SUMMARY 1. Streams and their adjacent riparian zones are closely linked by reciprocal flows of invertebrate prey. We review characteristics of these prey subsidies and their strong direct and indirect effects on consumers and recipient food webs. 2. Fluxes of terrestrial invertebrates to streams can provide up to half the annual energy budget for drift-feeding fishes such as salmonids, despite the fact that input occurs principally in summer. Inputs appear highest from closed-canopy riparian zones with deciduous vegetation and vary markedly with invertebrate phenology and weather. Two field experiments that manipulated this prey subsidy showed that it affected both foraging and local abundance of stream fishes. 3. Emergence of adult insects from streams can constitute a substantial export of benthic production to riparian consumers such as birds, bats, lizards, and spiders, and contributes 25–100% of the energy or carbon to such species. Emergence typically peaks in early summer in the temperate zone, but also provides a low-level flux from autumn to spring in ice-free streams. This flux varies with in-stream productivity, and declines exponentially with distance from the stream edge. Some predators aggregate near streams and forage on these prey during periods of peak emergence, whereas others rely on the lower subsidy from autumn through spring when terrestrial prey are scarce. Several field experiments that manipulated this subsidy showed that it affected the short-term behaviour, growth, and abundance of terrestrial consumers. 4. Reciprocal prey subsidies also have important indirect effects on both stream and riparian food webs. Theory predicts that allochthonous prey should increase density of subsidised predators, thereby increasing predation on in situ prey and causing a negative indirect effect via apparent competition. However, short-term experiments have produced either positive or negative indirect effects. These contrasting results may be due to characteristics of the subsidies and individual consumers, but could also result from differences in experimental designs. 5. New study approaches are needed to better determine the direct and indirect effects of reciprocal prey subsidies. Experiments coupled with comparative research will be required to measure their effects on individual consumer fitness and population demographics. Future work should investigate whether reciprocal prey fluxes stabilise linked stream–riparian ecosystems, explore how landscape context affects the magnitude and importance of subsidies, and determine how impacts of human disturbance can propagate between streams and riparian zones via these trophic linkages. Study of these

Tangled webs: reciprocal flows of invertebrate prey link streams and riparian zones

Cannibalism among generalist predators has implications for the dynamics of terrestrial food webs. Spiders are common, ubiquitous arthropod generalist predators in most natural and managed terrestrial ecosystems. Thus, the relationship of spider cannibalism to food limitation, competition, and population regulation has direct bearing on basic ecological theory and applications such as biological control. This review first briefly treats the different types of spider cannibalism and then focuses in more depth on evidence relating cannibalism to population dynamics and food web interactions to address the following questions: Is cannibalism in spiders a foraging strategy that helps to overcome the effects of a limited supply of calories and/or nutrients? Does cannibalism in spiders reduce competition for prey? Is cannibalism a significant density-dependent factor in spider population dynamics? Does cannibalism dampen spider-initiated trophic cascades?

Cannibalism, food limitation, intraspecific competition, and the regulation of spider populations

1. Riverine systems consist of a mosaic of patches and habitats linked by diverse processes and supporting highly complex communities. Invertebrates show a high taxonomic and functional diversity in riverine systems and are in several ways important components of these systems. Their distribution patterns, movements and effects on ecological flows, testify to their importance in various landscape ecological processes. This paper reviews the invertebrate literature with respect to patterns and processes in the riverine landscape.

2. The distribution of invertebrates in riverine habitats is governed by a number of factors that typically act at different scales. Hence, the local community structure can be seen as the result of a continuous sorting process through environmental filters ranging from regional or catchment-wide processes, involving speciation, geological history and climate, to the small-scale characteristics of individual patches, such as local predation risk, substratum porosity and current velocity.

3. Dispersal is an important process driving invertebrate distribution, linking different ecological systems across boundaries. Dispersal occurs within the aquatic habitat as well as into the terrestrial surrounding, and also over land to other waterbodies. New genetic techniques have contributed significantly to the understanding of aquatic invertebrate dispersal and revealed the importance of factors such as physical barriers, synchrony of emergence and taxonomic affiliation.

4. Invertebrates affect the cycling of nutrients and carbon by being a crucial intermediate link between primary producers, detritus pools or primary consumers, and predators higher up in the trophic hierarchy. Suspension feeders increase the retention of carbon. The subsidies of aquatic invertebrates to the terrestrial ecosystem have been shown to be important, as are reciprocal processes such as the supply of terrestrial invertebrates that fall into the water.

