Performance trade-off across a natural resource gradient

An important environmental factor determining both phytoplankton and zooplankton community composition is lake depth and thermal stratification. However, there is little information on how the interaction between zooplankton grazers and their phytoplankton food changes along an environmental gradient of lake depth. We contrasted resource availability for daphniid zooplankton populations living in two shallow, unstratified lakes and in two deep, stratified lakes using a novel growth bioassay. Stratified lakes had consistently lower resource richness than shallow unstratified lakes. To test whether resources were important in explaining differences in daphniid composition of shallow and deep lakes, we performed reciprocal transplant experiments. We raised daphniids typical of shallow (Ceriodaphnia reticulata) and deep (Daphnia dentifera) lakes in the resources from replicate shallow and deep lakes and monitored survival and reproduction. The two species exhibited a performance trade-off, measured by life table r and R ₀, across a gradient in natural resource richness. D. dentifera had higher relative fitness than C. reticulata when raised in the poorest resource environment from a deep lake. However, under richer resource conditions typical of shallow lakes, C. reticulata outperformed D. dentifera. We further created a gradient in natural resource quantity (by dilution) to test whether this trade-off in species relative fitness involved aspects of resource quality. No trade-off in species performance was evident across the dilution gradient, indicating that resource quality was important to the trade-off. We conclude that shifts in daphniid species composition along a gradient of lake depth involve an adaptive trade-off in ability to exploit rich versus poor resource quality. Received: 11 May 1998 / Accepted: 15 January 1999     

Patch use behaviour in benthic fish depends on their long-term growth prospects

Animals foraging in a heterogeneous environment may combine prior information on patch qualities and patch sample information to maximize intake rate. Prior information dictates the long-term expectations, whereas prior information in combination with patch sample information determines when to leave an individual food patch. We examined patch use behaviour of benthic feeding fish in their natural environment at different spatial scales to test if they could determine patch quality and if patch use behaviour was correlated with environmental quality. In seven lakes along a gradient of environmental quality (measured as maximum benthivore size), we made repeated measurements of giving-up density (GUD) in artificial food patches of different qualities. At the largest spatial scale, between lakes, we tested if giving-up densities revealed the long-term growth expectation of benthic fish. At the local scale of patches and micro patches we tested for the ability of benthic fish to assess patch quality, and how this ability depended on the patch exploitation levels between the different lakes. We found that GUD was positively related to maximum size of bream, suggesting that short-term behavioural decisions reflected long-term growth expectations. Benthic fish discriminated between nearby rich and poor patches, but not between rich and poor micropatches within a food patch. This suggests that the foraging scale of benthic fish lies between the patch and micro patch scale in our experiments. We conclude that patch use behaviour of benthic fish can provide a powerful measure of habitat quality that reveals how benthic fish perceive their environment.     

Grazing and detritivory in 20 stream food webs across a broad pH gradient

Acidity is a major driving variable in the ecology of fresh waters, and we sought to quantify macroecological patterns in stream food webs across a wide pH gradient. We postulated that a few generalist herbivore-detritivores would dominate the invertebrate assemblage at low pH, with more specialists grazers at high pH. We also expected a switch towards algae in the diet of all primary consumers as the pH increased. For 20 stream food webs across the British Isles, spanning pH 5.0–8.4 (the acid sites being at least partially culturally acidified), we characterised basal resources and primary consumers, using both gut contents analysis and stable isotopes to study resource use by the latter. We found considerable species turnover across the pH gradient, with generalist herbivore-detritivores dominating the primary consumer assemblage at low pH and maintaining grazing. These were joined or replaced at higher pH by a suite of specialist grazers, while many taxa that persisted across the pH gradient broadened the range of algae consumed as acidity declined and increased their ingestion of biofilm, whose nutritional quality was higher than that of coarse detritus. There was thus an increased overall reliance on algae at higher pH, both by generalist herbivore-detritivores and due to the presence of specialist grazers, although detritus was important even in non-acidic streams. Both the ability of acid-tolerant, herbivore-detritivores to exploit both autochthonous and allochthonous food and the low nutritional value of basal resources might render chemically recovering systems resistant to invasion by the specialist grazers and help explain the sluggish ecological recovery of fresh waters whose water chemistry has ameliorated.     

