Predator-Driven Nutrient Recycling in California Stream Ecosystems

Nutrient recycling by consumers in streams can influence ecosystem nutrient availability and the assemblage and growth of photoautotrophs. Stream fishes can play a large role in nutrient recycling, but contributions by other vertebrates to overall recycling rates remain poorly studied. In tributaries of the Pacific Northwest, coastal giant salamanders (Dicamptodon tenebrosus) occur at high densities alongside steelhead trout (Oncorhynchus mykiss) and are top aquatic predators. We surveyed the density and body size distributions of D. tenebrosus and O. mykiss in a California tributary stream, combined with a field study to determine mass-specific excretion rates of ammonium (N) and total dissolved phosphorus (P) for D. tenebrosus. We estimated O. mykiss excretion rates (N, P) by bioenergetics using field-collected data on the nutrient composition of O. mykiss diets from the same system. Despite lower abundance, D. tenebrosus biomass was 2.5 times higher than O. mykiss. Mass-specific excretion summed over 170 m of stream revealed that O. mykiss recycle 1.7 times more N, and 1.2 times more P than D. tenebrosus, and had a higher N:P ratio (8.7) than that of D. tenebrosus (6.0), or the two species combined (7.5). Through simulated trade-offs in biomass, we estimate that shifts from salamander biomass toward fish biomass have the potential to ease nutrient limitation in forested tributary streams. These results suggest that natural and anthropogenic heterogeneity in the relative abundance of these vertebrates and variation in the uptake rates across river networks can affect broad-scale patterns of nutrient limitation.     

Intra‐ and interspecific differences in nutrient recycling by European freshwater fish

1. We measured N and P excretion rates of 470 individuals belonging to 18 freshwater fish species widespread in Western Europe. We assessed the effect of body mass on excretion rates at both the intra‐ and interspecific levels. 2. The high variability in per capita N and P excretion rates was mainly determined by differences in body mass. The scaling coefficients of allometric relationships for both N and P excretion rates were significantly lower than 1 (mean ± SE, 0.95 ± 0.04 and 0.81 ± 0.05, respectively). 3. The slope of the allometric relationship between fish mass and nutrient excretion rate was significantly different among species. We did not detect any influence of phylogenetic conservatism on fish mass and on excretion rates. Further investigations are needed to understand the biological determinants of these differences. 4. This high intra‐ and interspecific variability in per capita excretion rates, coupled with differences in fish body mass, produce marked differences in biomass‐standardised excretion rates. These results thus indicate the necessity for further experimental and in situ investigations on the consequences of nutrient recycling by fish in freshwater ecosystems.     

Empirical models for predicting the excretion of nutrients (N and P) by aquatic metazoans: taxonomic differences in rates and element ratios

1. We developed empirical models for predicting the release of nutrients [nitrogen (N) and phosphorus (P)] by aquatic metazoans (zooplankton, mussels, benthic macroinvertebrates and fish). 2. The number of species represented in each model ranged from 9 to 74 (n = 40 – 1122), organism dry mass from 1 × 10^?5 to 8 × 10⁴ mg and water temperature from ?1.8 to 32 °C for all models. Organisms were from marine and freshwater (both lotic and lentic) environments. 3. Rates and ratios of nutrient excretion were modelled and intra‐ and intertaxon differences in excretion were examined. Rates of N and P excretion were not significantly different between marine and freshwater species within the same taxon (e.g. zooplankton). However, rates of excretion (as a function of organism dry mass and water temperature) were significantly different among different orders of zooplankton, mussels and fish. However, excretion of N was similar among different orders of benthic macroinvertebrates. 4. Detritivorous fish excreted both N and P at rates greater than all other taxa; whereas mussels excreted N and P generally at rates less than other taxa. There were no significant differences in the rate of N and P excretion between zooplankton and fish (i.e. the allometry of N and P excretion was similar between zooplankton and fish). 5. Molar N : P ratios of nutrients excreted increased with increasing organism dry mass for each group of metazoans, except for zooplankton and detritivorous fish (where N : P ratios declined with increasing organism dry mass). Molar N : P ratios in the excretions of aquatic metazoans were generally below the Redfield ratio of 16:1. 6. We examined the influence of variable abundance of zooplankton, benthic macroinvertebrates and fish on assemblage excretion rates. Rates of N and P excretion were calculated by applying our models to metazoan biomass and abundance data over seven consecutive years in two oligotrophic lakes. Rates of N and P excretion (g ha^?1 day^?1) increased linearly with increasing assemblage biomass (kg ha^?1). However, rates of N and P excretion were significantly and negatively correlated with the relative abundance of fish and positively correlated with the relative abundance of zooplankton.     

