Distribution and abundance of littoral benthic fauna in Canadian prairie saline lakes

The littoral benthos of 18 lakes in Alberta and Saskatchewan ranging in salinity from 3 to 126 (g1^–1 TDS) were investigated twice, in the spring and in the summer of 1986. Multiple Ekman dredge samples were taken at water depths of about 0.5, 1.0 and 2 metres in each transect. Two to three transects were used in each lake according to its estimated limnological diversity for a total of 114 stations. A total of 76 species was present varying from 29–31 species in the three lakes of lowest salinity (means of 3.1–5.55) to only 2 species in lakes exceeding 100. Species richness decreased rapidly in salinities greater than 15.Biomass maximum mean of 10.91 g m^–2 dry weight (maximum 63.0 g m^–2) occurred in culturally eutrophic Humboldt Lake (3.1) but one third as great in other low salinity lakes. However, biomass again increased to about 4.5 gm^–2 in two lakes of 15 As the salinity increased still further biomass declined steadily until a minimum of 0.0212 g m^–2 was recorded in most saline Aroma Lake (mean 119). Summer biomass (11 lakes) was greater than spring biomass (4 lakes) because some groups such as amphipods, corixids and ostracods became more abundant in summer. Wet weight biomass averaged 15.8 of dry weight biomass.Seasonality (spring or summer), sediment texture and organic matter content, water depth, pH, salinity (TDS) and the presence of aquatic plants ( plant cover) were considered in the matrix involving species dry weight biomass at each of 117 stations. TWINSPAN classification of the samples yielded a dendrogram with 18 indicator species. Successive dichotomies divided these indicator species into four main lake groups based on salinity, i.e., Group I: 3–10 (Gammarus, Glyptotendipes I, Chironomus cf. plumosus), Group II: 10–38%. (Hyalella, Enallagma,Bezzia), Group III: 38–63 (Hygrotus salinarius, Cricotopus ornatus), Group IV: >63 (Dolichopodidae, Ephydra hians). Each of these main groups was subdivided into smaller groups of lakes based on factors such as pH, seasonality (spring or summer species dominance), organic matter and plant cover. Depth of samples played no apparent role.     

Vertical heterogeneity of a forest floor invertebrate food web as indicated by stable-isotope analysis

Diverse populations of invertebrates constitute the food web in detritus layers of a forest floor. Heterogeneity in trophic interactions within such a species-rich community food web may affect the dynamic properties of biological communities such as stability. To examine the vertical heterogeneity in trophic interactions among invertebrates in litter and humus layers, we studied differences in species composition and variations in carbon and nitrogen stable-isotope ratios (δ¹³C and δ¹⁵N) using community-wide metrics of the forest floors of temperate broadleaf forests in Japan. The species composition differed between the two layers, and the invertebrates in the litter layer were generally larger than those in the humus layer, suggesting that these layers harbored separate food webs based on different basal resources. However, the δ¹³C of invertebrates, an indicator of differences in the basal resources of community food webs, did not provide evidence for separate food webs between layers even though plant-derived organic matter showed differences in stable-isotope ratios according to decomposition state. The minimum δ¹⁵N of invertebrates also did not differ between layers, suggesting sharing of food by detritivores from the two layers at lower trophic levels. The maximum and range of δ¹⁵N were greater in the humus layer, suggesting more trophic transfers (probably involving microorganisms) than in the litter layer and providing circumstantial evidence for weak trophic interactions between layers at higher trophic levels. Thus, the invertebrate community food web was not clearly compartmentalized between the detrital layers but still showed a conspicuous spatial (vertical) heterogeneity in trophic interactions.     

The trophic state of Lake Ladoga as indicated by late summer phytoplankton

As a part of the joint Russian-Finnish evaluation of human impact on Lake Ladoga, we studied the phytoplankton of the lake in order to find biological indicators for eutrophication. A second aim of the investigation was intercalibration of sampling and phytoplankton counting techniques between the Russian and Finnish laboratories. Phytoplankton samples were collected from 27 sampling stations in the lake and from the rivers Volkhov and Neva in 9–13 August 1993. In surface water samples the phytoplankton fresh weight biomass varied in the range 218–3575 mg m^–33. Highest biomass values were encountered in Sortavala Bay, and lowest ones in the western central part of the lake. Phytoplankton species composition varied considerably in the lake; blue-green and green algae predominated near-shore areas and Cryptophyceae in the offshore stations. Canonical correspondance analysis revealed close grouping of eutrophy indicating communities, dominated mainly by greens and blue-greens, in the most nutrient-rich parts of Lake Ladoga, the Volkhov and Svir Bays. Samples from the vicinity of the inflows of Vuoksi and Burnaya Rivers and off Pitkaranta formed a separate group, dominated by diatoms, most of which were typical to mesotrophic or eutrophy lakes. As judged by phytoplankton biomass values and chlorophyll a concentrations, Lake Ladoga may generally be classified as mesotrophic. Eutrophicated areas are found in the northern archipelago of the lake and in the areas influenced by large rivers.     

