Small stonefly predators affect microbenthic and meiobenthic communities in stream leaf packs

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Stepping in Elton's footprints: a general scaling model for body masses and trophic levels across ecosystems

Despite growing awareness of the significance of body-size and predator-prey body-mass ratios for the stability of ecological networks, our understanding of their distribution within ecosystems is incomplete. Here, we study the relationships between predator and prey size, body-mass ratios and predator trophic levels using body-mass estimates of 1313 predators (invertebrates, ectotherm and endotherm vertebrates) from 35 food-webs (marine, stream, lake and terrestrial). Across all ecosystem and predator types, except for streams (which appear to have a different size structure in their predator-prey interactions), we find that (1) geometric mean prey mass increases with predator mass with a power-law exponent greater than unity and (2) predator size increases with trophic level. Consistent with our theoretical derivations, we show that the quantitative nature of these relationships implies systematic decreases in predator-prey body-mass ratios with the trophic level of the predator. Thus, predators are, on an average, more similar in size to their prey at the top of food-webs than that closer to the base. These findings contradict the traditional Eltonian paradigm and have implications for our understanding of body-mass constraints on food-web topology, community dynamics and stability.     

Amphipod (Gammarus minus) responses to predators and predator impact on amphipod density

Recent theoretical work suggests that predator impact on local prey density will be the result of interactions between prey emigration responses to predators and predator consumption of prey. Whether prey increase or decrease their movement rates in response to predators will greatly influence the impact that predators have on prey density. In stream systems the type of predator, benthic versus water-column, is expected to influence whether prey increase or decrease their movement rates. Experiments were conducted to examine the response of amphipods (Gammarus minus) to benthic and water-column predators and to examine the interplay between amphipod response to predators and predator consumption of prey in determining prey density. Amphipods did not respond to nor were they consumed by the benthic predator. Thus, this predator had no impact on amphipod density. In contrast, amphipods did respond to two species of water-column predators (the predatory fish bluegills, Lepomis macrochirus, and striped shiners, Luxilus chrysocephalus) by decreasing their activity rates. This response led to similar positive effects on amphipod density at night by both species of predatory fish. However, striped shiners did not consume many amphipods, suggesting their impact on the whole amphipod “population” was zero. In contrast, bluegills consumed a significant number of amphipods, and thus had a negative impact on the amphipod “population”. These results lend support to theoretical work which suggests that prey behavioral responses to predators can mask the true impact that predators have on prey populations when experiments are conducted at small scales. Received: 21 March 1997 / Accepted: 15 December 1997     

Effects of prey dispersal on predator–prey interactions in streams

1. We studied the effect of mesh size (6 and 3 mm) on interactions between brown trout ( Salmo trutta ) and benthic invertebrates in enclosures placed in a stream in southern Sweden. We also compared how different prey exchange rates affected interactions between trout and invertebrates.
2. Trout had strong impacts on some benthic taxa, and different mesh sizes produced different patterns. Trout affected the abundance of 10 of the 21 taxa examined, six in enclosures with 3 mm mesh and six in enclosures with 6 mm mesh. The abundance of nine of the prey taxa was lower in the presence of trout, only leptocerids were more numerous in the presence of trout.
3. Our measurements of prey immigration/emigration, together with trout diet data, suggest that direct consumption by trout, rather than avoidance behaviour by prey, explains most decreases in prey abundance. There was avoidance behaviour by only two of the twenty-one prey taxa, with trout inducing emigration of the mayflies Baetis rhodani and Paraleptophlebia sp.
4. Trout indirectly increased periphyton biomass in both 3 and 6 mm enclosures. The effect of trout on periphyton was probably due to strong effects of trout on the grazer, Baetis rhodani , Heptagenia sp. and Paralepthoplebia sp.
5. Our results suggest that mesh size, through its effects on exchange rates of prey, may affect interactions between predators and prey in running waters, but that the effects of dispersal and predation on invertebrates are taxon specific.     