Does anthropogenic nitrogen deposition induce phosphorus limitation in herbivorous insects?

Anthropogenic nitrogen deposition has shifted many ecosystems from nitrogen (N) limitation to phosphorus (P) limitation. Although well documented in plants, no study to date has explored whether N deposition exacerbates P limitation at higher trophic levels, or focused on the effects of induced plant P limitation on trophic interactions. Insect herbivores exhibit strict N : P homeostasis, and should therefore be very sensitive to variations in plant N : P stoichiometry and prone to experiencing deposition‐induced P limitation. In the current study, we investigated the effects of N deposition and P availability on a plant‐herbivorous insect system. Using common milkweed (Asclepias syriaca) and two of its specialist herbivores, the monarch caterpillar (Danaus plexippus) and milkweed aphid (Aphis asclepiadis) as our study system, we found that experimental N deposition caused P limitation in milkweed plants, but not in either insect species. However, the mechanisms for the lack of P limitation were different for each insect species. The body tissues of A. asclepiadis always exhibited higher N : P ratios than that of the host plant, suggesting that the N demand of this species exceeds P demand, even under high N deposition levels. For D. plexippus, P addition increased the production of latex, which is an important defense negatively affecting D. plexippus growth rate. As a result, we illustrate that P limitation of herbivores is not an inevitable consequence of anthropogenic N deposition in terrestrial systems. Rather, species‐specific demands for nutrients and the defensive responses of plants combine to determine the responses of herbivores to P availability under N deposition.     

Climate change will alter amphibian‐mediated nutrient pathways: evidence from Rana temporaria tadpoles in experimental ponds

相似文献   

How<Emphasis Type="Italic"> Daphnia</Emphasis> copes with excess carbon in its food

Animals that maintain near homeostatic elemental ratios may get rid of excess ingested elements from their food in different ways. C regulation was studied in juveniles of Daphnia magna feeding on two Selenastrum capricornutum cultures contrasting in P content (400 and 80 C:P atomic ratios). Both cultures were labelled with ¹⁴C in order to measure Daphnia ingestion and assimilation rates. No significant difference in ingestion rates was observed between P-low and P-rich food, whereas the net assimilation of ¹⁴C was higher in the treatment with P-rich algae. Some Daphnia were also homogeneously labelled over 5 days on radioactive algae to estimate respiration rates and excretion rates of dissolved organic C (DOC). The respiration rate for Daphnia fed with high C:P algae (38.7% of body C day^-1) was significantly higher than for those feeding on low C:P algae (25.3% of body C day^-1). The DOC excretion rate was also higher when animals were fed on P-low algae (13.4% of body C day^-1) than on P-rich algae (5.7% of body C day^-1) . When corrected for respiratory losses, total assimilation of C did not differ significantly between treatments (around 60% of body C day^-1). Judging from these experiments, D. magna can maintain its stoichiometric balance when feeding on unbalanced diets (high C:P) primarily by disposing of excess dietary C via respiration and excretion of DOC.     

Impact of the removal of gizzard shad (Dorosoma cepedianum) on nutrient cycles in Lake Apopka,Florida

1. The St. Johns River Water Management District removed over 5.4 million kg of gizzard shad (Dorosoma cepedianum) from Lake Apopka, FL during 1993–2005, as a means of reducing lake phosphorus and phytoplankton concentrations and improving water clarity. Other steps included reduction of external nutrient inputs and operation of a treatment wetland. We measured nutrient excretion by Lake Apopka gizzard shad to quantify the nutrient effect of this biomanipulation. 2. Both N and P excretion were significantly affected by fish body mass and temperature. Larger fish had lower mass‐specific rates of excretion than smaller fish. 3. High water temperature increased P excretion to a much greater extent than N, resulting in a low N : P of nutrient excretion in midsummer. The N : P of excretion was lower than has been observed in other systems, probably because of higher water temperature. 4. Removal of gizzard shad >200 g prevented the annual release of 45 800 kg N year^?1 (3.46 kg N ha^?1 year^?1) and 7700 kg P year^?1 (0.62 kg P ha^?1 year^?1) on average. The actual impact on the P cycle varied substantially from year to year (range 7900–78 800 kg N year^?1; 1200–14 800 kg P year^?1), primarily because of fluctuations in the catch. 5. On an annual basis, the P directly removed in fish tissues was similar to that removed by the treatment wetland. The P excretion prevented by the removal of fish was approximately 20% of the reduction in external P loading achieved during 1993–2005. 6. In the short term, most of the P demand of planktonic primary producers is met through recycling of P, which greatly exceeds external P loading. Depending on population biomass, phosphorus excretion by the resident gizzard shad population was similar in magnitude to the P release by diffusive flux from the sediments.     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