The influence of environmental factors on seasonal changes in bacterial cell volume in two prairie saline lakes

Bacterial biovolumes of hypertrophic Humboldt Lake (total dissolved solids = 3.3 g liter^-1; 6 m deep) and oligotrophic Redberry Lake (total dissolved solids = 20.9 g liter^-1; 17 m deep), Saskatchewan, were measured concurrently with a variety of environmental variables to identify the major factors correlated with volume changes. There was no difference (P > 0.05) in mean bacterial volume between Redberry Lake (0.084 ± 0.034 m³ SD) and Humboldt Lake (0.083 ± 0.021 m³ SD). Statistical analyses suggested there were marked differences in the factors associated with the pronounced seasonality of bacterial cell volumes in these two lakes. Variance in bacterial volume in the epilimnion of Redberry Lake was best explained by a multivariate regression model which included ciliate abundance and chlorophyll concentration (r ² = 0.96). The model accounting for changes in hypolimnetic bacterial volume included ciliate numbers and primary production (r ² = 0.94), of the measured variables. Bacterial volume in Humboldt Lake was most highly correlated with primary production (r ² = 0.59). Bacterial production (estimated as the rate of thymidine incorporation into DNA) and growth (thymidine incorporation rate normalized to cell numbers) were not correlated to cell volume, with the exception of cocci volume in Humboldt Lake. Offprint requests to: R.D. Robarts.     

Patterns of seasonal phytoplankton distribution in prairie saline lakes of the northern Great Plains (U.S.A.)

Seasonal changes in freshwater phytoplankton communities have been extensively studied, but key drivers of phytoplankton in saline lakes are currently not well understood. Comparative lake studies of 19 prairie saline lakes in the northern Great Plains (USA) were conducted in spring and summer of 2004, with data gathered for a suite of limnological parameters. Nutrient enrichment assays for natural phytoplankton assemblages were also performed in spring and summer of 2006. Canonical correspondence analysis of 2004 data showed salinity (logCl), nitrogen, and phosphorus (N:P ratios) to be the main drivers of phytoplankton distribution in the spring, and phosphorus (C:P ratios), iron (logTFe), and nitrogen (logTN) as important factors in the summer. Despite major differences in nutrient limitation patterns (P-limitation in freshwater systems, N-limitation in saline systems), seasonal patterns of phytoplankton phyla changes in these saline lakes were similar to those of freshwater systems. Dominance shifted from diatoms in the spring to cyanobacteria in the summer. Nutrient enrichment assays (control, +Fe, +N, +P, +N+P) in 2006 indicated that nutrient limitation is generally more consistent within lakes than for individual taxa across systems, with widespread nitrogen and secondary phosphorus limitation. Understanding phytoplankton community structure provides insight into the overall ecology of saline lakes, and will assist in the future conservation and management of these valuable and climatically-sensitive systems.     

Genetic diversity of small eukaryotes in lakes differing by their trophic status

Small eukaryotes, cells with a diameter of less than 5 mum, are fundamental components of lacustrine planktonic systems. In this study, small-eukaryote diversity was determined by sequencing cloned 18S rRNA genes in three libraries from lakes of differing trophic status in the Massif Central, France: the oligotrophic Lake Godivelle, the oligomesotrophic Lake Pavin, and the eutrophic Lake Aydat. This analysis shows that the least diversified library was in the eutrophic lake (12 operational taxonomic units [OTUs]) and the most diversified was in the oligomesotrophic lake (26 OTUs). Certain groups were present in at least two ecosystems, while the others were specific to one lake on the sampling date. Cryptophyta, Chrysophyceae, and the strictly heterotrophic eukaryotes, Ciliophora and fungi, were identified in the three libraries. Among the small eukaryotes found only in two lakes, Choanoflagellida and environmental sequences (LKM11) were not detected in the eutrophic system whereas Cercozoa were confined to the oligomesotrophic and eutrophic lakes. Three OTUs, linked to the Perkinsozoa, were detected only in the Aydat library, where they represented 60% of the clones of the library. Chlorophyta and Haptophyta lineages were represented by a single clone and were present only in Godivelle and Pavin, respectively. Of the 127 clones studied, classical pigmented organisms (autotrophs and mixotrophs) represented only a low proportion regardless of the library's origin. This study shows that the small-eukaryote community composition may differ as a function of trophic status; certain lineages could be detected only in a single ecosystem.     