Influence of predator density on nonindependent effects of multiple predator species

Interactions between multiple predator species are frequent in natural communities and can have important implications for shared prey survival. Predator density may be an important component of these interactions between predator species, as the frequency of interactions between species is largely determined by species density. Here we experimentally examine the importance of predator density for interactions between predator species and subsequent impacts on prey. We show that aggressive interactions between the predatory shore crabs Carcinus maenas and Hemigrapsus sanguineus increased with predator density, yet did not increase as fast as negative interactions between conspecifics. At low density, interactions between conspecific and heterospecific predators had similar inhibitory impacts on predator function, whereas conspecific interference was greater than interference from heterospecifics at high predator density. Thus the impact of conspecific interference at high predator density was sufficient in itself that interactions with a second predator species had no additional impact on per capita predation. Spatial and temporal variability in predator density is a ubiquitous characteristic of natural systems that should be considered in studies of multiple predator species.     

Natural born killers: an invasive amphipod is predatory throughout its life-history

Introduced predators can have profound impacts on prey populations, with subsequent ramifications throughout entire ecosystems. However, studies of predator–prey interaction strengths in community and food-web analyses focus on adults or use average body sizes. This ignores ontogenetic changes, or lack thereof, in predatory capabilities over the life-histories of predators. Additionally, large individual predators might not be physically capable of consuming very small prey individuals. Both situations are important to resolve, as native prey may or may not therefore experience ontogenetic or size refuges from invasive predators. Here, we find that the freshwater amphipod invader, Gammarus pulex, is predatory throughout its development from juvenile through to adult. All size classes collected in the field had a common prey, nymphs of the mayfly Baetis rhodani, in their guts. In an experiment with predator, prey and experimental arenas scaled for body size, G. pulex juveniles and adults consumed B. rhodani in all size-matched categories. In a second experiment, the largest G. pulex individuals were able to prey on the smallest B. rhodani. Thus, the prey do not benefit from any ontogenetic or size refuge from the predator. This corroborates with the known negative population abundance relationships between this invasive predator and its native prey species. Understanding and predicting invasive predator impacts will be best served when interactions among all life-history stages of predator and prey are considered.     

Stream flow and predation effects on the spatial dynamics of benthic invertebrates

Field experiments were carried out to determine the influence of predation and prey movements on the accumulation of prey in enclosures. Experimental enclosures permitted exchange of prey with the benthos, but not of the large, predatory larvae of the caddisfly, Plectrocnemia conspersa (Curtis). Unseasonally heavy rainfalls during the experiment resulted in high flows and enabled us to examine the effects of a major, abiotic disturbance on invertebrate spatial dynamics. Prey colonization rates of cages without predators were determined in nine 24 h periods. Colonization rates increased exponentially with flow and were species-specific, depending on dispersal behaviour. Prey accumulation and predator impacts were measured in cages, with and without P. conspersa larvae, placed in the stream for 1, 2 or 3 weeks. Prey densities in cages increased with exposure time, but increases were not gradual and depended on flow regime. Flow was reduced within cages and they accumulated large numbers of invertebrates during high discharge. Analogous, naturally occurring refugia in the stream channel could be important for the recovery of lotic communities after major disturbances. Overall, prey densities were lowest in cages with predators. For fast colonizers, predation effects were detectable early in the experiment, but quickly obscured thereafter by continuous exchange of prey. For slow colonists, predation effects were detectable later, but persisted longer. Consumption rates for P. conspersa varied with prey density and flow regime. We suggest that the spatial dynamics of benthic invertebrates, especially as they are influenced by stochastic events, are important in understanding and detecting predation effects in stream communities.     