PHOSPHATE UPTAKE UNDER NITRATE LIMITATION BY SCENEDESMUS SP. AND ITS ECOLOGICAL IMPLICATIONS1

Under nitrogen limitation the phosphate content of Scenedesmus sp. shows little variation regardless of growth rate and the N/P atomic ratio of the medium. P uptake therefore can be calculated as the product of P content and N-dependent growth rate. The maximum rate of P uptake in N limitation is lower by a factor of about 8 than the rate in P limitation. As reported earlier, P uptake by this alga under P limitation is described by the kinetics resembling non-competitive enzyme inhibition, with one or several intracellular P fractions as inhibitors. These fractions include surplus P (water extractable) and inorganic polyphosphate fractions A (acid soluble) and B, C, and D (acid insoluble). In N limitation, the ratios of fractions A, B, C, and D are quite different from the ratios of P limitation at comparable growth rates. The concentrations of polyphosphate fraction A in N-limited cells are much, higher than the levels in P-limited cells, and this fraction becomes more predominant at low growth rates in N limitation. This fraction, if introduced as the inhibitor into the noncompetitive scheme, explains the uptake kinetics in both N- and P-limited cells and the low maximum uptake rate in N limitation. This finding may have two significant ecological implications: (1) A nutrient imbalance which brings about changes in the internal, level or the metabolism, of fraction A would affect P uptake. (2) Nitrogen sufficiency would cause a competitive advantage in P uptake. This advantage would be shared by N₂ fixers and algae with low optimum N/P ratios. In Scenedesmus sp. P limitation switches to N limitation and vice versa when the cell N/P atomic ratio is about 30.     

Inter- and intra-annual variability in the phytoplankton community of a high mountain lake: the influence of external (atmospheric) and internal (recycled) sources of phosphorus

1. The inter‐ and intra‐annual changes in the biomass, elemental (carbon (C), nitrogen (N) and phosphorus (P)) and taxonomical composition of the phytoplankton in a high mountain lake in Spain were studied during 3 years with different physical (fluctuating hydrological regime) and chemical conditions. The importance of internal and external sources of P to the phytoplankton was estimated as the amount of P supplied via zooplankton recycling (internal) or through ice‐melting and atmospheric deposition (external). 2. Inter‐annual differences in phytoplankton biomass were associated with temperature and total dissolved phosphorus. In 1995, phytoplankton biomass was positively correlated with total dissolved phosphorus. In contrast, the negative relationship between zooplankton and seston biomass (direct predatory effects) and the positive relationship between zooplankton P excretion and phytoplankton biomass in 1997 (indirect P‐recycling effects), reinforces the primary role of zooplankton in regulating the total biomass of phytoplankton but, at the same time, encouraging its growth via P‐recycling. 3. Year‐to‐year variations in seston C : P and N : P ratios exceeded intra‐annual variations. The C : P and N : P ratios were high in 1995, indicating strong P limitation. In contrast, in 1996 and 1997, these ratios were low during ice‐out (C : P < 100 and N : P < 10) and increased markedly as the season progressed. Atmospheric P load to the lake was responsible for the decline in C : P and N : P ratios. 4. Intra‐annual variations in zooplankton stoichiometry were more pronounced than the overall differences between 1995 and 1996. Thus, the zooplankton N : P ratio ranged from 6.9 to 40.1 (mean 21.4) in 1995, and from 10.4 to 42.2 (mean 24.9) in 1996. The zooplankton N : P ratio tended to be low after ice‐out, when the zooplankton community was dominated by copepod nauplii, and high towards mid‐ and late‐season, when these were replaced by copepodites and adults. 5. In 1995, the minimum demands for P of phytoplankton were satisfied by ice‐melting, atmospheric loading and zooplankton recycling over 100%. In order of importance, atmospheric inputs (> 1000%), zooplankton recycling (9–542%), and ice‐melting processes (0.37–5.16%) satisfied the minimum demand for P of phytoplankton during 1996 and 1997. Although the effect of external forces was rather sporadic and unpredictable in comparison with biologically driven recycle processes, both may affect phytoplankton structure and elemental composition. 6. We identified three conceptual models representing the seasonal phosphorus flux among the major compartments of the pelagic zone. While ice‐melting processes dominated the nutrient flow at the thaw, biologically driven processes such as zooplankton recycling became relevant as the season and zooplankton ontogeny progressed. The stochastic nature of P inputs associated with atmospheric events can promote rapid transitional changes between a community limited by internal recycling and one regulated by external load. 7. The elemental composition of the zooplankton explains changes in phytoplankton taxonomic and elemental composition. The elemental negative balance (seston N : P < zooplankton N : P, low N : P recycled) during the thaw, would promote a community dominated by species with a high demand for P (Cryptophyceae). The shift to an elemental positive balance (seston N : P > zooplankton N : P, high N : P recycled) in mid‐season would skew the N : P ratio of the recycled nutrients, favouring dominance by chrysophytes. The return to negative balance, as a consequence of the ontogenetic increase in zooplankton N : P ratio and the external P inputs towards the end of the ice‐free season, could alleviate the limitation of P and account for the appearance of other phytoplankton classes (Chlorophyceae or Dinophyceae).     

