on the trophic role of Posidonia oceanica L. Delile, the dominant primary producer, in a Mediterranean shallow environment (the Stagnone di Marsala, western Sicily). δ¹³C strongly discriminated between pelagic and benthic pathways, with the former based on phytoplankton and the latter on a mixed pool of seagrass detritus, epiphytes and benthic algae as carbon sources. A particularly important trophic role appears to be performed by the vegetal epiphytic community on seagrass leaves (δ¹³C = –14.9 ± 0.1‰), which supports most of the faunal seagrass community (i.e. Amphipoda, Isopoda, Tanaidacea; δ¹³C = –14.9 ± 0.1‰, –12.5 ± 0.1‰ and –14.8 ± 1.0‰, respectively). Although P. oceanica (δ¹³C = –11.3 ± 0.3‰) does not seem to be utilised by consumers via grazing (apart from a few Palaemonidae species with δ¹³C value of –10.8 ± 1.8‰), its trophic role may be via detritus. P. oceanica detritus may be exploited as a carbon source by small detritivore invertebrates, and above all seems to be exploited as a nitrogen reservoir by both bottom and water column consumers determining benthic–pelagic coupling. At least three trophic levels were detected in both the pelagic (mixture of phytoplankton and cyanobacteria, zooplankton, juvenile transient fish) and benthic (sedimentary organic matter and epiphytes, small seagrass-associated invertebrates, larger invertebrates and adult resident fish) pathways.     

The Coupling Between Grazing and Detritus Food Chains and the Strength of Trophic Cascades Across a Gradient of Nutrient Enrichment

A minimal food web model was constructed comprising one grazing and one detritus food chain coupled by nutrient cycling and generalist carnivores to investigate how prey preference by carnivores may affect the strength of trophic cascades across a gradient of nutrient enrichment. The equilibrium or mean abundance of each food web component and the magnitude of the carnivore effect on lower trophic levels were calculated for different values of the prey preference and nutrient input parameters. Our model predicts that nutrient enrichment increases the mean abundances of carnivores, autotrophs and detritus, but the magnitude of this effect is dependent on the prey preference term. On the other hand, herbivores and detritivores are relatively unaffected by enrichment but are strongly affected by carnivore preference. Carnivores have a negative effect on herbivores and a positive effect on autotrophs and detritus, whereas the effect on detritivores can be both positive and negative. At high preference for herbivores, carnivores have a positive effect on detritivores, because the positive effect of increased detritus availability due to reduced herbivore grazing outweighs the negative effect of predation. At high preference for detritivores, the balance is changed in the other direction. We argue that in systems where authochtonous primary production is the major source of detritus, herbivores can control the rates of detritus production and have indirect effects on detritivores, which may feed back into effects on herbivores through their shared enemies. This positive feedback is probably one mechanism affecting the resilience of alternative stable states in shallow lakes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  J.L.A. conceived the model and the study, J.R. wrote the Matlab programs and ran the simulations and J.L.A. wrote most of the paper.     

Trophic Cascades Induced by Lobster Fishing Are Not Ubiquitous in Southern California Kelp Forests

Fishing can trigger trophic cascades that alter community structure and dynamics and thus modify ecosystem attributes. We combined ecological data of sea urchin and macroalgal abundance with fishery data of spiny lobster (Panulirus interruptus) landings to evaluate whether: (1) patterns in the abundance and biomass among lobster (predator), sea urchins (grazer), and macroalgae (primary producer) in giant kelp forest communities indicated the presence of top-down control on urchins and macroalgae, and (2) lobster fishing triggers a trophic cascade leading to increased sea urchin densities and decreased macroalgal biomass. Eight years of data from eight rocky subtidal reefs known to support giant kelp forests near Santa Barbara, CA, USA, were analyzed in three-tiered least-squares regression models to evaluate the relationships between: (1) lobster abundance and sea urchin density, and (2) sea urchin density and macroalgal biomass. The models included reef physical structure and water depth. Results revealed a trend towards decreasing urchin density with increasing lobster abundance but little evidence that urchins control the biomass of macroalgae. Urchin density was highly correlated with habitat structure, although not water depth. To evaluate whether fishing triggered a trophic cascade we pooled data across all treatments to examine the extent to which sea urchin density and macroalgal biomass were related to the intensity of lobster fishing (as indicated by the density of traps pulled). We found that, with one exception, sea urchins remained more abundant at heavily fished sites, supporting the idea that fishing for lobsters releases top-down control on urchin grazers. Macroalgal biomass, however, was positively correlated with lobster fishing intensity, which contradicts the trophic cascade model. Collectively, our results suggest that factors other than urchin grazing play a major role in controlling macroalgal biomass in southern California kelp forests, and that lobster fishing does not always catalyze a top-down trophic cascade.     