In lakes but not in minds: stakeholder knowledge of invasive species in prairie lakes

Humans are key vectors in the spread and establishment of aquatic invasive species (AIS), and human behavior can exacerbate or help prevent further spread of non-native species. Therefore, stakeholders’ knowledge is critical to preventing establishment of AIS. However, stakeholders’ AIS knowledge in prairie lakes remains poorly understood. We used a survey questionnaire in Saskatchewan, Canada, to assess the state of AIS knowledge, identify predictors of knowledge, and optimize management strategies. Statistical analyses of the responses of 440 participants indicated a generally low level of AIS knowledge, suggesting low communication success. Respondents were generally more aware of non-native fishes than plants. Of concern was the observation of substantial knowledge gaps regarding non-native mussels and important preventative behaviors that may have devastating ecological, social, and economic consequences if left unaddressed. Better understanding of AIS issues was significantly associated with several trans-situational (age, sex and education), situational (recreational purpose and using multiple lakes), and lake-related knowledge (awareness of eutrophication) predictors. Exploitation of these predictors is recommended to improve effectiveness of outreach and communication efforts. Specifically, we propose that management strategies focus on improving communications by streamlining outreach messages, targeting low-knowledge groups (e.g., swimmers, cabin owners), and expanding education campaigns.     

Deepwater sediments and trophic conditions in Florida lakes

Sediment cores were taken from near maximum depth in 15 Florida lakes representing a wide range of trophic conditions. Chemical analyses of surface sediments showed Al, Fe, and Ca to be the most abundant elements in all samples, and the ratio of Al to Ca to be smaller for eutrophic lakes. Sediment organic matter increased with trophic state, as did the degree to which it was enriched in nitrogen. Corresponding sediment C/N ratios decreased with increasing lake trophic state and showed significant negative correlation with chlorophylla, total N, and total P in the water column. Concentrations of sedimentary chlorophyll derivatives showed some relation to trophic state but differences in basin morphometry hinder its use as an inter-lake index of chlorophyll production.     

The trophic state of shallow lakes in The Netherlands

A study has been carried out on the development of an ecological assessment method for shallow lakes in The Netherlands. Analyses of eutrophication characteristics of 93 lakes with, in total, 127 sampling localities gave insight into some of the steering variables, such as total-P, total-N, chlorophyll-a and transparency. On the basis of phytoplankton species composition, three main groups of lakes could be distinguished. The first group, characterized by persistent filamentous Cyanobacteria, had the highest summer means of chlorophyll-a and total-P. The second group was characterized by non-persistent filamentous Cyanobacteria, occurring in low abundancies, and at lower chlorophyll-a and total-P concentrations. The third group of lakes was characterized by the absence or very low abundance of the filamentous Cyanobacteria. As a result of these analyses, criteria for the abatement of eutrophication in shallow lakes could be refined.     

The fauna in the upper stony littoral of Danish lakes: macroinvertebrates as trophic indicators