北部湾鱼类碳、氮、磷生态化学计量特征

为探究北部湾鱼类生态化学计量特征,于2021年3月—4月采集分析了北部湾沿岸9个渔港的79种鱼类(382尾)碳(C)、氮(N)、磷(P)含量特征。结果表明北部湾鱼类C、N、P元素含量变化范围分别为33.87%—58.34%、6.31%—14.92%和0.77%—4.67%;C∶N、C∶P、N∶P的变化范围分别为3.43—9.72、19.15—173.06和5.04—33.68,其中P元素含量变化最大,导致C∶P和N∶P的变化。不同物种在科间的差异较大,科内的差异较小。不同体型和食性鱼类之间的C、N、P含量及比值具有显著差异(P<0.05)。鱼类的C∶P和N∶P与P含量呈显著负相关(R²=0.966,P<0.01;R²=0.877,P<0.01),P含量和Ca含量呈显著正相关(R²=0.919,P<0.01),P含量的变化可能与鱼类骨骼和鳞片的形成有关。总之,北部湾鱼类未保持严格的生态化学计量平衡,鱼类生态化学计量特征存在显著的种间差异,主要因鱼类不同的体型和食性差异所致。     

Nutrient recycling by fish in streams along a gradient of agricultural land use

Nutrient recycling is an essential ecosystem process provided by animals. In many aquatic systems, fish have been identified as important in ecosystem nutrient recycling; however, this importance can vary widely between systems. The factors controlling intersystem variation in animal‐mediated nutrient cycling have rarely been examined and as such it remains unclear what impact human landscape changes will have upon these processes. Here we examined rates of nutrient recycling for temperate stream fish assemblages along a gradient of agricultural land use (proportion cropland in the watershed: 1–59%). We quantified nutrient excretion rates of both ammonium–N (NH₄⁺–N) and phosphate (as soluble reactive phosphate: SRP) for fish assemblages at eight streams in southern Ontario, Canada with species‐specific excretion measurements and quantitative assemblage sampling. For both nutrients, total assemblage excretion exhibited a strong positive relationship with riparian cropland. The distance required for fish assemblages to turn over ambient nutrient pools was shorter for cropland systems, indicating that the relative importance of excreted nutrients was higher in these systems. Based on measured uptake rates of NH₄⁺–N in two streams (one higher cropland and one low cropland) and on modeled uptake rates for all streams, the proportion of ecosystem demand that can be satisfied by excretion is generally higher in the more agricultural streams. These patterns appear to be driven largely by disproportionate increases in fish assemblage biomass with increasing stream nutrient concentrations.     