A test of the Menge-Sutherland model of community organization in a tropical rocky intertidal food web

Summary Menge and Sutherland (1976) predicted that in physically benign habitats: (1) community structure will be most strongly affected be predation, (2) the effect of predation will increase with a decrease in trophic position in the food web, (3) trophically intermediate species will be influenced by both predation and competition, and (4) competition will occur among prey species which successfully escape consumers. These predictions were tested in a tropical rocky intertidal community on the Pacific coast of Panama. The most abundant mobile species included fishes and crabs, which occupied the top trophic level, and predaceous gastropods and herbivorous molluscs, which occupied intermediate trophic levels. The most abundant sessile organisms were encrusting algae, foliose algae, barnacles, and bivalves. Diets were broad and overlapping, and 30.3% of the consumers were omnivorous. Each consumer group had strong effects on prey occurring at lower trophic levels: (1) Fishes and crabs reduced the abundance of predaceous snails, herbivorous molluscs, foliose algae, and sessile invertebrates. (2) Predaceous gastropods reduced the abundance of herbivorous molluscs and sessile invertebrates. (3) Herbivorous molluscs reduced the abundance of foliose algae and young stages of sessile invertebrates, and altered relative abundances of the encrusting algae. The encrusting algae, although normally the dominant space occupiers, proved to be inferior competitors for space with other sessile organisms when consumers were experimentally excluded. However, the crusts escaped consumers by virtue of superior anti-herbivore defenses and competed for space despite intense grazing. Observations do not support the hypothesis that the trophically intermediate species compete. Hence, with the exception of this last observation, the predictions of the Menge and Sutherland model were supported. Although further work is needed to evaluate other predictions of the model in this community, evidence from this study joins an increasing body of knowledge supporting the model. Contradictory evidence also exists, however, indicating that aspects of the model require revision.     

Sea urchins predation facilitates coral invasion in a marine reserve

Macroalgae is the dominant trophic group on Mediterranean infralittoral rocky bottoms, whereas zooxanthellate corals are extremely rare. However, in recent years, the invasive coral Oculina patagonica appears to be increasing its abundance through unknown means. Here we examine the pattern of variation of this species at a marine reserve between 2002 and 2010 and contribute to the understanding of the mechanisms that allow its current increase. Because indirect interactions between species can play a relevant role in the establishment of species, a parallel assessment of the sea urchin Paracentrotus lividus, the main herbivorous invertebrate in this habitat and thus a key species, was conducted. O. patagonica has shown a 3-fold increase in abundance over the last 8 years and has become the most abundant invertebrate in the shallow waters of the marine reserve, matching some dominant erect macroalgae in abundance. High recruitment played an important role in this increasing coral abundance. The results from this study provide compelling evidence that the increase in sea urchin abundance may be one of the main drivers of the observed increase in coral abundance. Sea urchins overgraze macroalgae and create barren patches in the space-limited macroalgal community that subsequently facilitate coral recruitment. This study indicates that trophic interactions contributed to the success of an invasive coral in the Mediterranean because sea urchins grazing activity indirectly facilitated expansion of the coral. Current coral abundance at the marine reserve has ended the monopolization of algae in rocky infralittoral assemblages, an event that could greatly modify both the underwater seascape and the sources of primary production in the ecosystem.     

