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     

The effect of bacterivorous nematodes on detritus incorporation by macrofaunal detritivores: A study using stable isotope and fatty acid analyses

Several studies have investigated the effect of nematodes on microbial degradation of macrophyte detritus, but little is known about the potential effect of bacterivorous nematodes on productivity of macrofaunal detritivores. We investigated the influence of the bacterivorous nematode Rhabditis (Pellioditis) mediterranea on the incorporation rate of isotopically-labelled macroalgal detritus by the amphipod Paramoera chevreuxi in a laboratory microcosm. The fatty acid composition of amphipods, nematodes, and macroalgal detritus was characterized to help determine the pathway of detritus incorporation by amphipods. The potential importance of R. mediterranea as a source of highly unsaturated fatty acids (HUFAs) to higher trophic levels was also investigated.We found no clear evidence for an effect of nematodes on the incorporation rate of fresh macroalgal detritus by amphipods, although there was some indication that the type of detritus (i.e. the green Chaetomorpha sp. vs the red Polysiphonia sp.) is important in determining the nature and extent of the relationship between nematodes and macrofaunal detritivores. Fatty acid data indicated that nematodes did not contribute significantly to the diet of amphipods when detritus was present, and there was no evidence that nematodes affected the pathway of detritus incorporation by amphipods. Amphipods incorporated Chaetomorpha sp. detritus about 10 times faster than Polysiphonia sp. detritus despite the higher C/N ratio and lower HUFA content of the former. This suggests that other factors, such as the presence of grazer deterrent compounds, are important in determining the palatability of macroalgal detritus. Amphipods fed exclusively on nematodes retained high HUFA levels but suffered high mortality. The burrowing behaviour of nematodes is suggested as the most likely factor limiting their availability to predators.Results suggest a limited interaction between amphipods and bacterivorous nematodes in detrital food webs. Further experiments are needed to test the wider applicability of these findings to different nematode and macrofaunal taxa, and for different types of detritus.     

Relative strengths of top-down and bottom-up forces in a tropical forest community

We tested integrative bottom-up and top-down trophic cascade hypotheses with manipulative experiments in a tropical wet forest, using the ant-plant Piper cenocladum and its associated arthropod community. We examined enhanced nutrients and light along with predator and herbivore exclusions as sources of variation in the relative biomass of plants, their herbivores (via rates of herbivory), and resident predaceous ants. The combined manipulations of secondary consumers, primary consumers, and plant resources allowed us to examine some of the direct and indirect effects on each trophic level and to determine the relative contributions of bottom-up and top-down cascades to the structure of the community. We found that enhanced plant resources (nutrients and light) had direct positive effects on plant biomass. However, we found no evidence of indirect (cascading through the herbivores) effects of plant biomass on predators or top predators. In contrast, ants had indirect effects on plant biomass by decreasing herbivory on the plants. This top-down cascade occurred whether or not plant resources were enriched, conditions which are expected to modify top-down forces. Received: 9 August 1998 / Accepted: 1 December 1998     

Origin and trophic importance of detritus-evidence from stable isotopes in the benthos of a small,temperate estuary