Factors influencing nitrogen and phosphorus excretion rates of fish in a shallow lake

Summary 1. Fish excretion can be an important source of nutrients in aquatic ecosystems. Nitrogen (N) and phosphorus (P) excretion rates are influenced by many factors, including fish diet, fish growth rate and fish size. However, the relative influence of these and other factors on community‐level excretion rates of fish is unknown. 2. We used bioenergetics modeling to estimate excretion rates of eight fish species in a shallow, Minnesota (U.S.A.) lake over four months in both 2004 and 2005. Excretion rates of each species were summed for community‐level N and P excretion rates, as well as the N : P ratio of excretion. We then used a model‐selection approach to assess ability of independent variables to predict excretion rates, and to identify the most parsimonious model for predicting N : P excretion ratios and P and N excretion rates at the community scale. Predictive models were comprised of the independent variables water temperature and average fish density, fish size, fish growth rate, nutrient content of fish and nutrient content of fish diets at the community scale. 3. Fish density and nutrient content of fish diets (either N or P) were the most parsimonious models for predicting both N and P excretion rates, and explained 96% and 92% of the variance in N and P excretion, respectively. Moreover, fish density and nutrient models had 1400‐fold more support for predicting N and 21‐fold more support for predicting P excretion relative to models based on fish density only. Water temperature, fish size, fish growth rates and nutrient content of fish showed little influence on excretion rates, and none of our independent variables showed a strong relationship with N : P ratios of excretion. 4. Past work has focused on the importance of fish density as a driver of fish excretion rates on a volumetric basis. However, our results indicate that volumetric excretion rates at the community scale will also change substantially in response to changes in relative abundance of fish prey or shifts in relative dominance of planktivores, benthivores, or piscivores. Changes in community‐scale excretion rates will have subsequent influences on algal abundance, water clarity, and other ecosystem characteristics.     

Does the Redfield ratio infer nutrient limitation in the macroalga Spirogyra fluviatilis?

1. The cellular nutrient contents of microalgae, when growing at or approaching maximum rates, approximate the Redfield C : N : P (molar) ratio of 106 : 16 : 1. Deviations from this optimal ratio can be used to infer nutrient limitation of microalgal growth. However, this ratio may not be applicable to macroalgae, which are distinguished from microalgae by forming a thallus that is a discrete structure visible to the naked eye. The utility of the Redfield ratio to infer nutrient limitation of the growth of macroalgae was tested for Spirogyra fluviatilis in a field experiment conducted in tropical Australia. 2. The optimal cellular C : N : P ratio for S. fluvialitis was estimated by means of in situ nutrient addition. This was compared with S. fluvialitis cellular ratios determined from eight sites with a wide range of soluble N concentrations (<1–90 μg L^?1), a smaller range of soluble P concentrations (5–12 μg L^?1), and soluble molar N : P ratios of 0.11– 27. 3. Spirogyra fluviatilis had an optimal molar C : N : P ratio of 1800 : 87 : 1 which differs substantially from the Redfield ratio, and suggests that the latter ratio is not applicable to this macroalga. Concentrations of N and P in the river deviated from the optimal N : P ratio of 87 : 1, inferring nutrient limitation of growth. 4. C : P and C : N ratios of S. fluviatilis varied in accordance with general stoichiometric relationships for autotrophs under nutrient limitation of growth. Ratios of C : P and C : N increased, respectively, with increased severity of P‐ and N‐limitation. Additionally, C : P ratios increased with increased N : P ratios, whilst the C : N ratio increased with decreased N : P ratios. The C : N molar ratio however was an insensitive indicator of nutrient depletion compared with the C : P ratio. Under N‐limitation of growth, luxury amounts of P were stored by S. fluviatilis. 5. In aquatic environments where macroalgae are sufficiently abundant to be sampled, their cellular carbon, nitrogen and phosphorus stoichiometry can be used to infer nutrient limitation of growth when their optimal C : N : P ratio is known.     