The impact of vertebrate and invertebrate predators on a stream benthic community

I assessed the impact of both vertebrate and invertebrate predators on a lotic benthic community in a 1-month-long experiment, using enclosures containing cobble/gravel bottoms, with large-mesh netting that allowed invertebrates to drift freely. Brown trout (Salmo trutta) and leeches (Erpobdella octoculata) were used as predators and four treatments were tested: a predator-free control, leeches only, trout only, and leeches and trout together. A density of 26.7 leeches/m² (20 leeches/enclosure) and 1.3 trout/m² (one trout per enclosure) was stocked into the enclosures. The total biomass of invertebrate prey was significantly lower in the trout and trout plus leech treatments than in the leech and control treatments, which were due to strong negative effects of trout on Gammarus. On the individual prey taxon level, both trout and leeches affected the abundance of Asellus , Baetis and Ephemerella, whereas the abundance of Gammarus was only affected by trout, and the abundance of Orthocladiinae and Limnephilidae was only affected by leeches. In the treatment with trout and leeches together, the abundance of Ephemerella and Baetis was higher than when trout or leeches were alone, which was probably due to predator interactions. Leeches and trout had no effects on prey immigration but did affect per capita emigration rates. Both trout and leeches indirectly increased periphyton biomass in enclosures, probably due to their strong effects on grazers. Both trout and leeches were size-selective predators, with trout selecting large prey, and leeches selecting small prey. Size-selective predation by trout and leeches affected the size structure of five commonly consumed prey taxa. Trout produced prey populations of small sizes owing to consumption of large prey as well as increased emigration out of enclosures by these large prey. Leech predation produced prey assemblages of larger size owing to consumption and increased emigration of small prey. These results suggest that in lotic habits, predatory invertebrates can be as strong interactors as vertebrate predators. Received: 23 June 1997 / Accepted: 4 May 1998     

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito, Toxorhynchites towadensis

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito larva, Toxorhynchites towadensis, were investigated in the laboratory. Survivors of T. towadensis showed different developmental patterns in relation to prey age structure. All predatory larvae in containers with only second instar prey developed into the third instar. However, in several containers with fourth instar prey, mortality of predators was observed. During the third instar, no predatory larva died, but both prey density and prey instar significantly affected the survival of predators during their fourth instar. Large prey size promoted large predator adults, and predatory larvae which grew up in small surface containers responded by developing to large sizes than those in large containers. Larval developmental time of the predators differed in each treatment. During first and second instars, faster predator development was observed in containers with small surface areas and containing young prey individuals. However, when development was enhanced by the presence of old prey individuals, no surface effect was observed. The fastest predator development was observed with prey of mixed instars and high density. This study suggests that a small surface container containing prey of mixed instars and high density is suitable for development of predators.     

Facilitation and interference among three predators affect their consumption of a stream-dwelling mayfly

1. We experimentally tested if a multiplicative risk model accurately predicted the consumption of a common mayfly at risk of predation from three predator species in New Zealand streams. Deviations between model predictions and experimental observations were interpreted as indicators of ecologically important interactions between predators. 2. The predators included a drift‐feeding fish [brown trout (T), Salmo trutta], a benthivorous fish [galaxiid (G), koaro, Galaxias brevipennis] and a benthic predatory stonefly (S; Stenoperla sp.) with Deleatidium sp. mayflies as prey. Eight treatments with all predator species combinations and a predator‐free control were used. Experiments were performed in aquaria with cobbles as predator refuges for mayflies and we measured the proportion of prey consumed after 6 h for both day and night trials. 3. Trout consumed a higher proportion of prey than other predators. For the two predator treatments we found less than expected prey consumption in the galaxiid + trout treatment (G + T) for both day and night trials, whereas a higher than expected proportion of prey was consumed during night time in the stonefly + trout (S + T) treatment. 4. The results indicate interference (G + T) and facilitation (S + T) between predators depending on predator identity and time of day. Thus, to make accurate predictions of interspecific interactions, it is necessary to consider the ecology of individual species and how differences influence the direction and magnitude of interactions.     