5. Future studies are needed both to advance theoretical aspects of landscape ecology pertaining to the invertebrates in riverine systems and to intensify the experimental testing of hypotheses, for example with respect to the scaling of processes and to linkages between the terrestrial and aquatic systems. Another promising avenue is to take advantage of naturally steep environmental gradients, and of systems disturbed by humans, such as regulated rivers. By comparison with unimpaired reference sites, the mechanisms involved might be identified. The use of `natural' experiments, especially where environmental gradients are steep, is another technique with great potential.

Aquatic invertebrates in riverine landscapes

Summary

1
Riparian zones serve several ecological functions for bats. They provide a source of prey and likely provide favourable structural habitats and shelter from predators. Many studies have shown that bats use the space above streams, ponds or riparian vegetation as feeding habitat. These studies, however, have never distinguished between the effects of habitat structure and prey availability on the foraging activities of bats. Such effects can only be distinguished by an experimental approach. We predicted that bat activity along a stream is influenced by the number of emerged aquatic insects.

2
We evaluated the response of terrestrial consumers, insectivorous bats, to changes in the abundance of emergent aquatic insects by conducting a manipulative field experiment. In a deciduous riparian forest in Japan, aquatic insect flux from the stream to the riparian zone was controlled with an insect-proof cover over a 1·2 km stream reach.

3
We estimated the abundance of emergent aquatic and flying terrestrial arthropods near the treatment and control reaches using Malaise traps. The foraging activity of bats was evaluated in both treatment and control reaches using ultrasonic detectors.

4
The insect-proof cover effectively reduced the flux of emergent aquatic insects to the riparian zone adjacent to the treatment reach. Adjacent to the control reach, adult aquatic insect biomass was highest in spring, and then decreased gradually. Terrestrial insect biomass increased gradually during the summer at both treatment and control reaches.

5
Foraging activity of bats was correlated with insect abundance. In spring, foraging activity of bats at the control reach was significantly greater than at the treatment reach, and increased at both sites with increasing terrestrial insect abundance.

6
Our result suggests that the flux of aquatic insects emerging from streams is one of the most important factors affecting the distribution of riparian-foraging bats. As is the case with other riparian consumers, resource subsidies from streams can directly enhance the performance or population density of riparian-dependent bats. To conserve and manage bat populations, it is important to protect not only forest ecosystems, but also adjacent aquatic systems such as streams.

https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1365-2656.2006.01146.x

Effect of emergent aquatic insects on bat foraging in a riparian forest

Ecosystems are differentially open to subsidies of energy, material and organisms. This fundamental ecosystem attribute has long been recognized but the influence of this property on community regulation has not been investigated. We propose that this environmental attribute may explain variation in the strength of trophic cascades among ecosystems. Simply because of gravity, we should predict that systems with convex profiles receive low amounts of subsidies whereas systems with concave profiles act as spatial attractors, and receive high amounts of subsidies. The subsidy hypothesis states that ecosystems with high amounts of allochthonous inputs will experience the strongest trophic cascades. To test this hypothesis, we derive ecosystem models and investigate the effect of location and magnitude of subsidies on the strength of trophic cascades. Predictions from our models support the subsidy hypothesis and highlight the need to consider ecosystems as open to allochthonous flows.

Subsidy hypothesis and strength of trophic cascades across ecosystems.

Rivers produce an abundance of aquatic insects that traverse land, where they can have bottom-up effects on predators, who, in turn, can have top-down effects on terrestrial herbivores. This effect can cascade down to plants. These trophic relationships were demonstrated in a field of stinging nettles, Urtica dioica, along a river in Germany. At the shore compared to similar microhabitats 30-60 m away the abundance and biomass of: midges were highest, spiders were also highest, while herbivorous leafhoppers were lowest. At the shore, nettle plants were less damaged by herbivores and thus had less regrowth. Spiders regularly captured both aquatic midges as well as terrestrial leafhoppers and they captured more individuals of both groups at the shore than further away. Midges supported high densities of shore spiders. This was inferred from correlation of distribution and diet in the absence of other environmental gradients. Removal of spiders from experimental plots caused leafhoppers to increase at the shore, causing more plant damage. These effects were not evident at spider-removal sites away from the shore. This demonstrated that spiders depressed leafhoppers and decreased herbivory on plants only at the shore. It is concluded that aquatic insects had a bottom-up effect on spiders and that this subsidy facilitated a top-down effect that cascaded from spiders to leafhoppers to plants. Similar effects would explain the distribution of arthropods along many rivers. Allochthony connects river food webs with shore food webs, making both components essential for each other.

Helmut Stumpf

Papers

Allochthonous aquatic insects increase predation and decrease herbivory in river shore food webs

Mass-length relationships of spiders and harvestmen (Araneae and Opiliones)

Increase of arachnid abundance and biomass of water shores