1. The macroinvertebrate fauna living on stones in the exposed stony littorals of thirty-nine Danish lakes were examined by multivariate numerical methods. The data were derived from 125 semi-quantitative samples and a species list of 126 taxa. The mean number of individuals per sample was 960, and among the most common taxa were Asellus aquaticus, Gammarus, Oulimnius, Tinodes, Cricotopus and Dicrotendipes. 2. The total number of species and fourteen individual taxa were positively correlated to mean depth of the lakes and eleven taxa were correlated to the total phosphorus concentration. The Shannon diversity was negatively correlated to the chlorophyll a concentration ([Chl a]). 3. Community patterns were examined by detrended correspondence analysis (DCA), and the relationship between species data and selected environmental variables was analysed by canonical correspondence analysis (CCA). Mean lake depth was found to be the strongest environmental variable in explaining the species data. The [Chl a] and Secchi depth also explained significant variation in the distribution of the stony littoral invertebrates. Wind fetch and relative exposure did not explain any variation in the faunal composition among sites. 4. The abilities of the macroinvertebrates to predict the lake trophic state, expressed as log ([Chl a]), were explored by means of weighted averaging (WA) regression and calibration. Two tolerance-weighted WA models using inverse and classical regression for deshrinking are presented. The models were assessed by the root mean square error (RMSE) of prediction, using bootstrapping as cross validation, and by the correlation between observed and inferred log ([Chl a]). The model using inverse deshrinking had a RMSE_boot = 0.41 and r² = 0.63. By using classical regression, the predictability in the ends of the gradient was improved but the RMSE increased: RMSE_boot = 0.46. 5. Although the factors determining faunal distribution patterns in the Danish lowland lakes were highly multivariate and difficult to disentangle, it seems reasonable to use the WA estimated species optima and tolerances to [Chl a] in a bio-assessment model.     

Size‐based interactions across trophic levels in food webs of shallow Mediterranean lakes

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Shifts in trophic interactions with forest type in soil generalist predators as indicated by complementary analyses of fatty acids and stable isotopes

Human impact on structure and functioning of ecosystems is rapidly increasing. Virtually all European forests are managed with major implications for diversity and structure of food webs. Centipedes (Chilopoda: Lithobiidae) are abundant arthropod predators in European temperate forest soils with a generalistic feeding behaviour. However, little is known on the variability in the prey spectrum of centipedes with land use and the responsible factors. Combining fatty acid (FA) analysis, which allows determination of the relative contribution of different prey to predator nutrition, and stable isotope analysis, providing insight into the trophic structure of decomposer food webs, we investigated variations in trophic niches of two dominant centipede species, Lithobius mutabilis and Lithobius crassipes, in differently aged beech and spruce forests. FA composition of the two centipede species differed significantly with bacterial marker FAs being more abundant in L. crassipes as compared to L. mutabilis. Differences were most pronounced in spruce as compared to beech forests. The results suggest that dense needle litter in coniferous forests may restrict prey availability to the larger L. mutabilis and confine foraging to the litter surface whereas the smaller L. crassipes is able to also exploit prey of deeper litter layers. Lithobius crassipes was significantly more enriched in ¹⁵N and ¹³C compared to L. mutabilis suggesting that, compared to L. mutabilis, the smaller L. crassipes occupies higher trophic levels and relies more on root derived carbon. The results indicate that trophic niches of centipedes vary in a species specific way between forest types with body size and habitat structure being major determinants of the variations in the prey spectrum. Combining techniques for delineating predator–prey interactions allowed insights into variations in trophic interrelationships and their driving forces in temperate forest soil food webs.     

Heterogeneity in shortgrass prairie vegetation: the role of playa lakes

Abstract. We examined the role of playa lakes in promoting regional heterogeneity on the southern High Plains. The goals of this paper were to: (1) describe vegetation types and zonation patterns within playas, (2) evaluate patterns of species distribution and abundance within and among playas, and (3) assess patterns of heterogeneity within and among playas on a portion of the southern High Plains. Perennial grasses were the most abundant species in playa vegetation. Playa vegetation exhibited distinct and repeatable vegetation zones at the majority of sites, but the number of distinct zones varied from site to site. Agropyron smithii, Buchloë dactyloides, and Panicum obtusum were the most important species of playa interior zones, and Bouteloua gracilis, Buchloë dactyloides, and Schedonnardus paniculatus were important upland species outside of playas. Species distribution and abundance were positively correlated at 38 of 40 sites. The distributions of species occurrences among sites were unimodal both locally and regionally. The degree of heterogeneity varied from playa to playa. Local heterogeneity within playas was found to increase regional heterogeneity; therefore, playas increase both local and regional heterogeneity of vegetation. Long-term monitoring will be necessary to understand the spatial and temporal response of vegetation within and among playas to stochastic climatic factors on the southern High Plains of North America.     