Who eats whom in a pool? A comparative study of prey selectivity by predatory aquatic insects

Predatory aquatic insects are a diverse group comprising top predators in small fishless water bodies. Knowledge of their diet composition is fragmentary, which hinders the understanding of mechanisms maintaining their high local diversity and of their impacts on local food web structure and dynamics. We conducted multiple-choice predation experiments using nine common species of predatory aquatic insects, including adult and larval Coleoptera, adult Heteroptera and larval Odonata, and complemented them with literature survey of similar experiments. All predators in our experiments fed selectively on the seven prey species offered, and vulnerability to predation varied strongly between the prey. The predators most often preferred dipteran larvae; previous studies further reported preferences for cladocerans. Diet overlaps between all predator pairs and predator overlaps between all prey pairs were non-zero. Modularity analysis separated all primarily nectonic predator and prey species from two groups of large and small benthic predators and their prey. These results, together with limited evidence from the literature, suggest a highly interconnected food web with several modules, in which similarly sized predators from the same microhabitat are likely to compete strongly for resources in the field (observed Pianka's diet overlap indices >0.85). Our experiments further imply that ontogenetic diet shifts are common in predatory aquatic insects, although we observed higher diet overlaps than previously reported. Hence, individuals may or may not shift between food web modules during ontogeny.     

Differences in predator composition alter the direction of structure‐mediated predation risk in macrophyte communities

Structural complexity strongly influences the outcome of predator–prey interactions in benthic marine communities affecting both prey concealment and predator hunting efficacy. How habitat structure interacts with species‐specific differences in predatory style and antipredatory strategies may therefore be critical in determining higher trophic functions. We examined the role of structural complexity in mediating predator–prey interactions across several macrophyte habitats along a gradient of structural complexity in three different bioregions: western Mediterranean Sea (WMS), eastern Indian Ocean (EIO) and northern Gulf of Mexico (NGM). Using sea urchins as model prey, we measured survival rates of small (juveniles) and medium (young adults) size classes in different habitat zones: within the macrophyte habitat, along the edge and in bare sandy spaces. At each site we also measured structural variables and predator abundance. Generalised linear models identified biomass and predatory fish abundance as the main determinants of predation intensity but the efficiency of predation was also influenced by urchin size class. Interestingly though, the direction of structure‐mediated effects on predation risk was markedly different between habitats and bioregions. In WMS and NGM, where predation by roving fish was relatively high, structure served as a critical prey refuge, particularly for juvenile urchins. In contrast, in EIO, where roving fish predation was low, predation was generally higher inside structurally complex environments where sea stars were responsible for much of the predation. Larger prey were generally less affected by predation in all habitats, probably due to the absence of large predators. Overall, our results indicate that, while the structural complexity of habitats is critical in mediating predator–prey interactions, the direction of this mediation is strongly influenced by differences in predator composition. Whether the regional pool of predators is dominated by visual roving species or chemotactic benthic predators may determine if structure dampens or enhances the influence of top–down control in marine macrophyte communities.     

The role of trout in stream food webs: integrating evidence from field surveys and experiments