Responses of tropical cladocerans to a gradient of resource quality

1. The response of three tropical cladocerans to a gradient of resource quality was compared in a series of growth bioassays using seston collected from five lakes of different depth and trophic structure in Michigan, U.S.A. To assess the food quality in terms of digestibility, assimilation experiments were performed with ³²P‐labelled seston from the same lakes. Animals were also analysed for P‐content in their tissues at the end of these assays. 2. In general, assimilation efficiency was higher when animals fed on seston from shallow compared to deep lakes, and was significantly correlated with growth rates, suggesting that shallow lakes have the best food resources in terms of digestibility and P availability. 3. Results also showed that all cladoceran species responded similarly to the resource gradient, with lower growth rates in deep lakes and higher growth rates in shallow lakes, although the strength of response (sensitivity) was different among the species tested. 4. The cladoceran Moina micrura was the most sensitive species, and also displayed the highest P‐content and maximal growth rate, a pattern consistent with the growth rate hypothesis. 5. However, seston C : P ratio and growth rates in the different resources did not correlate with the animals’ P‐contents, showing an uncoupling between RNA‐phosphorus demands for growth and seston food quality. 6. In conclusion, our results support the idea that digestion resistance in algae is a major constraint to cladocerans in natural plankton communities.     

Food quantity and quality shapes reproductive strategies of Daphnia

In freshwater environments, one of the challenges aquatic grazers face are periods of suboptimal food quantity and quality. In a life table experiment, the effects of food quantity (a gradient of algae concentration) and quality (a diet of cyanobacteria) on the life histories and resource allocation strategy in Daphnia magna were tested. Growth‐related traits were similarly affected under different food regimes while the reproductive strategies differed in animals exposed to low food quantity and quality. The per‐clutch investment (clutch volume) did not differ between Daphnia fed with cyanobacteria and underfed mothers, but resources were differently allocated; underfed mothers increased their per‐offspring investment by producing fewer, but larger eggs, whereas cyanobacteria‐fed mothers invested in a greater number of eggs of smaller size. I argue that both strategies of resource allocation (number vs. size of eggs) may be adaptive under the given food regime. The results of the study show that the cyanobacteria diet‐driven fitness losses are comparable to losses caused by food quantity, which is only slightly above the growth capability threshold for Daphnia.     

Resource Defence or Exploded Lek? – A Question of Perspective

It becomes increasingly obvious that animal mating systems cannot be classified into distinct categories, but transitions between mating system classes are continuous. Positioning a certain mating system at this continuum is often not straightforward, however. Depending on which characteristic is considered, a mating system may end up at very different positions on this gradient. Here, we explore the potential conflict between mating system classifications that may arise when they are based on different criteria by investigating the mating system of the cichlid fish Simochromis pleurospilus in which males defend small patches of homogeneously distributed food resources (turf algae) vigorously against food competitors, but they allow specific females to use them. We hypothesized that male defence may generate high‐quality feeding patches serving to attract females, and hence male territoriality constitutes a form of courtship. Our field data show that males selectively allow approximately one‐third of the visiting females to feed on their territory and that females preferentially feed in male territories and usually sample several territories successively. As males protect food patches against other algae grazers and guard females from harassment by food competitors, females gain nutritional benefits from visiting male territories. Hence, males appear to generate essential resources for females, which is the key feature of resource‐defence mating systems, although the distributions of resources and of males and females are characteristics of an exploded lek.     

‘House and garden’: larval galleries enhance resource availability for a sedentary caddisfly

1. Sedentary grazers can be numerous in fresh waters, despite the constraints on resource availability and the increased predation risk inherent in this lifestyle. The retreats of sedentary grazers have been assumed to provide protection to the resident (a ‘house’), but also may provide additional fertilised food for the grazer (i.e. a ‘garden’). If retreats function as a garden, then they should (i) contain a higher quality and/or quantity of food than the alternative food source. Furthermore, the proportion of retreat‐derived carbon and nitrogen assimilated by the resident should be (ii) related to overall resource availability (more when resources are limited) or (iii) perhaps also to larval density. Alternatively, if retreats provide a less risky food source, then (iv) assimilation of material from the retreat is likely to be greater under conditions in which the risk of emerging from the retreat is high. 2. We tested these four hypotheses for the common and widespread gallery‐building grazing caddisfly Tinodes waeneri. Resource availability, larval density and biomass, and exposure were measured for populations from six lakes of differing productivity in August, October and January. 3. Galleries always contained more algal food than the surrounding epilithon, suggesting that gardening is effective. Furthermore, gallery chlorophyll a content in August, and the disparity in food quality (assessed from the C : N ratio) between gallery and epilithon (quality higher in the former) in October were positively related to the proportion of larval biomass that was derived from the gallery. Larval density and wave exposure parameters were not related to larval assimilation of gallery material. 4. Galleries that are fertilised by the occupant provide more, and sometimes also better quality, food (in terms of the C : N ratio) than is otherwise available. Thus, the gallery plays a substantial role in larval nutrition, and this role is greater at key times of food shortage.     