Phytoplankton food quality control of planktonic food web processes

We developed a mechanistic model of nutrient, phytoplankton, zooplankton and fish interactions to test the effects of phytoplankton food quality for herbivorous zooplankton on planktonic food web processes. When phytoplankton food quality is high strong trophic cascades suppress phytoplankton biomass, the zooplankton can withstand intense zooplanktivory, and energy is efficiently transferred through the food web sustaining higher trophic level production. Low food quality results in trophic decoupling at the plant-animal interface, with phytoplankton biomass determined primarily by nutrient availability, zooplankton easily eliminated by fish predation, and poor energy transfer through the food web. At a given nutrient availability, food quality and zooplanktivory interact to determine zooplankton biomass which in turn determines algal biomass. High food quality resulted in intense zooplankton grazing which favored fast-growing phytoplankton taxa, whereas fish predation favored slow-growing phytoplankton. These results suggest algal food quality for herbivorous zooplankton can strongly influence the nature of aquatic food web dynamics, and can have profound effects on water quality and fisheries production. Handling editor: D. Hamilton     

Impacts of two invasive mollusks, <Emphasis Type="Italic">Rapana venosa</Emphasis> (Gastropoda) and <Emphasis Type="Italic">Corbicula fluminea</Emphasis> (Bivalvia), on the food web structure of the Río de la Plata estuary and nearshore oceanic ecosystem

This paper quantifies the impacts of two invasive species, Rapana venosa (Gastropoda, Muricidae) and Corbicula fluminea (Bivalvia, Corbiculidae), in the food web of the Río de la Plata estuary and adjacent nearshore oceanic ecosystem. We analyzed certain functional traits of these mollusks assessed by a mass balance trophic model previously constructed for the years 2005–2007. This model incorporates 37 functional groups: three marine mammals species, one coastal bird, 17 fishes, 12 invertebrates, two zooplankton, one phytoplankton, and detritus. The model also includes 5 fishing fleets operating in the area. The results showed that the two invasive species affect multiple ecosystem components both directly and indirectly. R. venosa and the whitemouth croaker, Micropogonias furnieri, exhibited a high level of niche overlap (91%), while C. fluminea exhibited a high level of niche overlap with Mytilidae (94%), which suggests in both cases high levels of competition for similar resources. R. venosa had mixed trophic impacts but exhibited a predominantly top down effect on most bivalves. R. venosa could be a threat to natural resources in the area including to the fishing fleets. C. fluminea negatively influenced phytoplankton and detritus biomass and its positive effects on higher trophic level groups suggest a central bottom-up role in the food web as a bentho-pelagic coupler. Both species had negative impacts on the five fleets modeled, showing that the effects of these invasive species could extend to the socio-economic sector.     

Functional significance of crayfish in stream food webs: roles of omnivory,substrate heterogeneity and sex

Heterogeneity of species interactions in food webs can result from characteristics of substrates as well as attributes of top consumers. We performed a streamside channel experiment to evaluate the impact of crayfish on lower trophic levels in detritus‐based (leaf packs) and algal‐based food webs (hard‐bottoms). After 43 days, both male and female crayfish had dramatically promoted leaf decomposition, with males processing material at a faster rate. However, the difference in leaf processing rates was not related to a greater level of male activity. Despite the sex‐related difference in residual leaf dry mass, densities of invertebrates in leaf packs were similarly low in the presence of crayfish of either sex, due to resource consumption, physical dislodgment (bioturbation) and/or predation. No trophic cascade was evident in the leaf pack assemblage. In the hard‐bottom assemblage, the results confirmed circumstantial field evidence that crayfish reduce predatory Tanypodinae and indirectly increase collector‐gatherer Chironominae following the prediction of a trophic cascade. However, no other taxa were indirectly facilitated, because of strong direct effects of crayfish on algal abundance (through direct consumption and bioturbation). Overall, impacts of crayfish on lower trophic levels were more pronounced in the structurally complex, detritus‐based assemblages than in its hard‐bottom, algal‐based counterpart. This conflicts with the expectation that net predation effects should be weaker where structural complexity is greater but is mainly a consequence of the profound engineering effects of crayfish in reducing colonisable substrate when they shred and disturb detrital material. Effects of crayfish may therefore propagate differently and with varying strength depending on substrate. Moreover, engineering activities and predation by crayfish appear to have been of overwhelming significance with subtle sex differences in leaf processing rates failing to lead to differences in invertebrate densities.     