1. We evaluated the effects of brown trout on boreal stream food webs using field surveys and enclosure/exclosure experiments. Experimental results were related to prey preference of uncaged trout in the same stream, as well as to a survey of macroinvertebrate densities in streams with vs. without trout. Finally, we assessed the generality of our findings by examining salmonid predation on three groups of macroinvertebrate prey (chironomid midges, epibenthic grazers, invertebrate predators) in a meta-analysis. 2. In a preliminary experiment, invertebrate predators showed a strong negative response to trout, whereas chironomids benefited from trout presence. In the main experiment, trout impact increased with prey size. Trout had the strongest effect on invertebrate predators and cased caddis larvae, whereas Baetis mayfly and chironomid larvae were unaffected. Trout impact on the largest prey seemed mainly consumptive, because prey emigration rates were low and independent of fish presence. Despite strong effects on macroinvertebrates, trout did not induce a trophic cascade on periphyton. Uncaged trout showed a strong preference for the largest prey items (predatory invertebrates and aerial prey), whereas Baetis mayflies and chironomids were avoided by trout. 3. Densities of invertebrate predators were significantly higher in troutless streams. Baetis mayflies also were less abundant in trout streams, whereas densities of chironomids were positively, although non-significantly, related to trout presence. Meta-analysis showed a strong negative impact of trout on invertebrate predators, a negative but variable impact on mobile grazers (mainly mayfly larvae) and a slightly positive impact on chironomid larvae. 4. Being size-selective predators, salmonid fishes have a strong impact on the largest prey types available, and this effect spans several domains of scale. Discrepancies between our experimental findings and those from the field survey and meta-analysis show, however, that for most lotic prey, small-scale experiments do not reflect fish impact reliably at stream-wide scales. 5. Our findings suggest that small-scale experiments will be useful only if the experimental results are evaluated carefully against natural history information about the experimental system and interacting species across a wide array of spatial scales.     

Sub-lethal effects of disturbance on a predatory net-spinning caddisfly

1. The sub-lethal effects of hydrologic disturbances on stream invertebrates are poorly understood, but integral to some models of how disturbances influence population and community dynamics. Carnivorous larvae of a net-spinning caddisfly, Plectrocnemia conspersa , have a strong predatory impact in some streams. Their silken nets, however, are vulnerable to high flow disturbance and the consequent destruction of nets could reduce predatory impacts and have life history consequences.
2. In a laboratory experiment, we manipulated the frequency of disturbances that destroyed the nets of P. conspersa , in the presence and absence of potential prey. Animals were housed individually and each trial lasted 8 days. We estimated net size, cumulative mass of silk produced, net allocation (net mass expressed as a proportion of body mass), per capita prey consumption and growth or mass loss of larvae.
3. In the absence of prey, increased disturbance frequency was accompanied by increased loss of body mass, a reduction of net size and an increase in the cumulative mass of silk produced. At the highest disturbance frequency, larvae eventually gave up producing nets. The ratio of net mass to body mass decreased with increasing disturbance, suggesting a trade-off in the allocation of resources, with a decreasing proportion of resources available for foraging. In the presence of prey, increased disturbance frequency was accompanied by a reduction in per capita prey consumption. Although foraging success offset the costs of silk production, growth rate decreased with increasing disturbance and could eventually lead to reduced body size and fecundity of adults.
4. These sub-lethal effects suggest that hydrologic disturbances could impose metabolic costs and reduce foraging efficiency of this predator. Thus, disturbances may reduce predator impact on prey populations and reduce predator population size without any direct mortality or loss of individuals.     

Differential effects of native vs. invasive predators on a common Caribbean reef fish

Predators may have consumptive (lethal) and non-consumptive (sub-lethal) effects on prey. Non-consumptive effects include altered behavior and reduced growth and fecundity. Native prey may not recognize non-native predators as a threat, and therefore may suffer pronounced effects. Additionally, non-native predators may elicit different behavioral responses from prey compared to native predators. Theory predicts that consumptive effects should be greater for non-native predators (due to prey naiveté), and non-consumptive effects should be greater for native predators (due to predator recognition). To test these hypotheses, I monitored bicolor damselfish (Stegastes partitus) in the presence of invasive predatory Pacific lionfish (Pterois spp.), a native predator (graysby, Cephalopholis cruentata), and an egg predator (bluehead wrasse, Thalassoma bifasciatum). Body size and location of lionfish and graysby were monitored on reefs in the Bahamas. Bicolor fecundity was measured as the number and size of egg-masses that individual fish laid. Bicolor fecundity was negatively correlated with lionfish density but not graysby or bluehead density. Neither predator had a detectable effect on bicolor body size, but lionfish density was negatively correlated with the size of mature adult damselfish. I observed behavioral responses of bicolors to the two piscivores, to bluehead wrasse, and to two herbivorous fishes (Acanthurus coeruleus, Scarus spp.) as non-aggressive controls. Bicolors changed behavior (feeding and aggression) in the presence of all native fishes, but not in the presence of lionfish. Thus, differential effects exist between native and non-native predators, and invasive lionfish pose a non-consumptive threat to bicolor damselfish via reduced growth and fecundity.     