Direct evaluation of macroalgal removal by herbivorous coral reef fishes

Few studies have examined the relative functional impacts of individual herbivorous fish species on coral reef ecosystem processes in the Indo-Pacific. This study assessed the potential grazing impact of individual species within an inshore herbivorous reef fish assemblage on the central Great Barrier Reef (GBR), by determining which fish species were able to remove particular macroalgal species. Transplanted multiple-choice algal assays and remote stationary underwater digital video cameras were used to quantify the impact of local herbivorous reef fish species on 12 species of macroalgae. Macroalgal removal by the fishes was rapid. Within 3 h of exposure to herbivorous reef fishes there was significant evidence of intense grazing. After 12 h of exposure, 10 of the 12 macroalgal species had decreased to less than 15% of their original mass. Chlorodesmis fastigiata (Chlorophyta) and Galaxaura sp. (Rhodophyta) showed significantly less susceptibility to herbivorous reef fish grazing than all other macroalgae, even after 24 h exposure. Six herbivorous and/or nominally herbivorous reef fish species were identified as the dominant grazers of macroalgae: Siganus doliatus, Siganus canaliculatus, Chlorurus microrhinos, Hipposcarus longiceps, Scarus rivulatus and Pomacanthus sexstriatus. The siganid S. doliatus fed heavily on Hypnea sp., while S. canaliculatus fed intensively on Sargassum sp. Variation in macroalgal susceptibility was not clearly correlated with morphological and/or chemical defenses that have been previously suggested as deterrents against herbivory. Nevertheless, the results stress the potential importance of individual herbivorous reef fish species in removing macroalgae from coral reefs.     

Soil phosphorus supply controls P nutrition strategies of beech forest ecosystems in Central Europe

Phosphorus availability may shape plant–microorganism–soil interactions in forest ecosystems. Our aim was to quantify the interactions between soil P availability and P nutrition strategies of European beech (Fagus sylvatica) forests. We assumed that plants and microorganisms of P-rich forests carry over mineral-bound P into the biogeochemical P cycle (acquiring strategy). In contrast, P-poor ecosystems establish tight P cycles to sustain their P demand (recycling strategy). We tested if this conceptual model on supply-controlled P nutrition strategies was consistent with data from five European beech forest ecosystems with different parent materials (geosequence), covering a wide range of total soil P stocks (160–900 g P m^?2; <1 m depth). We analyzed numerous soil chemical and biological properties. Especially P-rich beech ecosystems accumulated P in topsoil horizons in moderately labile forms. Forest floor turnover rates decreased with decreasing total P stocks (from 1/5 to 1/40 per year) while ratios between organic carbon and organic phosphorus (C:P_org) increased from 110 to 984 (A horizons). High proportions of fine-root biomass in forest floors seemed to favor tight P recycling. Phosphorus in fine-root biomass increased relative to microbial P with decreasing P stocks. Concomitantly, phosphodiesterase activity decreased, which might explain increasing proportions of diester-P remaining in the soil organic matter. With decreasing P supply indicator values for P acquisition decreased and those for recycling increased, implying adjustment of plant–microorganism–soil feedbacks to soil P availability. Intense recycling improves the P use efficiency of beech forests.     

Diel variation of the cellular carbon to nitrogen ratio of Chlorella autotrophica (Chlorophyta) growing in phosphorus‐ and nitrogen‐limited continuous cultures

We investigated the relationship between daily growth rates and diel variation of carbon (C) metabolism and C to nitrogen (N) ratio under P‐ and N‐limitation in the green algae Chlorella autotrophica. To do this, continuous cultures of C. autotrophica were maintained in a cyclostat culture system under 14:10 light:dark cycle over a series of P‐ and N‐limited growth rates. Cell abundance, together with cell size, as reflected by side scatter signal from flow cytometric analysis demonstrated a synchronized diel pattern with cell division occurring at night. Under either type of nutrient limitation, the cellular C:N ratio increased through the light period and decreased through the dark period over all growth rates, indicating a higher diel variation of C metabolism than that of N. Daily average cellular C:N ratios were higher at lower dilution rates under both types of nutrient limitation but cell enlargement was only observed at lower dilution rates under P‐limitation. Carbon specific growth rates during the dark period positively correlated with cellular daily growth rates (dilution rates), with net loss of C during night at the lowest growth rates under N‐limitation. Under P‐limitation, dark C specific growth rates were close to zero at low dilution rates but also exhibited an increasing trend at high dilution rates. In general, diel variations of cellular C:N were low when dark C specific growth rates were high. This result indicated that the fast growing cells performed dark C assimilation at high rates, hence diminished the uncoupling of C and N metabolism at night.     