Ecosystem-level evidence for top-down and bottom-up control of production in a grassland stream system

Ecosystem-wide effects of introduced brown trout (Salmo trutta L.) and native river galaxias (Galaxiaseldoni McDowall) were studied by analysing ecosystem production budgets for two adjacent tributaries of a grassland stream-system in the South Island of New Zealand. One tributary was inhabited by brown trout, the other by river galaxias. No other fish species were present in either stream. The budget for the river galaxias stream indicated little top-down control of invertebrates by fish predation (river galaxias consumed ∼18% of available prey production). A large proportion of annual net primary production was required to support production by invertebrates (invertebrates consumed an average of ∼75% of available primary production), and mean surplus primary production (i.e. not consumed) was not significantly different from zero. Primary and secondary production were presumably mutually limiting in this system (i.e. controlled by simultaneous top-down and bottom-up mechanisms). In contrast, the budget for the brown trout stream indicated extreme top-down control of invertebrate populations by fish predation; essentially all invertebrate production (∼100%) was required to support trout production. Invertebrate production required only a minor portion of annual net primary production (∼21%) and primary production was presumably controlled by mechanisms other than grazing (e.g. sloughing, nutrient limitation). Predatory invertebrates had little quantitative effect on prey populations in either stream. Recent experimental studies of invertebrate behaviour, fish behaviour, and food-web structure in New Zealand streams with physically stable channels indicate that a trophic cascade should be observed in streams inhabited by brown trout, in contrast to those inhabited by native fish. The results reported here provide ecosystem-level evidence supporting this prediction. Received: 10 March 1997 / Accepted: 12 December 1997     

Cascading Effects of Ocean Acidification in a Rocky Subtidal Community

Temperate marine rocky habitats may be alternatively characterized by well vegetated macroalgal assemblages or barren grounds, as a consequence of direct and indirect human impacts (e.g. overfishing) and grazing pressure by herbivorous organisms. In future scenarios of ocean acidification, calcifying organisms are expected to be less competitive: among these two key elements of the rocky subtidal food web, coralline algae and sea urchins. In order to highlight how the effects of increased pCO₂ on individual calcifying species will be exacerbated by interactions with other trophic levels, we performed an experiment simultaneously testing ocean acidification effects on primary producers (calcifying and non-calcifying algae) and their grazers (sea urchins). Artificial communities, composed by juveniles of the sea urchin Paracentrotus lividus and calcifying (Corallina elongata) and non-calcifying (Cystoseira amentacea var stricta, Dictyota dichotoma) macroalgae, were subjected to pCO₂ levels of 390, 550, 750 and 1000 µatm in the laboratory. Our study highlighted a direct pCO₂ effect on coralline algae and on sea urchin defense from predation (test robustness). There was no direct effect on the non-calcifying macroalgae. More interestingly, we highlighted diet-mediated effects on test robustness and on the Aristotle''s lantern size. In a future scenario of ocean acidification a decrease of sea urchins'' density is expected, due to lower defense from predation, as a direct consequence of pH decrease, and to a reduced availability of calcifying macroalgae, important component of urchins'' diet. The effects of ocean acidification may therefore be contrasting on well vegetated macroalgal assemblages and barren grounds: in the absence of other human impacts, a decrease of biodiversity can be predicted in vegetated macroalgal assemblages, whereas a lower density of sea urchin could help the recovery of shallow subtidal rocky areas affected by overfishing from barren grounds to assemblages dominated by fleshy macroalgae.     