Gardening by the psychomyiid caddisfly Tinodes waeneri: evidence from stable isotopes

Sedentary species face a trade-off between the benefits of exploiting food close to their homes and the cost of defending it. In aquatic systems, it has been suggested that some sedentary grazers can increase the range of circumstances under which they are at an advantage over mobile grazers by enhancing food resources within their feeding territories through ‘gardening’. We examined this for the retreat-building sedentary larvae of the caddis Tinodes waeneri, which are often dominant in the littoral of lakes. We hypothesised that T. waeneri gardens by fertilising its retreat (a fixed ‘gallery’ on which algae and other microorganisms grow), and that gardening would be more important in lower productivity lakes. We tested this by analysing the carbon and nitrogen stable isotope ratios of larvae, their galleries and the general background epilithon, collected from rocks in the littoral zones of six lakes spread across a natural nutrient gradient. We found evidence of nutrient recycling within the Tinodes gallery community in all lakes. Galleries were ¹⁵N-depleted compared to the epilithon, suggesting that algae on galleries preferentially assimilated ¹⁴N from larval excretions that were themselves ¹⁵N-depleted relative to the larval food source. Mixing model results indicate that galleries formed an important larval carbon and nitrogen source, with mean gallery dietary contributions of over 40% in at least one sample period in all lakes studied. Gallery contributions were greater between April and October than in January and, contrary to our initial hypothesis, greater in the more productive lakes of those surveyed. Nevertheless, T. waeneri galleries do act as a fertilised garden. ‘Gardening’ appears to be widespread in this species, and may affect productivity and patterns of nitrogen retention within the stony littoral of lakes.     

Some like it deep: Intraspecific niche segregation in ruffe (Gymnocephalus cernua)

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Interactions between metazoans,autotrophs, mixotrophs and bacterioplankton in nutrient‐depleted high DOC environments: a long‐term experiment

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Relationships between oral morphology and feeding style in the Ungulata: a phylogenetically controlled evaluation

In ungulates it is argued that specialization in the consumption of a particular type of food (feeding style) is reflected in morphological adaptations of the organs involved in the selection, processing and digestion of food. We analysed the differences in size and morphology of some oral traits that have been functionally related to food-selection ability (muzzle width, incisor-arcade shape, incisor shape), prehension of food (incisor protrusion), food comminution (molar occlusal surface area, hypsodonty (high-crowned molars)) and intake rate (incisor breadth) between ungulate species with different feeding styles (browser, mixed feeder, grazer). Grazers were characterized by large-body-size species. After controlling only for body mass, we found that grazers had wider muzzles and incisors, more-protruding incisors and more-bulky and higher-crowned molars than did mixed feeders and browsers. When the analyses took into account both body mass and phylogeny, only body mass and two out of the three hypsodonty indexes used remained significantly different between feeding styles. Browsers were smaller, on average, than mixed feeders and grazers, whilst grazers and mixed feeders did not differ in size. Also, browsers had shorter and less-bulky molars than did mixed feeders and grazers; the latter two feeding styles did not differ from each other in any of the hypsodonty indexes. We conclude that the adaptation to different dietary types in most of the oral traits studied is subsumed by the effects of body mass and the sharing of common ancestors. We hypothesize that differences in the ability to exploit different food resources primarily result from differences in body mass between species, and also discuss why hypsodonty characterizes feeding styles.     

Macrophytes in tropical shallow lakes: An important food item to benthic entomofauna or an underused resource?