Ratios of ¹³C/¹²C and ¹⁵N/¹⁴N were measured in dissolved inorganic carbon (DIC), marginal vegetation, benthic macrodetritus (diameter > 1 mm) and selected invertebrate consumers in the Gamtoos estuary, South Africa to: (1) trace the provenance of benthic detrital deposits, and (2) determine the extent to which three abundant species of macroinvertebrates utilise this resource. DIC was strongly depleted in ¹³C with average ¹³C values (–9.5±0.5) being typical of limnetic waters. Benthic detrital particles (¹³C–24.1±0.3) originated mainly from marginal vegetation (¹³C–25.7±0.3), but their slightly elevated carbon ratio suggests additional input from ¹³C-rich sources-possibly C₄ plants cultivated on the floodplain. Populations of the fossorial ghost shrimp Callianassa kraussi, the bentho-pelagic amphipod Grandidierella lignorum and the epifaunal crab Hymenosoma robiculare together account for 96% of total benthic biomass in the upper regions of this estuary. Marked differences in trophic niches were evident among these three consumer species. Ghost shrimp (¹³C –32.5±0.3) foraged by filter-feeding on fine suspended particulate organic matter (¹³C–31.2±0.5). Amphipods (¹³C–28.0±0.6) utilised some benthic detritus but fed mainly on suspended material. Only the relatively rare crabs (¹³C–23.8±1.5) appeared to utilise benthic detrital particles to any significant extent. In the benthic consumer community of the upper Gamtoos estuary, suspension feeders make up 98% of biomass and thus clearly dominate over deposit feeders. This can be traced to the low contribution of higher plants (c. 13%) to overall carbon production, and detritus originating from macrophytes is consequently relatively unimportant in supporting invertebrate secondary production in this particular system.     

Dual isotope analyses indicate efficient processing of atmospheric nitrate by forested watersheds in the northeastern U.S.

Nitrogen from atmospheric deposition serves as the dominant source of new nitrogen to forested ecosystems in the northeastern U.S. By combining isotopic data obtained using the denitrifier method, with chemical and hydrologic measurements we determined the relative importance of sources and control mechanisms on nitrate (NO₃ ⁻) export from five forested watersheds in the Connecticut River watershed. Microbially produced NO₃ ⁻ was the dominant source (82–100%) of NO₃ ⁻ to the sampled streams as indicated by the δ¹⁵N and δ¹⁸O of NO₃ ⁻. Seasonal variations in the δ¹⁸O–NO₃ ⁻ in streamwater are controlled by shifting hydrologic and temperature affects on biotic processing, resulting in a relative increase in unprocessed NO₃ ⁻ export during winter months. Mass balance estimates find that the unprocessed atmospherically derived NO₃ ⁻ stream flux represents less than 3% of the atmospherically delivered wet NO₃ ⁻ flux to the region. This suggests that despite chronically elevated nitrogen deposition these forests are not nitrogen saturated and are retaining, removing, and reprocessing the vast majority of NO₃ ⁻ delivered to them throughout the year. These results confirm previous work within Northeastern U.S. forests and extend observations to watersheds not dominated by a snow-melt driven hydrology. In contrast to previous work, unprocessed atmospherically derived NO₃ ⁻ export is associated with the period of high recharge and low biotic activity as opposed to spring snowmelt and other large runoff events.     

Top-down and bottom-up factors in tidepool communities

Recent studies suggest that nutrient variation influences rocky intertidal community structure, however empirical evidence is rare. In the Gulf of Maine, tidepools that occur on seagull feeding roosts are potentially subjected to regular nutrient loading from seagull guano. The results of a survey conducted on Swan's Island, ME show that roost tidepools have very low macroinvertebrate and macroalgal diversity as well as very high phytoplankton biomass compared to non-roost tidepools. An experiment presented here tested basic food chain hypotheses in tidepool communities. These basic food chain models predict that in a tidepool with one trophic level (phytoplankton only), phytoplankton biomass will increase when nutrients are enriched. In contrast, these models predict that in two trophic level tidepools (phytoplankton and mussels) herbivory will prevent an increase in phytoplankton biomass when nutrients are enriched. A short term 2×2 factorially designed field experiment was used to test this basic conceptual model using herbivory by mussels and enrichment with nitrogen as the main effects. The results of this investigation are consistent with the predictions of basic food chain models, and indicate that over the short time interval of a few days, herbivory by mussels is sufficient to maintain low phytoplankton levels following enrichment with nitrogen. Experimental enrichment with phosphorus in this study had no effect on phytoplankton biomass. The results of this study suggest that periodic pulses of nitrogen into tidepools will have little effect on phytoplankton biomass when mussels are present and that longer-term chronic nitrogen influxes may be driving the patterns of community structure in tidepools occurring on roosts.     

