Per Nyström

THE INFLUENCE OF MULTIPLE INTRODUCED PREDATORS ON A LITTORAL POND COMMUNITY

In a replicated field experiment we studied the effects of natural densities of two exotic consumers, the predatory and herbivorous signal crayfish (Pacifastacus leniusculus) and the predatory rainbow trout (Oncorhynchus mykiss), on multiple trophic levels of a pond community. The goals were to: (1) determine the individual and combined effects of predators on macroinvertebrates, macrophytes, and periphytic algae; (2) evaluate the strength of direct and indirect interactions in a food web influenced by omnivores; and (3) evaluate the relative importance of direct and indirect predator effects on mortality and growth of a native frog species, Rana temporaria. The experiment showed that both signal crayfish and rainbow trout had strong effects on multitrophic levels of a littoral pond community, through direct consumption and indirect effects on lower trophic levels. Crayfish had weak but significant negative effects on the biomass of predatory invertebrates and greatly reduced the biomass of snails, the most abundant invertebrate grazers. Although the number of active herbivorous tadpoles tended to be higher in crayfish cages relative to control cages, the proportion of surviving froglets was lower in crayfish cages than in control cages, possibly due to crayfish predation on injured tadpoles. The size of surviving froglets did not differ from controls, but tadpoles in crayfish cages often suffered tail injuries. Macrophyte coverage decreased as a result of crayfish consumption and nonconsumptive fragmentation. However, the biomass of periphyton increased in crayfish cages relative to controls, probably due to reduced grazing from snails. In contrast, trout had strong negative effects on the biomass of both predatory invertebrates and insect grazers, whereas trout had less effect on snail biomass than did crayfish. Also, in contrast to crayfish cages, the number of active tadpoles in trout cages was lower than in controls, probably due to a combination of trout predation and trout-induced reduced tadpole activity. Trout had a strong negative impact on froglet survival, and froglets in trout cages metamorphosed at a smaller size and had reduced growth rates compared to froglets in crayfish and control cages. As with crayfish, the biomass of periphyton increased in trout cages relative to controls, which may be due to a combination of both density and trait-mediated trout effects on tadpole grazing. In treatments with multiple predators the effects of crayfish and trout on caged communities were independent, and there were few interactions. Mostly effects of combined predators reflected those in single predator cages. Our results demonstrate that noninteracting, introduced multiple predators can have strong direct and indirect effects on multiple trophic levels in pond communities. Trophic cascades may develop in aquatic food webs even with omnivores such as crayfish, and in complex habitats with trout. These strong indirect effects are mediated through both predation on important grazers (i.e., the crayfish-snail-periphyton link) and a combination of density and behavioral responses of grazers to predators (i.e., the trout-tadpole-periphyton link). When two noninteracting predators have strong but different effects on prey survival or activity, their combined effects on intermediate trophic levels reflect responses to the more dangerous predator. (Less)

Influence of an Exotic and a Native Crayfish Species on a Littoral Benthic Community

The aim of this study was to compare the effects of the introduced signal crayfish (Pacifastacus leniusculus) and the native noble crayfish (Astacus astacus) on a benthic food web. We mimicked the habitat of a pond littoral in 4.5-m 2 plastic pools stocked with natural densities of macrophytes, invertebrates and either signal crayfish, noble crayfish, or kept as crayfish free controls. After two summer months, all invertebrates and macrophytes were collected from each pool, and periphyton was sampled on one substratum exposed and two substrata not exposed to crayfish grazing. Samples for stabile isotope analysis of benthos were collected in pools with noble crayfish. 15 N ratios showed that crayfish were top consumers, and 13 C ratios indicated that they received most of their carbon from invertebrates, but less from primary producers. Crayfish did not affect the biomass of predatory invertebrates, dominated by active swimmers among Heteroptera and Coleoptera, but had a strong impact on grazers dominated by thin-shelled Lymmaea snails. Hard-shelled Bithynia snails were also reduced in numbers, but the largest of these snails were consumed less than thin-shelled Lymmaea snails. The reduced biomass of snails had an indirect positive effect on periphyton biomass on all three substrata. Crayfish grazed selectively on macrophytes and reduced the biomass of Chara, whereas Elodea was less affected. The exotic signal crayfish had, overall, a stronger impact on the biomass of macrophytes and grazers than the native noble crayfish. The results indicate that crayfish may structure food webs through consumption from many food levels. The short-term influence of crayfish on other trophic levels depends on crayfish feeding efficiency, food preferences and species-specific consumption rates.

