Animals can be important in modulating ecosystem-level nutrient cycling, although their importance varies greatly among species and ecosystems. Nutrient cycling rates of individual animals represent valuable data for testing the predictions of important frameworks such as the Metabolic Theory of Ecology (MTE) and ecological stoichiometry (ES). They also represent an important set of functional traits that may reflect both environmental and phylogenetic influences. Over the past two decades, studies of animal-mediated nutrient cycling have increased dramatically, especially in aquatic ecosystems. Here we present a global compilation of aquatic animal nutrient excretion rates. The dataset includes 10,534 observations from freshwater and marine animals of N and/or P excretion rates. These observations represent 491 species, including most aquatic phyla. Coverage varies greatly among phyla and other taxonomic levels. The dataset includes information on animal body size, ambient temperature, taxonomic affiliations, and animal body N:P. This data set was used to test predictions of MTE and ES, as described in Vanni and McIntyre (2016; Ecology DOI: 10.1002/ecy.1582).

Pelagic crustacean zooplankton were collected from 336 Norwegian lakes covering a wide range of latitude, altitude, lake area, mean depth, production (as chlorophyll a), and fish community structure. Mean zooplankton species richness during the ice-free season was generally low at high latitudes and altitudes. Further, lower species richness was recorded in western lakes, possibly reflecting constraints on migration and dispersal. However, despite obvious spatial limitations, geographic boundaries were only weak predictors of mean zooplankton richness. Similarly, lake surface area did not contribute positively to mean richness such as seen in other ecosystem surveys. Rather, intrinsic factors such as primary production and fish community (planktivore) structure were identified by regression analysis as the major predictors of zooplankton diversity, while a positive correlation was observed between species richness and total zooplankton biomass. However, in spite of a large number of variables included in this study, the predictive power of multiple regression models was modest (< 50% variance explained), pointing to a major role for within-lake properties, as yet unidentified intrinsic forces, stochasticity, or dispersal as constraints on zooplankton diversity in these lakes.

Phosphorus (P) to chlorophyll ratios and zooplankton–phytoplankton (Z:P) biomass ratios were assessed in 400 temperate lakes over a gradient of phosphorus (P) and with different fish communities. Most of the lakes in this survey were oligotrophic, with a median total P of 7.3 μg P L−1. Thus, the survey provided information on food web effects during the early phase of eutrophication. There was no tendency toward a reduced yield of autotrophs per unit of P over the gradient covered in this survey. The zooplankton yield per unit of P or chlorophyll a decreased slightly with increased nutrient concentrations, and Z:P biomass ratios decreased with fish community classes, reflecting increased fish predation pressure. However, the variability in biomass ratios within a given range of P and fish class was some 100 times higher than the difference over the gradients. This finding suggests that lake-specific properties, community composition, and food quality are by far the most important determinants of biomass ratios and probably also trophic efficiency in lakes; it further suggests that these factors are superimposed on the general effect of eutrophication, at least up to 30 μg P L−1.

A survey on phytoplankton:zooplankton biomass ratios was performed in 342 Norwegian lakes, covering a wide range in lake size and productivity (total phosphorus: 3–246 μg l−1), but with most localities being oligo- to mesotrophic. Mean phytoplankton biomass was 88 μg C l−1, yet with the majority below 50 μg C l−1and a median of 25 μg C l−1. Total zooplankton biomass displayed a mean and median of 37 and 26 μg C l−1, respectively. Cladocerans were by far the dominant group, making up a median of almost 60% of total zooplankton biomass. Total zooplankton biomass as well as that of major aggregated metazoan taxa (cladocerans, calanoid copepods, cyclopoid copepods and rotifers) all showed a positive, but weak correlation with total phytoplankton biomass. These weak correlations suggest that algal biomass per se is a poor predictor of zooplankton biomass. An average phyto-:zooplankton biomass ratio (C:C) of 2.8 (SD±4.7) was found. 30% of the lakes had a phyto-:zooplankton biomass ratio below unity. While there was no correlation between the phyto-:zooplankton biomass ratio with increasing productivity in terms of P concentration, there was a higher biomass ratio in lakes with high fish predation pressure. The low ratio of phyto-:zooplankton biomass suggest major requirements from non-algal sources of C in the zooplankton diet. The need for dietary subsidizing is also supported by the fact that more than 75% of the lakes had algal biomass less than the estimated threshold for net positive growth of zooplankton, although it should be kept in mind that a high share of picoplankton would imply an underestimation of autotroph biomass in these lakes. Since the C-deficiency apparently is most pronounced in oligotrophic systems, it contradicts the view that the detritus pathways plays a predominant role in highly productive systems only, but while the source of detritus probably is mostly of autochthonous origin in eutrophic lakes, allochthonous detritus will be more important in oligotrophic systems.