Seasonal variation in top-down and bottom-up processes in a grassland arthropod community

Both top-down and bottom-up processes are common in terrestrial ecosystems, but how these opposing forces interact and vary over time is poorly understood. We tested the variation of these processes over seasonal time in a natural temperate zone grassland, a field site characterized by strong seasonal changes in abiotic and biotic conditions. Separate factorial experiments manipulating nutrients and cursorial spiders were performed in the wet and dry seasons. We also performed a water-addition experiment during the summer (dry season) to determine the degree of water limitation during this time. In the spring, nutrient addition increased plant growth and carnivore abundance, indicating a bottom-up control process. Among herbivores, sap-feeders were significantly enhanced while grazers significantly declined resulting in no net change in herbivore abundance. In the summer, water limitation was predominant increasing plants and all herbivores while nutrient (N) effects were non-significant. Top-down processes were present only in the spring season and only impacted the guild of grazing herbivores. These results show that bottom-up limitation is present throughout the season in this grassland, although the specific limiting resource changes as the season progresses. Bottom-up processes affected all trophic levels and many different guilds, while top-down effects were limited to a select group of herbivores and did not extend to the plant trophic level. Our results show that the relative strengths of top-down and bottom-up processes can shift over relatively short periods of time in habitats with a strong seasonal component.     

Determination of food sources and trophic position in Malaysian tropical highland streams using carbon and nitrogen stable isotopes

Stable isotope analysis has been extensively used as an effective tool in determination of trophic relationship in ecosystems. In freshwater ecosystem, aquatic invertebrates represent main component of a river food web. This study was carried out to determine potential food sources of freshwater organism together with pattern of trophic position along the river food web. In this study, rivers of Belum-Temengor Forest Complex (BTFC) has been selected as sampling site as it is a pristine area that contains high diversity and abundance of organisms and can be a benchmark for other rivers in Malaysia. Stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) were applied to estimate trophic position and food web paradigm. Analysis of stable isotopes based on organic material collected from the study area revealed that the highest δ¹³C value was reported from filamentous algae (? 22.68 ± 0.126⁰/₀₀) and the lowest δ¹³C was in allocthonous leaf packs (? 31.58 ± 0.187⁰/₀₀). Meanwhile the highest δ¹⁵N value was in fish (8.45 ± 0.177⁰/₀₀) and the lowest value of δ¹⁵N was in autochthonous aquatic macrophyte (2.00 ± 1.234⁰/₀₀). Based on the δ¹⁵N results, there are three trophic levels in the study river and it is suggested that the trophic chain begins with organic matter followed by group of insects and ends with fish (organic matter < insects < fish).     

The effect of dietary protein quality on nitrogen isotope discrimination in mammals and birds

We tested the competing hypotheses that (1) nitrogen discrimination in mammals and birds increases with dietary nitrogen concentration or decreasing C:N ratios and, therefore, discrimination will increase with trophic level as carnivores ingest more protein than herbivores and omnivores or (2) nitrogen discrimination increases as dietary protein quality decreases and, therefore, discrimination will decrease with trophic level as carnivores ingest higher quality protein than do herbivores. Discrimination factors were summarized for five major diet groupings and 21 different species of birds and mammals. Discrimination did not differ between mammals and birds and decreased as protein quality (expressed as biological value) increased with trophic level (i.e., herbivores to carnivores). Relationships between discrimination factors and dietary nitrogen concentration or C:N ratios were either the opposite of what was hypothesized or non-significant. Dietary protein quality accounted for 72% of the variation in discrimination factors across diet groupings. We concluded that protein quality established the baseline for discrimination between dietary groupings, while other variables, such as dietary protein intake relative to animal requirements, created within-group variation. We caution about the care needed in developing studies to understand variation in discrimination and subsequently applying those discrimination factors to estimate assimilated diets of wild animals.     