Top-down and bottom-up processes in grassland and forested streams

The influence of predatory fish on the structure of stream food webs may be altered by the presence of forest canopy cover, and consequent differences in allochthonous inputs and primary production. Eight sites containing introduced brown trout (Salmo trutta) and eight sites that did not were sampled in the Cass region, South Island, New Zealand. For each predator category, half the sites were located in southern beech (Nothofagus) forest patches (range of canopy cover, 65–90%) and the other half were in tussock grassland. Food resources used by two dominant herbivores-detritivores were assessed using stable isotopes. 13C/12C ratios were obtained for coarse particulate organic matter (CPOM), fine particulate organic matter (FPOM), algal dominated biofilm from rocks, and larvae of Deleatidium (Ephemeroptera) and Olinga (Trichoptera). Total abundance and biomass of macroinvertebrates did not differ between streams with and without trout, but were significantly higher at grassland sites than forested sites. However, taxon richness and species composition differed substantially between trout and no-trout sites, irrespective of whether streams were located in forest or not. Trout streams typically contained more taxa, had low biomass of predatory invertebrates and large shredders, but a high proportion of consumers with cases or shells. The standing stock of CPOM was higher at forested sites, but there was less FPOM and more algae at sites with trout, regardless of the presence or absence of forest cover. The stable carbon isotope range for biofilm on rocks was broad and encompassed the narrow CPOM and FPOM ranges. At trout sites, carbon isotope ratios of Deleatidium, the most abundant invertebrate primary consumer, were closely related to biofilm values, but no relationship was found at no-trout sites where algal biomass was much lower. These results support a role for both bottom-up and top-down processes in controlling the structure of the stream communities studied, but indicate that predatory fish and forest cover had largely independent effects.

Effects of Enrichment on Simple Aquatic Food Webs

Simple models, based on Lotka‐Volterra types of interactions between predator and prey, predict that enrichment will have a destabilizing effect on populations and that equilibrium population densities will change at the top trophic level and every second level below. We experimentally tested these predictions in three aquatic food web configurations subjected to either high or low nutrient additions. The results were structured by viewing the systems as either food chains or webs and showed that trophic level biomass increased with enrichment, which contradicts food chain theory. However, within each trophic level, food web configuration affected the extent to which different functional groups responded to enrichment. By dividing trophic levels into functional groups, based on vulnerability to consumption, we were able to identify significant effects that were obscured when systems were viewed as food chains. The results support the prediction that invulnerable prey may stabilize trophic‐level dynamics by replacing other, more vulnerable prey. Furthermore, the vulnerable prey, such as Daphnia and edible algae, responded as predicted by the paradox of enrichment hypothesis; that is, variability in population density increased with enrichment. Hence, by describing ecosystems as a matrix of food web interactions, and by recognizing the interplay between interspecific competition and predation, a more complete description of the ecosystem function was obtained compared to when species were placed into distinct trophic levels.

Stable isotopes as an indicator of diet in omnivorous crayfish (Pacifastacus leniusculus): the influence of tissue, sample treatment, and season

Stable isotopes have been used to analyse food webs and (or) trace movements of animals for about 30 years. There has been some debate on the use of different tissues and treatments before isotope analysis, as well as on seasonal effects. We found different crayfish (Pacifastacus leniusculus) tissues (muscle, hepatopancreas, exoskeleton, gill, and whole body) to have different isotope values. Lipid extraction made whole-body carbon isotope values higher but had no effect on nitrogen isotope values. Acidification made whole-body isotope values lower. For crayfish, there was no seasonal or interannual variation in isotope values. In contrast to studies based on gut content analysis, we found adult crayfish to be at least as carnivorous as young-of-the-year crayfish. Earlier studies often have assumed that each food source contributes both nitrogen and carbon in equal proportions. Omnivores do not fit easily into this view. We suggest that nitrogen and carbon in an organism could come from different sources. Adopting this view for a pond food web could render crayfish both predators and detritivores as crayfish prey on nitrogen sources (other invertebrates) and consume large amounts of detritus to satisfy their carbon demand. (Less)

Crayfish predation on the common pond snail (Lymnaea stagnalis): the effect of habitat complexity and snail size on foraging efficiency