Photochemical mineralization of dissolved organic nitrogen to ammonia in prairie lakes

We examined the rate of photoammonification in 16 lakes from Saskatchewan, Canada. Lakes were selected to encompass a broad range in dissolved organic nitrogen (DON) (269–1,435 μg l⁻¹). Lake filtrate (<0.2 μm) was exposed to artificial solar radiation for 4 h. Rates of photoammonification were significant in 7 of the 16 study lakes. Ammonia (NH₃) concentrations increased 0.84–2.85 μg l⁻¹ over control values. This is a 4–92% increase in NH₃ concentration and a conversion of 0.18–0.3% of the DON pool to NH₃. We developed an empirical model to predict photoammonification rates across aquatic ecosystems. Photoammonification rates and ancillary parameters (i.e., pH, DOC and DON concentrations, DOC:DON ratios, and a350:DOC) were obtained from published studies to expand our dataset for model development. Model selection was conducted with Akaike’s Information Criterion (AIC). DON concentration and pH were selected as model predictors by AIC. Our model explains 49% of the variance in photoammonification rate across a diverse set of aquatic systems. This model may be useful in estimating photoammonification rates in other aquatic systems.     

Lability of organic carbon in lakes of different trophic status

1. We used first‐order kinetic parameters of biological oxygen demand (BOD), the constant of aerobic decomposition (k) and the asymptotic value of BOD (BOD_ult), to characterise the lability of organic carbon pools in six lakes of different trophic state: L. Naroch, L. Miastro and L. Batorino (Belarus), L. Kinneret (Israel), L. Ladoga (Russia) and L. Mendota (U.S.A.). The relative contributions of labile and refractory organic carbon fractions to the pool of total organic carbon (TOC) in these lakes were quantified. We also determined the amounts of labile organic carbon within the dissolved and particulate TOC pools in the three Belarus lakes. 2. Mean annual chlorophyll concentrations (used as a proxy for lake trophic state) ranged from 2.3 to 50.6 μg L⁻¹, labile organic carbon (OC_L = 0.3BOD_ult) from 0.75 to 2.95 mg C L⁻¹ and k from 0.044 to 0.14 day⁻¹. 3. Our data showed that there were greater concentrations of OC_L but lower k values in more productive lakes. 4. In all cases, the DOC fraction dominated the TOC pool. OC_L was a minor component of the TOC pool averaging about 20%, irrespective of lake trophic state. 5. In all the lakes, most (c. 85%) of the DOC pool was refractory, corresponding with published data based on measurements of bacterial production and DOC depletion. In contrast, a larger fraction (27–55%) of the particulate organic carbon (POC) pool was labile. The relative amount of POC in the TOC pool tended to increase with increasing lake productivity. 6. Long‐term BOD incubations can be valuable in quantifying the rates of breakdown of the combined particulate and dissolved organic carbon pools and in characterising the relative proportions of the labile and recalcitrant fractions of these pools. If verified from a larger number of lakes our results could have important general implications.     

Flagellate nutritional versatility as a key to survival in two contrasting Antarctic saline lakes

1. Seasonal patterns of grazing and photosynthesis were investigated in two saline Antarctic lakes (Highway and Ace) in the Vestfold Hills (68°S). The phototrophic nanoflagellate (PNAN) community was dominated by Pyramimonas gelidicola and two morphological forms of a cryptophyte species that occurred throughout the year. Both species were mixotrophic on bacteria, and in Highway Lake they also exploited dissolved organic carbon as determined by the uptake of fluorescently labelled dextrans. 2. Clearance rates ranged between 0.02 and 0.21 nL h^?1 cell^?1 in Ace Lake and 0.004–1.05 nL h^?1 cell^?1 in Highway Lake. On occasion cryptophyte grazing equalled that of the heterotrophic nanoflagellates (HNAN). 3. Photosynthetic rates showed similar trends in both lakes, but there were differences in chlorophyll a specific rates and photosynthetic efficiency, probably related to the meromictic characteristic of Ace Lake. Primary production was measurable in winter and peaked in summer following the maxima of mixotroph grazing. 4. The HNAN community of Highway Lake achieved clearance rates of 0.02–1.80 nL h^?1 cell^?1 and removing between 50 and 693 ng bacterial carbon L^?1 day^?1, with highest impact in winter when HNAN were most abundant. The HNAN also ingested fluorescently labelled dextrans showing a preference for 4 and 500 kDa molecules. The more diverse HNAN community of Ace Lake had lower clearance rates (0.04–0.37 nL h^?1 cell^?1) and exerted a lower grazing pressure on bacterioplankton. In Highway Lake, where the HNAN community was dominated by the choanoflagellate Diaphanoeca grandis, there was a significant correlation between mean cell volume and clearance rate. 5. The major feature was that the microbial plankton functioned throughout the year by employing nutritional versatility.     