Movement of carbon among estuarine habitats and its assimilation by invertebrates

We measured the extent of movement of carbon and its assimilation by invertebrates among estuarine habitats by analysing carbon stable isotopes of invertebrates collected along transects crossing the boundary of two habitats. The habitats were dominated by autotrophs with distinct isotope values: (1) mudflats containing benthic microalgae (mean −22.6, SE 0.6‰) and (2) seagrass and its associated epiphytic algae (similar values, pooled mean −9.8, 0.5‰). Three species of invertebrates were analysed: a palaemonid shrimp, Macrobrachium intermedium, and two polychaete worms, Nephtys australiensis and Australonereis ehlersi. All species had a similar narrow range of isotope values (−9 to −14‰), and showed no statistically significant relationship between position along transect and isotope values. Animals were relying on carbon from seagrass meadows whether they were in seagrass or on mudflats hundreds of metres away. Particulate organic matter collected from superficial sediments along the transects had similar values to animals (mean −11.1, SE 1.3‰) and also showed no significant relationship with position. The isotope values of these relatively immobile invertebrates and the particulate detritus suggest that carbon moves from subtidal seagrass meadows to mudflats as particulate matter and is assimilated by invertebrates. This assimilation might be direct in the case of the detritivorous worm, A. ehlersi, but must be via invertebrate prey in the case of the carnivorous worm, N. australiensis and the scavenging shrimp, M. intermedium. The extent of movement of carbon among habitats, especially towards shallower habitats, is surprising since in theory, carbon is more likely to move offshore in situations such as the current study where habitats are in relatively open, unprotected waters.     

Nitrogen isotope ratios and fatty acid composition as indicators of animal diets in belowground systems

This study analyses trophic interactions between soil fungi, micro- and mesofauna in microcosm experiments. The trophic shift of ¹⁵N and fatty acids (FAs) was investigated in different food chains, which comprised either two (fungi and grazers) or three (fungi, nematodes and Collembola) levels. Contrary to the widely accepted assumption of ¹⁵N enrichment in trophic cascades the experiments revealed enrichment, depletion or no change in ¹⁵N of consumers compared to their diet. Factors responsible for this pattern were suggested to be: (1) the main metabolic pathway used for N excretion in ammonotelic nematodes to be similar or depleted in the heavier isotope, and uricotelic Collembola mostly enriched in the heavier isotope; (2) a higher shift in ¹⁵N with a high-protein diet (e.g. for predators); (3) compensation due to low-quality food altering the fractionation of ¹⁵N. Analysis of the lipid composition showed phospholipids to be generally unaffected and neutral lipids closely related to the FA pattern of the food source. Dietary routing of FAs into neutral lipids occurred, as evidenced by corresponding frequencies of FAs in host and consumer profiles. Additionally, several FAs were only detected in the grazer when present in the food source. Oleic acid showed a shift over three trophic levels, from fungi to nematodes to Collembola. The assimilation of dietary FAs resulted in a more diverse neutral lipid profile, i.e. animals higher in the food chain contained more individual FAs compared to animals lower in the food chain. The results indicate that monoenoic C18 and monoenoic C20 FAs have the potential to act as tools for the bioindication of feeding strategies in belowground systems. We suggest that primary consumers will have no or only trace amounts of monoenoic C20 acids in their neutral lipid profile, whereas consumers feeding on a eukaryote diet will show a considerably higher frequency.     

Fine-scale movement and assimilation of carbon in saltmarsh and mangrove habitat by resident animals