Optimal foraging theory was used to explain selective foraging by the introduced signal crayfish (Pacifastacus leniusculus) on the thin-shelled common pond snail (Lymnaea stagnalis). Crayfish predation efficiency was studied in relation to habitat complexity and snail size. In a pool experiment (area 1.3 m2) single adult crayfish were allowed to feed on four size classes of snails for one week. A pair-wise preference trial (aquarium experiment) tested if adult crayfish selectively predated on particular size classes of snail and if prey value (expressed as snail dry mass per handling time) could explain the size range of snails chosen. Crayfish preferred the smallest size classes of snails in both pool and aquaria experiments. In the pool experiment crayfish had a strong effect on snail survival. Habitat complexity did not affect overall snail survival, but resulted in reduced predation pressure on the smallest size classes of snails. Handling time and shell-thickness increased exponentially with increasing snail size, and the two smallest size classes had the highest prey values. The results suggest that crayfish can structure the abundance and size distribution of thin-shelled snails, through size-selective predation and reduction of macrophytes. The mechanisms behind the choice of snails may be based on prey value and reduced exposure time to predators and conspecifics. Crayfish effects on snail size distribution may be less pronounced in complex habitats such as macrophyte beds.

Crayfish predation on amphibian eggs and larvae

We experimentally evaluated the impact of the introduced signal crayfish (Pacifastacus leniusculus) and the native noble crayfish (Astacus astacus) on eggs and larvae of seven species of amphibians, likely to co-occur with crayfish in southern Sweden. In aquarium experiments eggs and tadpoles of all amphibian species were consumed by both crayfish species. The consumption of amphibian eggs by signal crayfish increased with temperature. The noble crayfish consumed more tadpoles than the signal crayfish, but the latter caused more sub-lethal damage to tadpoles. Tadpoles of the common toad (Bufo bufo) were sometimes killed but left uneaten by both crayfish species. In pool experiments, signal crayfish consumed more tadpoles of Hyla arborea in a less complex habitat and significantly reduced survival of Hyla tadpoles and the biomass of aquatic macrophytes.

Consumption patterns, complexity and enrichment in aquatic food chains

The interactions between consumers and prey, and their impact on biomass distribution among trophic levels, are central issues in both empirical and theoretical ecology. In a long–term experiment, where all organisms, including the top predator, were allowed to respond to environmental conditions by reproduction, we tested predictions from ‘prey–dependent’and ‘ratio–dependent’models. Prey–dependent models made correct predictions only in the presence of strong interactors in simple food chains, but failed to predict patterns in more complex situations. Processes such as omnivory, consumer excretion, and unsuitable prey–size windows (invulnerable prey) increased the complexity and created patterns resembling ratio–dependent consumption. However, whereas the prey–dependent patterns were created by the mechanisms predicted by the model, ratio–dependent patterns were not, suggesting that they may be ‘right for the wrong reason’. We show here that despite the enormous complexity of ecosystems, it is possible to identify and disentangle mechanisms responsible for observed patterns in community structure, as well as in biomass development of organisms ranging in size from bacteria to fish.

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.

Does risk of intraspecific interactions induce shifts in prey-size preference in aquatic: predators?

Abstract Interactions between foragers may seriously affect individual foraging efficiency. In a laboratory study of handling time, prey value and prey-size preference in northern pike and signal crayfish, we show that risk of intraspecific interactions between predators does not affect handling time or value of prey. However, the presence of agonistic intraspecific interactors shifts prey-size preference in these predators. Neither northern pike nor signal crayfish foraging alone show a prey-size preference, while pike foraging among conspecifics prefer small prey, and crayfish foraging in groups prefer large prey. We ascribe the different outcomes in prey preference to differences in susceptibility to interactions: northern pike under risk avoid large prey to avoid long handling times and the associated risk of interactions, while signal crayfish foraging among conspecifics may defend themselves and their prey during handling, and thus select prey to maximise investment. In addition, the value of pike prey (roach) is low for very small prey, maximises for small prey, and then decreases monotonically for larger prey, while crayfish prey (pond snail) value is low for very small prey, has a maximum at small prey, but does not decrease as much for larger prey. Therefore, a large and easily detected snail prey provides a crayfish with as much value as a small prey. We conclude that interaction risk and predator density affect prey-size preference differently in these aquatic predators, and therefore has different potential effects on prey-size structure and population and community dynamics.