Feedback between size balance and consumption strongly affects the consequences of hatching phenology in size‐dependent predator–prey interactions

In many size‐dependent predator–prey systems, hatching phenology strongly affects predator–prey interaction outcomes. Early‐hatched predators can easily consume prey when they first interact because they encounter smaller prey. However, this process by itself may be insufficient to explain all predator–prey interaction outcomes over the whole interaction period because the predator–prey size balance changes dynamically throughout their ontogeny. We hypothesized that hatching phenology influences predator–prey interactions via a feedback mechanism between the predator–prey size balance and prey consumption by predators. We experimentally tested this hypothesis in an amphibian predator–prey model system. Frog tadpoles Rana pirica were exposed to a predatory salamander larva Hynobius retardatus that had hatched 5, 12, 19 or 26 days after the frog tadpoles hatched. We investigated how the salamander hatch timing affected the dynamics of prey mortality, size changes of both predator and prey, and their subsequent life history (larval period and size at metamorphosis). The predator–prey size balance favoured earlier hatched salamanders, which just after hatching could successfully consume more frog tadpoles than later hatched salamanders. The early‐hatched salamanders grew rapidly and their accelerated growth enabled them to maintain the predator‐superior size balance; thus, they continued to exert strong predation pressure on the frog tadpoles in the subsequent period. Furthermore, frog tadpoles exposed to the early‐hatched salamanders were larger at metamorphosis and had a longer larval period than other frog tadpoles. These results suggest that feedback between the predator‐superior size balance and prey consumption is a critical mechanism that strongly affects the impacts of early hatching of predators in the short‐term population dynamics and life history of the prey. Because consumption of large nutrient‐rich prey items supports the growth of predators, a similar feedback mechanism may be common and have strong impacts on phenological shifts in size‐dependent trophic relationships.     

Are Infaunal Predators Important in Structuring Marine Soft-Bottom Communities?

The role of infaunal predators in structuring marine soft-bottomcommunities was evaluated according to these predators': 1)effects on prey density based on manipulative field experiments,2) feeding rates, 3) effects on prey distribution, 4) effectson species diversity, and 5) interactions with their prey. Estimatesof feeding rates indicate that many predatory taxa have thepotential to reduce the size of prey populations and suggestthat nemerteans are likely to have a larger impact on infaunalabundances than polychaetes. Infaunal predators have been demonstratedto have a significant effect on infaunal densities and to affectthe spatial and temporal distribution of their prey. The effectsof these predators on species diversity apparently depend onthe predator and the diversity of the system. These conclusionsmay not be applicable to all soft-bottom habitats or all groupsof infaunal predators because they are based on studies of veryfew taxa conducted almost exclusively in intertidal, unvegetated,mud habitats. Additional studies are needed on the effects ofpredation by infauna on infaunal population dynamics and onthe mechanisms of interactions between predator and prey. Furtherinvestigation will probably reveal that different groups ofinfaunal predators play different roles in structuring soft-bottomcommunities.     