Despite theories of large-scale movement and assimilation of carbon in estuaries, recent evidence suggests that in some estuaries much more limited exchange occurs. We measured the fine-scale movement and assimilation of carbon by resident macroinvertebrates between adjacent saltmarsh and mangrove habitats in an Australian estuary using ¹³C analysis of animals at different distances into adjacent patches of habitat. ¹³C values of crabs (Parasesarma erythrodactyla –15.7 ± 0.1, Australoplax tridentata –14.7 ± 0.1) and slugs (Onchidina australis –16.2 ± 0.3) in saltmarsh closely matched that of the salt couch grass Sporobolus virginicus (–15.5 ± 0.1). In mangroves, ¹³C values of crabs (P. erythrodactyla –22.0 ± 0.2, A. tridentata –19.2 ± 0.3) and slugs (–19.7 ± 0.3) were enriched relative to those of mangroves (–27.9 ± 0.2) but were more similar to those of microphytobenthos (–23.7 ± 0.3). The ¹³C values of animals across the saltmarsh-mangrove interface fitted a sigmoidal curve, with a transition zone of rapidly changing values at the saltmarsh-mangrove boundary. The width of this transition indicated that the movement and assimilation of carbon is limited to between 5 and 7 m. The ¹³C values of crabs and slugs, especially those in saltmarsh habitat, clearly indicate that the movement and assimilation of carbon between adjacent saltmarsh and mangrove habitat is restricted to just a few metres, although some contribution from unmeasured sources elsewhere in the estuary is possible. Such evidence demonstrating the extent of carbon movement and assimilation by animals in estuarine habitats is useful in determining the spatial arrangement of habitats needed in marine protected areas to capture food web processes.     

The brown-world role of insectivores: Frogs reduce plant growth by suppressing detritivores in an alpine meadow

Predators of plant-suppressing herbivores have long been known to indirectly enhance plant biomass, while more recent work has revealed that predators of plant-facilitating detritivores can have the opposite effect on plant biomass. Generalist predators, such as frogs that typically facilitate plant growth in green food webs, may potentially negatively affect plant growth by consuming prey from brown food webs. In a marshy Tibetan alpine meadow, we tested the hypothesis that the locally abundant frog Rana kukunoris could negatively affect plant growth through suppression of dung-decomposing detritivores that promote plant growth by enhancing nutrient recycling. We conducted a factorial experiment (presence/absence of predators × presence/absence of dung) using replicate field enclosures over a growing season. Where dung was present, frogs significantly reduced the number of dung-feeding beetles and dung-feeding flies (including fly eggs and maggots) per dung pat, thereby decreasing dung mass loss and, indirectly, aboveground plant biomass (by 22%) surrounding dung pats. Where dung was absent, frogs did not affect plant biomass. Moreover, the number of dung beetles was positively associated with dung mass loss and soil soluble N concentration (but not total N concentrations), which in turn positively correlated with aboveground plant biomass. These results indicate that a generalist predator species standing in green food webs may play a contrasting role in brown food webs and indicate that a more nuanced appreciation of the functional role of predators in tri-trophic systems is required to accurately predict their cascading effects. Future studies must assess the relative strength of cascading effects mediated through brown and green channels in order to assess the net cascading effects of generalist predators in natural food webs.     

The diet composition of immature loggerheads: Insights on trophic niche, growth rates, and fisheries interactions

For immature animals, diet quality and composition influence expression of life history traits such as growth rates and ultimately life stage duration and age to maturity. Circumglobally distributed loggerhead turtles (Caretta caretta) exhibit a multi-decade immature stage that generally occupies neritic habitats and is characterized by slow growth and an omnivorous diet. Although adult nesting populations are geographically distinct, foraging areas for immature loggerheads show a high degree of mixing of individuals that originate from multiple nesting stocks. Furthermore, despite their generalist foraging ecology, immature loggerheads have been observed to supplement their natural diets with fish from fishery discards and/or caught in fishing gear. However, whether trophic opportunism results in variation in loggerhead growth rates within or among feeding areas has not been investigated. In Core Sound, North Carolina (NC), USA, immature loggerheads demonstrate highly variable size-specific growth rates, in contrast to other studies that report discernible somatic growth functions in immature marine turtles. To determine whether inter-individual variation in growth rates at this site was due to variation in diet composition, and specifically variation in consumption of fish, we analyzed carbon and nitrogen stable isotope ratios of loggerhead blood plasma and of tissue samples of putative loggerhead prey, as well as commercially important fish species. Our results indicated that growth rates were not related to trophic levels at which individual turtles fed, but rather probably reflected inter-individual variation in overwintering or foraging behavior (i.e. nearshore vs. offshore). Furthermore, loggerhead diets were highly diverse, and comprised mainly blue crabs and/or whelks, as well as small proportions of cannonball jellies. Fish were unimportant dietary components for loggerheads. Although loggerheads in NC do not appear to feed on fish catch or discards, immature turtles showed dietary preferences for prey items that are also valuable to or are commonly taken as bycatch in commercial fisheries (e.g. blue crabs and whelks, respectively) in the region. Thus, the status of these prey items/fishery stocks as well as trends in loggerhead populations should be monitored to mitigate potential competitive interactions between fisheries activities and loggerhead turtles.     