Trophic structure and major trophic links in conventional versus organic farming systems as indicated by carbon stable isotope ratios of fatty acids

Using bulk tissue and fatty acid ¹³C analysis we investigated major trophic pathways from soil microorganisms to microbial consumers to predators in conventional versus organic farming systems planted for the first time with maize. Organic farming led to an increase in microbial biomass in particular that of fungi as indicated by phospholipid fatty acids (PLFAs). Microbial PLFAs reflected the conversion from C₃ to C₄ plants by a shift in δ¹³C of 2‰, whereas the isotopic signal in fatty acids (FAs) of Collembola was much more pronounced. In the euedaphic Protaphorura fimata the δ¹³C values in maize fields exceeded that in C₃ (soybean) fields by up to 10‰, indicating a close relationship between diet and vegetation cover. In the epedaphic Orchesella villosaδ¹³C values shifted by 4‰, suggesting a wider food spectrum including carbon of former C₃ crop residues. Differences in δ¹³C of corresponding FAs in consumers and resources were assessed to assign food web links. P. fimata was suggested as root and fungal feeder in soybean fields, fungal feeder in conventional and leaf consumer in organically managed maize fields. O. villosa likely fed on root and bacteria under soybean, and bacteria and fungi under maize. Comparison of δ¹³C values in FAs of the cursorial spider Pardosaagrestis and O. villosa implied the latter as important prey species in soybean fields. In contrast, the web‐building spider Mangora acalypha showed no predator–prey relationship with Collembola. The determination of δ¹³C values in trophic biomarker FAs allowed detailed insight into the structure of the decomposer food web and identified diet‐shifts in both consumers at the base of the food web and in top predators in organic versus conventional agricultural systems. The results indicate changes in major trophic links and therefore carbon flux through the food web by conversion of conventional into organic farming systems.     

Aquatic macrophytes in saline lakes of the Canadian prairies

Vascular macrophyte species richness decreases with increasing salinity. Only three species of submerged plants (Potamogeton pectinatus, Ruppia maritima, R. occidentalis) tolerate hypersaline waters (>50 g l^-1, total of ionic constituents). Eight emergent species occur in more saline habitats but only five (Scirpus maritimus var. paludosus, Distichlisstricta, Puccinellia nuttalliana, Scirpus americanus, Triglochin maritima) occur commonly over a range of saline lakes into the hypersaline category. Usually, species tolerant of high salinities are found over the entire saline spectrum and even extend into subsaline waters (<3 g l^-1) and thrive there. A major increase in the number of species occurs below 5 g l^-1. As the water recedes plants such as Salicornia rubra, Suaeda calceoliformes, Hordeum jubatum and Sonchus arvensis invade.Submerged angiosperm distribution is controlled by total ion concentration and substrate texture plays no apparent role. Although angiosperms normally grow in all kinds of substrates, they occupy coarse substrates in Wakaw lake because suitable fine substrates are densely colonized by charophytes. In this lake light limited growth occurs to a depth of 5% of surface light. Light was not limiting in Redberry Lake but angiosperm growth was limited to the upper 8 m (10% or more of surface light). Thermal stratification and depth (pressure) were probably limiting istead. In meromictic Waldsea Lake the depth of the chemocline (6 m, 5% surface light) delimits angiosperm growth.     

Predictive power of food web models based on body size decreases with trophic complexity

Food web models parameterised using body size show promise to predict trophic interaction strengths (IS) and abundance dynamics. However, this remains to be rigorously tested in food webs beyond simple trophic modules, where indirect and intraguild interactions could be important and driven by traits other than body size. We systematically varied predator body size, guild composition and richness in microcosm insect webs and compared experimental outcomes with predictions of IS from models with allometrically scaled parameters. Body size was a strong predictor of IS in simple modules (r² = 0.92), but with increasing complexity the predictive power decreased, with model IS being consistently overestimated. We quantify the strength of observed trophic interaction modifications, partition this into density‐mediated vs. behaviour‐mediated indirect effects and show that model shortcomings in predicting IS is related to the size of behaviour‐mediated effects. Our findings encourage development of dynamical food web models explicitly including and exploring indirect mechanisms.