Using empirical data from 466 temperate to arctic lakes covering a total phosphorus (TP) gradient of 2-1036 mug L-1, we describe how the relative contributions of resource supply, and predator control change along a nutrient gradient. We argue that (a) predator control on large-bodied zooplankton is unimodally related to TP and is highest in the most nutrient-rich and nutrient-poor lakes and generally higher in shallow than deep lakes, (b) the cascading effect of changes in predator control on phytoplankton decreases with increasing TP, and (c) these general patterns occur with significant variations-that is, the predation pressure can be low or high at all nutrient levels. A quantile regression revealed that the median share of the predator-sensitive Daphnia to the total cladoceran biomass was significantly related unimodally to TP, while the 10% and 90% percentiles approached 0 and 100%, respectively, at all TP levels. Moreover, deep lakes (more than 6 m) had a higher percentage of Daphnia than shallow (less than 6 m) lakes. The median percentage of Daphnia peaked at 0.15 mg L-1 in shallow lakes and 0.09 mg L-1 in deep lakes. The assumption that fish are responsible for the unimodality was supported by data on the abundance of potential planktivorous fish (catch net(-1) night(-1) gill nets with the different mesh sizes [CPUE]). To elucidate the potential cascading effect on phytoplankton, we examined the zooplankton phytoplankton biomass ratio. Even though this ratio was inversely related to CPUE at all TP levels, we found an overall higher ratio in oligotrophic lakes that declined toward low values (typically below 0.2) in hypertrophic lakes. These results suggest that planktivorous fish have a more limited effect on the grazing control of phytoplankton in oligotrophic lakes than in eutrophic lakes, despite similar predator control of large-bodied zooplankton. Accordingly, the phytoplankton yield, expressed as the chlorophyll a-TP ratio, did not relate to CPUE at low TP, but it increased significantly with CPUE at high TP. We conclude that the chances of implementing a successful restoration program using biomanipulation as a tool to reduce phytoplankton biomass increase progressively with increasing TP, but that success in the long term is most likely achieved at intermediate TP concentrations.

Phytoplankton carbon and particulate organic carbon (POC), nitrogen (PON), and phosphorus (POP) (POC :PON: POP) were analyzed in 109 temperate lakes covering a wide span in productivity and other key parameters. Seasonal means of total POC (four samples) ranged from 206 to 7160 µg C L −1 , with a grand mean of 960 µg C L −1 , whereas estimated phytoplankton C ranged 12 to 1,770 µg C L −1 , with a mean of 217 mg C L −1 . Sestonic C: P ratios ranged from 59 to 553 (atom : atom), with a mean of 207. The elemental contributions from phytoplankton and other sestonic compartments (mainly detritus) were analyzed with a simple regression model, in which autochthonous and allochthonous components were separated. Model‐derived estimates for N: P ratios of phytoplankton and allochthonous seston compartments were nearly equal (15.4 ± 2.5 and 16.0 ± 2.0) and were not significantly different from the Redfield N: P ratio (16). The estimated C: P ratio of allochthonous detritus was 2.7 times higher than that for phytoplankton (123 ± 15), which again was not significantly different from the Redfield C: P ratio (106). Altogether, this indicates that sestonic components of autochthonous origin should be closer to Redfield proportions in eutrophic than in oligotrophic lakes. It also indicates that major contributions of allochthonous detrital C in oligotrophic lake seston may explain deviations from the Redfield ratio and calls for caution when interpreting elemental ratios in algae versus total seston. The regression model indicates that live phytoplankton cells rarely exceed 40% of total POC, yet it suggests that a major fraction of detritus is derived from autotrophs. This close link between live and dead cells could explain why total seston apparently carries the stoichiometric and biochemical footprints from the phytoplankton. Judged from algal biomass alone, Daphnia would face severe food limitation in a majority of lakes, while if we were to include total seston, Daphnia would be above threshold food levels in all lakes. Likewise, the effect of food quality limitation related to C: P ratios will turn out differently if total seston or only the phytoplankton fraction is considered.

During the late 1980s and early 1990s, the cladoceran Limnosida frontosa invaded several lakes within its natural range in southeastern Norway. In this project, we wanted to study the types of lakes preferred by Limnosida. We also wanted to evaluate the potential competitive effects on other zooplankton species. In a survey of 65 Norwegian lakes, Limnosida showed preference for lakes of low Ca concentrations and low productivity. This is probably due to decreased competition from species with higher Ca requirements and a lower fish predation on zooplankton in these lakes compared to more productive lakes. Particle size preferences of Limnosida were studied and compared with those of the microfiltrator Daphnia magna, as there was no published information on the food preference of Limnosida.When fed monodisperse fluorescent latex beads (0.5, 1.0, 6.0 μm), Limnosida strongly selected the largest beads, while D. magna had a more nonselective feeding behaviour. Mesh sizes of Limnosida's filtering appendages were 0.4–1.2 μm depending on the animal size, and the particle selection was correlated with the filter mesh sizes. Limnosida should thus be considered a low efficiency grazer on bacteria and μ-algae. Hence, this species probably does not interfere significantly with microfiltrators like Diaphanosoma brachyurumand most daphnids. This was supported by community analysis of lakes with and without Limnosida. In general, Limnosida commonly co-occurred with a number of filter-feeding cladocerans, and we found no sign of competitive exclusion in lakes where the species has become established.