Quantifying the trophic base for benthic secondary production in the Nakdong River estuary of Korea using stable C and N isotopes

The overall dependence of benthic secondary production on the main primary producers at three different habitats in the Nakdong River estuarine system, Korea, was estimated. Inventories of macrobenthic invertebrate biomass were combined with multiple-isotope-mixing models to evaluate the trophic base, comparing Scirpus triqueter-dominated and Phragmites australis-dominated marshes and bare intertidal flat. The feasible contributions of four main food sources, marsh macrophytes, the microphytobenthos, and riverine and marine suspended particulate organic matter (RPOM and MPOM), to the consumer biomasses were calculated using the isotopic mixing model. After weighting the feasible contributions of food sources to each taxon by the consumer biomass, the resultant values were summed for all the consumers at each habitat to quantify the trophic base of the benthic invertebrate community. Dual-isotope-mixing model calculations verified the varying dependence on those potential food sources among the functional feeding groups. In addition, the dependence on each source of the same functional group varied between bare intertidal and salt-marsh habitats, shifting from a dominance of benthic and pelagic microalgal sources on the former habitat to a mixed food source at the latter habitat. The biomasses of the species comprising each functional group differed among habitats and sampling dates, so that each functional group made a different contribution to the whole benthic community and its basal food source. Given the calculation of the overall dependence of macrozoobenthic community on each food source, our results indicate that the microphytobenthic source dominates (nearly half) the trophic base in all the intertidal habitats of different vegetational compositions. Marsh-macrophyte-derived organic matter and RPOM served as considerable subsidies only to salt-marsh food webs, reflecting the use of the mixed food source by salt-marsh-bed consumers. Conversely, the dominance of MPOM in the total food base was equal to that of the microphytobenthos in the bare intertidal ecosystem but increased during spring−summer in the salt-marsh systems. Our results also suggest that the river discharge concentrated during the summer monsoon does not lead to any shift in trophic base for estuarine secondary production.     

Seasonal carbon isotope discrimination in a grassland community

Summary Grassland communities of arid western North America are often characterized by a seasonal increase in ambient temperature and evaporative demand and a corresponding decline in soil moisture availability. As the environment changes, particular species could respond differently, which should be reflected in a number of physiological processes. Carbon isotope discrimination varies during photosynthetic activity as a function of both stomatal aperture and the biochemistry of the fixation process, and provides an integrated measure of plant response to seasonal changes in the environment. We measured the seasonal course of carbon isotope discrimination in 42 grassland species to evaluate changes in gas exchange processes in response to these varying environmental factors. The seasonal courses were then used to identify community-wide patterns associated with life form, with phenology and with differences between grasses and forbs. Significant differences were detected in the following comparisons: (1) Carbon isotope discrimination decreased throughout the growing season; (2) perennial species discriminated less than annual species; (3) grasses discriminated less than forbs; and (4) early flowering species discriminated more than the later flowering ones. These comparisons suggested that (1) species active only during the initial, less stressful months of the growing season used water less efficiently, and (2) that physiological responses increasing the ratio of carbon fixed to water lost were common in these grassland species, and were correlated with the increase in evaporative demand and the decrease in soil moisture.     