Although macrophytes are known to increase benthic diversity in lakes, the importance of this resource as food for the insects living at the bottom of these ecosystems are still poorly understood. This study assessed the diets of benthic Chironomidae and Campsurus (Ephemeroptera) in two environments: a lake with macrophytes (M+) and another without macrophytes (M?). We expected a differential use of food resources in M+, where plant tissue is particularly important for the aquatic insects’ diet. The diet of 734 individuals from 16 taxa were analyzed. Contrary to expectations, benthic insects consumed low amounts of plant tissue. This finding led us to investigate whether the presence of macrophytes in lakes would indirectly contribute to the benthic insects’ feeding, as more food resources were explored in M+ and a spatial variation of resources intake was observed in this lake, in contrast to the homogeneous feeding in M?. We highlight that macrophytes were responsible for the organic matter build‐up in the sediment, especially at the lake region dominated by these plants, and contributed to increase the deposition of high‐quality amorphous organic matter, which favored taxa in M+ that fed exclusively on this item. The lower diversity of food items exploited in M?, and the Tanypus alga‐based diet in this lake, indicates the low quality of organic resources in its sediment. Although macrophytes were indirectly beneficial for benthic insects’ feeding, we found that this is not an attractive resource for prompt ingestion by most benthic taxa.     

The influence of moss on grazers in high‐altitude streams: food,refuge or both?

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Quality matters: resource quality for hosts and the timing of epidemics

Epidemiologists increasingly realize that species interactions (e.g. selective predation) can determine when epidemics start and end. We hypothesize here that resource quality can also strongly influence disease dynamics: epidemics can be inhibited when resource quality for hosts is too poor and too good. In three lakes, resource quality for the zooplankton host ( Daphnia dentifera ) was poor when fungal epidemics ( Metschnikowia bicuspidata ) commenced and increased as epidemics waned. Experiments using variation in algal food showed that resource quality had conflicting effects on underlying epidemiology: high-quality food induced large production of infective propagules (spores) and high birth rate but also reduced transmission. A model then illustrated how these underlying correlations can inhibit the start of epidemics (when spore production/birth rate are too low) but also catalyse their end (when transmission becomes too low). This resource quality mechanism is likely to interface with other ones controlling disease dynamics and warrants closer evaluation.     

Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Food quality of heterotrophic bacteria for Daphnia magna: evidence for a limitation by sterols

The quality of heterotrophic bacteria as food for metazoan grazers has been investigated poorly. We conducted growth experiments with juvenile Daphnia magna feeding on different strains of heterotrophic bacteria that represent typical pelagic bacteria of five phylogenetically distinct groups. The bacterial food suspensions were supplemented with cholesterol and/or the polyunsaturated fatty acid eicosapentaenoic acid (EPA), two essential nutrients that are either absent or scarcely represented in bacteria. Our data imply that the selected heterotrophic bacteria are of poor food quality for D. magna, which was indicated either by very low somatic growth rates or by high mortality. However, with four out of six bacterial strains tested, the somatic growth rates increased significantly upon supplementation with cholesterol, which shows that the lack of sterols in bacteria is a major food quality constraint. We did not find clear evidence for a limitation by EPA on bacterial diets within our growth experiments. High mortality was observed when D. magna was fed with Hydrogenophaga sp. or Pseudomonas sp., which indicates that these two bacterial strains are toxic to D. magna. Our findings highlight the limitations of bacteria as a carbon source for Daphnia and point to a so far underestimated diversity of interactions between grazers and its bacterial food.     

Food quantity and quality regulation of trophic transfer between primary producers and a keystone grazer (Daphnia) in pelagic freshwater food webs

The transfer of energy and nutrients from plants to animals is a key process in all ecosystems. In lakes, inefficient transfer of primary producer derived energy can result in low animal growth rates, accumulation of nuisance phytoplankton blooms and dissipation of energy from the ecosystem. Most research on carbon transfer efficiency in pelagic food webs has focused on either food quantity or food quality, with the latter considered separately as either elemental stoichiometry or biochemical composition. The natural occurrence and magnitude of these types of growth limitations and their combined effects on Daphnia , a keystone grazer in pelagic freshwater ecosystems, are largely unknown. Our empirical models predict that the strength and nature of food quantity and quality limitation varies greatly with lake trophic state (total phosphorus, TP) and that Daphnia growth rates and thus energy and nutrient transfer efficiency are highest in lakes with intermediate trophic status (TP 10–25 μg l⁻¹). We predict that food availability place the greatest constraint on Daphnia growth in nutrient poor lakes (TP≤4 μg l⁻¹). Phosphorus limitation of Daphnia growth increased with decreasing TP, but the overall effect was never predicted to be the dominant constraining factor. Eicosapentaenoic acid (EPA, 20:5ω3) limitation was predicted to occur in both nutrient poor and nutrient rich lakes and placed the primary constraint on food quality in the most productive lakes. Two contrasting EPA-models gave different results on the magnitude of EPA-limitation, implying that additional food quality factors decrease Daphnia growth at high TP. In conclusion, the model predicts that Daphnia growth should peak in mesotrophic lakes, food quantity will place the greatest constraint on growth in oligotrophic lakes and EPA will primarily limit growth in eutrophic lakes.     