Herbivory in omnivorous fishes: effect of plant secondary metabolites and prey stoichiometry

1. Many animals that consume freshwater macrophytes are omnivorous (i.e., they include both plant and animal matter in their diet). For invertebrate omnivorous consumers, selection of macrophyte species depends partly on the presence of secondary metabolites in plants, plant carbon/nutrient balances and/or physical structure of plants. However, little is known about the mechanisms influencing consumption of macrophytes in aquatic vertebrates. 2. For two fish species, the omnivorous rudd (Scardinius erythrophthalmus) and herbivorous grass carp (Ctenopharyngodon idella), feeding preferences were determined in three choice experiments. We tested (i) whether the presence of secondary metabolites and macrophyte stoichiometry affects macrophyte species selection by fish, (ii) the importance of macrophyte stoichiometry by manipulating the macrophytes experimentally and (iii) the rate of herbivory when the most palatable macrophyte is offered simultaneously with a common animal prey. 3. In a choice experiment with five species of submerged macrophytes (Callitriche sp., Chara globularis, Elodea nuttallii, Myriophyllum spicatum and Potamogeton pectinatus), Myriophyllum was clearly consumed least by both fishes, which strongly correlated with the highest phenolic concentration of this macrophyte. Additionally, a significant negative relationship was found between consumption and C : N ratio of the five macrophytes. The two most consumed macrophytes also had the lowest dry matter concentration (DMC). 4. In a second choice experiment, the C : N ratio of the least (Myriophyllum) and most (Potamogeton) palatable plants was manipulated by growing the macrophytes under fertilised and unfertilised conditions and subsequently feeding them to rudd. The avoidance of consumption of the chemically defended Myriophyllum by rudd was partly alleviated by the lowered C : N ratio. 5. The third choice experiment showed that both fishes preferred animal prey (the amphipod Gammarus pulex) over the most palatable macrophyte (Potamogeton) when offered simultaneously. The C : N ratio of the amphipods was about half that of the lowest C : N ratio measured in the macrophytes. Consumption by the fishes could not clearly be related to C : P or N : P ratios of prey items in any of the experiments. 6. We conclude that omnivorous fish avoid macrophytes that are chemically defended. However, when these defences are only minor, stoichiometry (C : N ratio) in combination with DMC may be a determining factor for consumption by vertebrate facultative herbivores.     

Herbivory facilitates growth of a key reef‐building Caribbean coral

The decline of reef‐building corals in conjunction with shifts to short‐lived opportunistic species has prompted concerns that Caribbean reef framework‐building capacity has substantially diminished. Restoring herbivore populations may be a potential driver of coral recovery; however, the impact of herbivores on coral calcification has been little studied. We performed an exclusion experiment to evaluate the impact of herbivory on Orbicella faveolata coral growth over 14 months. The experiment consisted of three treatments: full exclusion cages; half cage procedural controls; and uncaged control plates, each with small O. faveolata colonies. We found that herbivorous fish exclusion had a substantial impact on both macroalgal cover and coral growth. Fleshy macroalgae reached 50% cover within some exclusion cages, but were almost absent from uncaged control plates. Critically, O. faveolata calcification rates were suppressed by almost half within exclusion cages, with monthly coral growth negatively related to overgrowth by fleshy macroalgae. These findings highlight the importance of herbivorous fishes for coral growth and the detrimental impact of macroalgal proliferation in the Caribbean. Policy makers and local managers should consider measures to protect herbivorous fishes and reduce macroalgal proliferation to enable coral communities to continue to grow and function.     

Stoichiometry of nutrient recycling by vertebrates in a tropical stream: linking species identity and ecosystem processes

Ecological stoichiometry offers a framework for predicting how animal species vary in recycling nutrients, thus providing a mechanism for how animal species identity mediates ecosystem processes. Here we show that variation in the rates and ratios at which 28 vertebrate species (fish, amphibians) recycled nitrogen (N) and phosphorus (P) in a tropical stream supports stoichiometry theory. Mass-specific P excretion rate varied 10-fold among taxa and was negatively related to animal body P content. In addition, the N : P ratio excreted was negatively related to body N : P. Body mass (negatively related to excretion rates) explained additional variance in these excretion parameters. Body P content and P excretion varied much more among taxonomic families than among species within families, suggesting that familial composition may strongly influence ecosystem-wide nutrient cycling. Interspecific variation in nutrient recycling, mediated by phylogenetic constraints on stoichiometry and allometry, illustrates a strong linkage between species identity and ecosystem function.