Top-down impact through a bottom-up mechanism: the effect of limpet grazing on growth,productivity and carbon allocation of Zostera marina L. (eelgrass)

The unusual appearance of a commensal eelgrass limpet [Tectura depicta (Berry)] from southern California at high density (up to 10 shoot^–1) has coincided with the catastrophic decline of a subtidal Zostera marina L. meadow in Monterey Bay, California. Some commensal limpets graze the chloroplast-rich epidermis of eelgrass leaves, but were not known to affect seagrass growth or productivity. We evaluated the effect on eelgrass productivity of grazing by limpets maintained at natural densities (8±2 shoot^–1) in a natural light mesocosm for 45 days. Growth rates, carbon reserves, root proliferation and net photosynthesis of grazed plants were 50–80% below those of ungrazed plants, but biomass-specific respiration was unaffected. The daily period of irradiance-saturated photosynthesis (H _sat) needed to maintain positive carbon balance in grazed plants approached 13.5 h, compared with 5–6 h for ungrazed plants. The amount of carbon allocated to roots of ungrazed plants was 800% higher than for grazed plants. By grazing the chlorophyll-rich epidermis, T. depicta induced carbon limitation in eelgrass growing in an other-wise light-replete environment. Continued northward movement of T. depicta, may have significant impacts on eelgrass production and population dynamics in the northeast Pacific, even thought this limpet consumes very little plant biomass. This interaction is a dramatic example of top-down control (grazing/predation) of eelgrass productivity and survival operating via a bottom-up mechanism (photosynthesis limitation).     

Stable isotopes and trophic positions of littoral fishes from a Mediterranean marine protected area

Stable isotope analyses were employed to explore feeding and foraging habitats and trophic levels of littoral fishes in a western Mediterranean Marine Protected Area (Egadi Islands, Sicily, Italy). Carbon and nitrogen stable isotope ratios were measured in primary producers, invertebrates and fishes collected in December 2001 and January 2002. Fishes of the littoral region of the Egadi Islands had isotopic signatures that fell into a wider range for δ ¹³C (about 6‰) than for δ ¹⁵N (about 3‰). Carbon isotope ratios were consistent with a food web based on mixed sources and two trophic pathways leading to different fish species. Differences in the isotopic composition between islands were higher for benthivorous than for planktivorous fishes. The overall picture gained from this study is of a isotopic distinction between planktivorous and benthivorous fishes, resource partitioning facilitating the coexistence of similar species within the same ecosystem, and spatial variability in the isotopic signatures and trophic level of fishes. Asymmetrical analysis of variance showed that estimated trophic levels were lower in the area with the highest level of protection (Zone A) for only two out of the nine fishes analysed. As a consequence, overall spatial differences do not seem to be a consequence of protection, since in most cases trophic levels did not change significantly between zone A and zones C where professional fishing (trawling apart) is permitted, but of natural sources of variation (e.g. variability in food availability and site-specific food preferences of fishes). However, the results of this study suggest a different response at the species compared to the community level.     

The influence of productivity and width of littoral zone on the trophic position of a large-bodied omnivore

Omnivory is common in many food webs. Omnivores in different habitats can potentially change their feeding behaviour and alter their trophic position and role according to habitat conditions. Here we examine the trophic level and diet of the omnivorous signal crayfish (Pacifastacus leniusculus) in gradients of trophic status and lake size, both of which have been previously suggested to affect trophic position of predators separately or combined as productive space. We found the trophic position of omnivorous crayfish to be positively correlated with lake trophic status, but found no evidence for any influence of lake size or productive space on crayfish trophic position. The higher trophic position of crayfish in eutrophic lakes was largely caused by a shift in crayfish diet and not by an increase in trophic links in basal parts of the food web. Hence, our results support the “productivity hypothesis,” suggesting that food chains can be longer in more productive systems. Furthermore, stable isotope data indicated that larger crayfish are more predatory than smaller crayfish in lakes with wider littoral zones. Wider littoral zones promoted the development of intrapopulation differences in trophic position whereas narrow littoral zones did not. Hence, differences in habitat quality between and within lakes seem to influence the trophic positions of omnivorous crayfish. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.