Are zooplankton food resources poor in the vegetated littoral zone of shallow lakes?

1. The distribution of zooplankton in shallow lakes is negatively related to macrophyte density. However, the abundance of their food along density gradients of macrophytes is unknown. A common but untested assumption is that food quantity and quality for pelagic zooplankton is poor in the littoral zone owing to the deleterious influence of macrophytes on phytoplankton. 2. We tested this assumption with a combination of a field survey and laboratory experiments. We collected seston samples from the littoral and pelagic zones of four shallow temperate lakes and related food quantity (phytoplankton biovolume) and quality to macrophyte abundance (per cent volume infested). Seston food quality was assessed in three ways: N/C and P/C ratios, polyunsaturated fatty acid content and phytoplankton community composition. In the laboratory, we measured the growth and reproduction of Daphnia pulicaria on diets consisting of seston from the littoral and pelagic zones in one lake. 3. In our four study lakes, food quantity was not significantly influenced by macrophyte abundance, and food quality was generally high. Laboratory experiments showed increased juvenile growth, but no significant change in D. pulicaria reproduction, when feeding on littoral resources compared to pelagic resources. 4. Our results suggest that there is no nutritional cost to pelagic zooplankton inhabiting the littoral zone. Therefore, it is likely that other factors (e.g. predation, abiotic factors) are involved in determining zooplankton habitat use.     

Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers

1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. 2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. 3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 × 5 km cells across repeated censuses. 4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. 5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.     

Benthic and pelagic food resources for zooplankton in shallow high-latitude lakes and ponds

1. Shallow lakes and ponds are a major component of the northern landscape and often contain a high zooplankton biomass despite clear waters that are poor in phytoplankton. 2. In this study we quantified zooplankton food sources and feeding rates in the shallow waters of two contrasting high‐latitude biomes: subarctic forest tundra (Kuujjuarapik, Quebec) and high arctic polar desert (Resolute, Nunavut). Five substrate types were tested (beads, bacteria, picophytoplankton, filamentous plankton and microbial mats). Special attention was given to the role of benthos, a component that is usually poorly integrated into models of aquatic foodwebs. 3. Consistent with observations elsewhere in the circumpolar region, high concentrations of adult macrozooplankton occurred in all sites (up to 17 100 crustaceans m^?3) while phytoplankton concentrations and primary productivity were low. The communities were composed of multiple species, including Daphnia middendorfiana, Hesperodiaptomus arcticus, Leptodiaptomus minutus, Artemiopsis stefanssoni and Branchinecta paludosa. 4. Detritus made 89–98% of the planktonic resource pool and bacteria contributed the highest biomass (up to 29 mg C m^?3) of the planktonic food particles available to zooplankton. Benthic resources were dominated by microbial mats that grew in nutrient‐rich conditions at the base of the ponds and which dominated overall ecosystem biomass and productivity. 5. All species were flexible in their feeding but there were large, order of magnitude differences in clearance rates among taxa. These differences likely resulted from different grazing strategies among cladocerans, copepods and fairy shrimps, and possibly also from adaptation to specific food types and size ranges that occur locally in these waters. 6. The subarctic cladocerans Ceriodaphnia quadrangula and D. middendorfiana, and the arctic fairy shrimp B. paludosa were observed to graze directly on the microbial mats and the feeding experiments confirmed their assimilation of benthic substrates. The other zooplankton species showed a more pelagic feeding mode but were capable of using microbial mat filaments, thus may be indirectly linked to benthic processes via resuspension. 7. Our study indicates that the classical aquatic food web in which phytoplankton provide the sole production base for grazers does not apply to northern shallow lakes and ponds. Instead, microbial mats increase the physical complexity of these high latitude ecosystems and likely play a role in sustaining their high zooplankton biomass.