The budget model of ecosystem of a shallow highly eutrophic lake

A model of energy budget of Lake Bolshoi Okunenok ecosystem was based on the data received during field studies from May through November 1986. The model takes into account 36 components including dissolved organic matter, bacteria, phytoplankton, zooplankton, meiobenthos, macrobenthos, fish, suspended and sediment detritus. The growing season has been divided into 16 intervals according to the number of observations. The balance equation for each live component describes the change in its biomass for a time interval between two successive sampling dates. The change is considered as a balance of energy input with assimilation or feeding, and energy loss due to respiration, excretion, predation, natural mortality, fishery catchment or and emergence of imago insects. For non-live components we estimate an increase and a decrease in their mass due to the activity of living organisms, as well as organic matter exchange between water and sediments. Seasonal value of balance elements for each component are equal to sums of appropriate interval value. Comparison of energy flows through different links of a trophic web has shown that the role of a bacterial-detrial link was extremely important in Lake Bolshoi Okunenok for the growth season of 1986. Detritus constituted 58% of seasonal diet of non-predatory zooplankton, 39% of diet of predatory zooplankton, 50% of diet of planktivorous fish (fry of whitefish) and 92% of diet of benthivorous fish (fry of carp). The contribution of bacteria to the total seasonal decomposition amounted to 46%. Approximately 57% of the forage phytoplankton production, 86% of non-predatory benthos production, and 23-38% of the other trophic groups production were consumed by all grazers. "Coefficient of energy transformation" is proposed. It is calculated as: CET(s, k) = Ps(k)/Pk, where Ps(k) is production of consumers "s", built due to consumption of source "k"; Pk is production of source "k" itself. In Lake Bolshoi Okunenok only 14% of energy built by phytoplankton were accumulated in organic matter of zooplankton due to direct consumption.     

Contributions of stable-isotope data to elucidating food webs of Mediterranean rocky littoral fishes

The food webs of rocky infra-littoral ecosystems in the Mediterranean have been little studied. In this investigation stable isotopes and dietary data were compared in an attempt to describe features of the food webs concerned. δ¹³C and δ¹⁵N were determined for plants, invertebrates and fishes from the Bay of Calvi, Corsica. Dietary data were derived from the literature. δ¹³C of plants ranged from –8.59‰ to –33.74‰, of benthic invertebrates from –17.0‰ to –20.52‰, of planktonic invertebrates from –20.08‰ to –22.34‰ and of fishes from –16.27‰ to –19.59‰. δ¹⁵N was generally greater at higher trophic levels. δ¹⁵N of plants was 0.95–2.92‰, of benthic invertebrates 1.69–6.54‰, of planktonic invertebrates 3.51–6.82‰ and of fishes 4.63–9.77‰. ¹³C enrichment tended to be associated with benthic food chains and ¹³C depletion with planktonic chains. Stable-isotope data suggested more varied diets for many species than implied by gut-contents data. Omnivory and trophic plasticity were widespread, and many consumers fed lower down the food chain than previous studies had suggested. Both stable-isotope and gut-contents analysis resolved differences between fishes feeding on planktonic and benthic prey and indicated that the herbivorous fish Sarpa salpa fed on a diet substantially different from that of other fishes. Zooplankton were important in the diets of several consumers (both primary and secondary), as was plankton derived detritus. One species of fish previously identified as planktivorous was shown to feed largely on benthic organisms, whilst several species of benthic invertebrates may feed on plankton-derived detritus. Although herbivores seemed to obtain most of their C from macroalgae, δ¹⁵N data suggested that many of these animals supplemented their intake of N, although gut-contents analysis did not provide evidence for such uptake. The isotopic data have elucidated several features of the food web which we would not otherwise have detected. Received: 26 April 1999 / Accepted: 24 September 1999     

Coupling of phytoplankton and detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

An oscillating steady state is described of phytoplankton, dominated by Prochlorothrix hollandica and Oscillatoria limnetica, and sestonic detritus in shallow, eutrophic Lake Loosdrecht (The Netherlands). A steady-state model for the coupling of the phytoplankton and detritus is discussed in relation to field and experimental data on phytoplankton growth and decomposition. According to model predictions, the phytoplankton to detritus ratio decreases hyperbolically at increasing phytoplankton growth rate and is independent of a lake's trophic state. The seston in L. Loosdrecht contains more detritus than phytoplankton as will apply to many other lakes. The model provides a basis for estimating the loss rate of the detritus, including decomposition, sedimentation and hydraulic loss. In a shallow lake like L. Loosdrecht detritus will continue to influence the water quality for years.