Based on data from the four Nordic countries: Denmark, Sweden, Norway and Finland, regressions have been developed relating phytoplankton abundance as chlorophyll-a, chl-a, to nutrient concentrations (as total phosphorus, TP, and total nitrogen, TN). Regressions were developed for the full data set from each country and for three ranges of trophy (expressed as TP < 20 mg · m-3, 20 mg · m-3 > TP > 200 mg · m-3, and TP > 200 mg · m-3). Our results suggest that trophic level is a stronger determinant for characterising the chl-a = f(TP) and Chl-a = g(TN) relationships than the regional localisation of the lakes. Thus, in contrast to many other pollution criteria, (e.g., acidification) equal guideline- or criteria values for phosphorus- or nitrogen- loads across regions will give equal end-point effects in lakes. Total phosphorus seems to limit phytoplankton growth when TP < 20 mg · m-3. There is no clear indication which of the nutrients, TP or TN, that limit phytoplankton growth for 20 mg · m-3 > TP > 200 mg · m-3. There are signs that neither TP nor TN are limiting factors for phytoplankton growth at TP > 200 mg · m-3.

1. Data on submerged and floating‐leafed macrophytes, phytoplankton, nutrients (N, P) and calcium were collected from twenty‐four small lakes ( 1 km2) over a wide range of latitudes in Norway. The majority of the investigated lakes were mesotrophic or eutrophic, and most of the lakes were markedly affected by diffuse and point‐source runoff from agriculture. According to their macrophyte species composition, the majority of the lakes can be classified as Potamogeton lakes or Chara lakes, or a combination of these.2. This study is consistent with the ‘two alternative stable states’ hypothesis. We observed clearwater lakes with dense macrophyte cover over a wider range of total P concentration than has been reported previously: from 30 to more than 700 mg P m–3. The clearwater state was only observed in lakes with mean depths of less than 1.9 m.3. Most clear lakes with high cover of submerged vegetation showed indications of N limitation.4. In this study nearly all the macrophyte‐dominated lakes with P concentrations above 30 mg m–3 had dense stands of Ceratophyllum demersum (L.). This indicates that Ceratophyllum may also play an important role in stabilizing and maintaining a clearwater state at high P concentrations.

A survey of 346 Norwegian lakes revealed some consistent patterns with regard to the occurrence of Holopedium gibberum versus Daphnia spp. Both biomass and frequency of Holopedium were negatively correlated with pH and Ca-concentration, and this species rarely occurred at pH > 8, or Ca-concentrations > 10 mg l(-1). On the contrary, the various Daphnia species, although showing different susceptibilities to fish predation and different preferences for lake productivity, were all positively correlated with Ca. Statistical analysis as well as laboratory experiments and biochemical considerations, suggest that Holopedium may be a superior competitor at very low Ca-concentrations, but is replaced by Daphnia at higher pH and increasing Ca-content. High pH in itself is possibly disadvantageous for Holopedium by preventing formation of the muco-polysaccaride mantle.

In general the pooling of major taxa did not provide a basis for classifying zooplankton communities in 342 large Norwegian lakes, as neither cladocerans nor calanoids varied systematically with lake productivity or fish predation pressure. At the species level, most herbivorous cladocerans and calanoids, which constituted three quarters of the metazoan zooplankton biomass, differed in their preference for lake productivity and fish community and could be distinctly grouped according to these variables by canonical correspondance analysis. The analysis pointed out one oligotrophic and one eutrophic specialist among the herbivorous cladocerans, while two of the calanoids were oligotrophic specialists. The biomasses of cladocerans, calanoids, or daphnids were poorly correlated with both lake productivity and fish predation, whereas shifts in average size and species distribution could be attributed to these variables. At low lake productivity, chemical variables such as pH and Ca, as well as the species' physiological adaptations, appear as the main determinants for the competitive advantage and relative success of herbivorous species. Fish community composition changes with increasing lake productivity, but only at very high fish predation intensity (cyprinid communities) did the effects of predation become the main determinant of the zooplankton community, superimposed on the lake productivity.

Cells of the green algaSelenastrum capricornutum were immobilized in alginate beads. The alga was able to grow inside these beads without being grazed by zooplankton. For P-limited immobilized cells, however, a lower µ m and initial slope of the Monod growth curve µ m /K s were found than for free cells.

"Habitat shift in roach (Rutilus rutilus) induced by the introduction of pikeperch (Stizostedion lucioperca)." Internationale Vereinigung für theoretische und angewandte Limnologie: Verhandlungen, 25(4), p. 2123

Changes in the fish community structure and habitat use were followed after the introduction of pikeperch (Stizostedion lucioperca) to the roach-dominated Lake Gjersjøen. Quantitative echosounding showed that the density of juvenile roach (Rutilus rutilus) was dramatically reduced in pelagic areas, from 12 000–15 000 fish/ha to 250 fish/ha, while total fish density remained unchanged in littoral areas. At the same time, the habitat segregation between different size groups of roach was altered as larger roach utilized the pelagic zone after pikeperch introduction. The loss of the pelagic refuge for juvenile roach increased the availability of juvenile roach to littoral predators, notably perch. In littoral areas, the fish community changed from one dominated by roach (> 95%) to one dominated by perch (> 50%).