Biodiversity as both a cause and consequence of resource availability: a study of reciprocal causality in a predator-prey system

1. One of the oldest questions in ecology is how species diversity in any given trophic level is related to the availability of essential resources that limit biomass (e.g. water, nutrients, light or prey). Researchers have tried to understand this relationship by focusing either on how diversity is influenced by the availability of resources, or alternatively, how resource abundance is influenced by species diversity. These contrasting perspectives have led to a seeming paradox '... is species diversity the cause or the consequence of resources that limit community biomass?' 2. Here we present results of an experiment that show it is possible for species diversity and resource density to exhibit reciprocal causal relationships in the same ecological system. Using a guild of ladybeetle predators and their aphid prey, we manipulated the number of predator species in field enclosures to examine how predator diversity impacts prey population size. At the same time, we manipulated the abundance of aphid prey in discrete habitat patches within each enclosure to determine how smaller-scale spatial variation in resource abundance affects the number of co-occurring predator species. 3. We found that the number of ladybeetle species added to enclosures had a significant impact on aphid population dynamics because interference competition among the predators reduced per capita rates of predation and, in turn, the overall efficiency of the predator guild. At the same time, spatial variation in aphid abundance among smaller habitat patches generated variation in the observed richness of ladybeetles because more species occurred in patches where predators aggregated in response to high aphid density. 4. The results of our experiment demonstrate that it is possible for species diversity to simultaneously be a cause and a consequence of resource density in the same ecological system, and they shed light on how this might occur for groups of mobile consumers that exhibit rapid responses to spatial and temporal variation in their prey.     

Is prey predation risk influenced more by increasing predator density or predator species richness in stream enclosures?

The direct lethal impacts and the indirect effects predators have on prey characteristics, such as behavior, have fitness consequences for the prey. Whether the level of predation risk that prey face in the presence of multiple predator species can be predicted from a null model that sums the risk from each predator species in isolation is unclear. In field enclosures, we tested whether the predation risk experienced by Stenonema mayfly larvae from a dragonfly larva (Boyeria vinosa) and a hellgrammite (Corydalus cornutus) together matched the predictions of the multiplicative risk model. We then compared whether any deviations from the models predictions were larger in the presence of two predator species than in the presence of an equivalent density of individuals from either predator species alone, to determine if unique effects arise for the prey in the presence of multiple predator species. We also determined if prey moved preferentially into predator-free refuge spaces or decreased their movement in the presence of predators. Stenonemas risk of predation was reduced compared to the models prediction, but no unique multiple predator species effects were present because this risk reduction was comparable in magnitude to the level exhibited in the presence of each predator species alone. The prey did not move into predator-free refuge spaces in the presence of predators in the field enclosures. Thus, these predators appear to interfere interspecifically and intraspecifically, which may facilitate the coexistence of the predators and the prey.     

Systematic deviations from linear size spectra of lake fish communities are correlated with predator–prey interactions and lake‐use intensity

Size structure of organisms at logarithmic scale (i.e. size spectrum) can often be described by a linear function with a negative slope; however, substantial deviations from linearity have often been found in natural systems. Theoretical studies suggest that greater nonlinearity in community size spectrum is associated with high predator–prey size ratios but low predator–prey abundance ratios; however, empirical evaluation of the effects of predator–prey interactions on nonlinear structures remains scarce. Here, we aim to empirically explore the pattern of the size‐specific residuals (i.e. deviations from the linear regression between the logarithmic fish abundance and the logarithmic mean fish size) by using size spectra of fish communities in 74 German lakes. We found that nonlinearity was strong in lakes with high predator–prey abundance ratios but at low predator–prey size ratios. More specifically, our results suggest that only large predators, even if occurring in low abundances, can control the density of prey fishes in a broad range of size classes in a community and thus promote linearity in the size spectrum. In turn, the lack of large predator fishes may cause high abundances of fish in intermediate size classes, resulting in nonlinear size spectra in these lakes. Moreover, these lakes were characterized by a more intense human use including high fishing pressure and high total phosphorus concentrations, which have negative impacts on the abundance of large, predatory fish. Our findings indicate that nonlinear size spectra may reflect dynamical processes potentially caused by predator–prey interactions. This opens a new perspective in the research on size spectrum, and can be relevant to further quantify the efficiency of energy transfer in aquatic food webs.