Consumer-specific responses to riverine subsidy pulses in a riparian arthropod assemblage

1. Aquatic resource fluxes from streams can provide significant subsidies for riparian consumers. Because aquatic resource fluxes can be highly variable in space and time, the subsidy efficiency (i.e. transfer to the recipient food web) is controlled by the short‐term aggregative response of riparian consumers. 2. Field manipulations of stream‐derived invertebrate prey subsidies were used to examine specific aggregative responses of ground‐dwelling arthropods to riverine subsidy pulses in a braided‐river (Tagliamento River, NE Italy). Subsidy manipulation comprised short‐term reductions of natural stream‐derived subsidies and increased subsidies of stream‐derived invertebrate prey during four seasons. 3. We hypothesised that specific aggregative responses of riparian arthropods depend on their specialisation on aquatic insects which was inferred from stable isotope analysis. Natural riverine subsidy sources including aquatic insect emergence and surface‐drifting organisms were quantified. 4. Arthropods responded significantly with a reduction in abundance by 51%, at reduced subsidies and an increase by 110% at increased subsidies, when averaged over all seasons. Different arthropod taxa responded differently to subsidy manipulations in relation to their specialisation on aquatic subsidies: ground beetles with a diet consisting predominantly of aquatic insects responded only to subsidy reductions, indicating that their local abundance was not limited by natural stream‐derived subsidies; lycosid spiders with a partly aquatic diet showed no significant response; and ants, although relying on a terrestrial diet, responded positively to added stream‐derived invertebrate prey, indicating that stranding of surface‐drifting terrestrial invertebrates represented an important subsidy pathway. 5. Ground beetles and lycosid spiders were seasonally separated in their use of aquatic subsidies. Results indicate that the life‐history characteristics of riparian consumers can control the subsidy efficiency for the recipient community. By the effective uptake of pulsed riverine‐derived subsidies, riparian arthropods can enhance the transfer of riverine food sources to the riparian food web.     

More than a barrier: The complex effects of ecotone vegetation type on terrestrial consumer consumption of an aquatic prey resource

Community structure and dynamics can be influenced by resource transfers between ecosystems, yet little is known about how boundary structure determines both the magnitude of exchanges and their effects on recipient and donor communities. Aquatic and terrestrial ecosystems are often linked by resource fluxes and riparian vegetation is commonly affected by anthropogenic alterations to land use or river hydrological regime. I investigated whether shrubs at the freshwater–terrestrial interface alter the supply, distribution and importance of aquatic prey resources to terrestrial consumers. Shrubs were predicted to alter the larval community composition of aquatic insects and the emergence of winged adults, thus affecting aquatic prey subsidies to terrestrial consumers. In addition, shrubs were hypothesized to alter the microclimatic suitability of the riparian zone for adult aquatic insects, act as a physical barrier to their dispersal and affect terrestrial community composition, particularly the abundance and type of predators that could benefit from the aquatic prey resource. Stable isotope dietary analyses and a survey of shrub‐dominated and open grassland riparian habitats revealed that larval densities of aquatic insects (EPTM: Ephemeroptera, Plecoptera, Trichoptera and Megaloptera) were higher in shrub than grassland habitats; however, reduced emergence and lateral dispersal in shrub areas led to lower densities of adults. The temperature and relative humidity of the riparian zone did not differ between the habitats. Ground‐active terrestrial invertebrate communities had a higher proportion of cursorial spiders in grassland, coinciding with greater abundances of aquatic prey. Aquatic prey contribution to cursorial spider diet matched adult aquatic insect abundances. Overall, riparian shrubs reduced the magnitude, or at least altered the timing, of cross‐ecosystem subsidy supply, distribution and use by consumers through mechanisms operating in both the aquatic and terrestrial ecosystems. Thus, the structure of ecosystem boundaries has complex effects on the strength of biological interactions between adjacent systems.     

Trophic overlap between fish and riparian spiders: potential impacts of an invasive fish on terrestrial consumers

Studies on resource sharing and partitioning generally consider species that occur in the same habitat. However, subsidies between linked habitats, such as streams and riparian zones, create potential for competition between populations which never directly interact. Evidence suggests that the abundance of riparian consumers declines after fish invasion and a subsequent increase in resource sharing of emerging insects. However, diet overlap has not been investigated. Here, we examine the trophic niche of native fish, invasive fish, and native spiders in South Africa using stable isotope analysis. We compared spider abundance and diet at upstream fishless and downstream fish sites and quantified niche overlap with invasive and native fish. Spider abundance was consistently higher at upstream fishless sites compared with paired downstream fish sites, suggesting that the fish reduced aquatic resource availability to riparian consumers. Spiders incorporated more aquatic than terrestrial insects in their diet, with aquatic insects accounting for 45–90% of spider mass. In three of four invaded trout rivers, we found that the average proportion of aquatic resources in web‐building spider diet was higher at fishless sites compared to fish sites. The probability of web‐building and ground spiders overlapping into the trophic niche of invasive brown and rainbow trout was as high as 26 and 51%, respectively. In contrast, the probability of spiders overlapping into the trophic niche of native fish was always less than 5%. Our results suggest that spiders share resources with invasive fish. In contrast, spiders had a low probability of trophic overlap with native fish indicating that the traits of invaders may be important in determining their influence on ecosystem subsidies. We have added to the growing body of evidence that invaders can have cross‐ecosystem impacts and demonstrated that this can be due to niche overlap.     

Flow regime alters body size but not the use of aquatic subsidies in a riparian predatory arthropod

Alterations to river flow conditions have wide impacts on riparian organisms in terms of behavior and biomass. However, little is known about natural flood impacts on prey use and individual growth of riparian predators. Using stable carbon isotope analysis, we investigated flood impacts on aquatic-prey use and the size structure of an orb-web spider, Nephila clavata, during 3 years under different flood conditions in a black locust forest in the middle reaches of the Chikuma River. Large floods depressed aquatic-prey abundance, but did not affect terrestrial-prey abundance in the riparian forest. Consequently, spider growth was stunted after large floods. Spider body size was positively correlated with the body sizes of both aquatic and terrestrial insects in spider webs, where terrestrial insects were significantly larger than aquatic insects. The δ¹³C of aquatic insects was about 8‰ higher than that of terrestrial insects, and the δ¹³C of both insect groups did not vary significantly between months or among years. A negative relationship was found between body size and δ¹³C in spiders under different subsidies levels. Our results showed that flow regime altered spider growth through changes in aquatic subsidies level, but not aquatic-prey use by the spiders due to relative body sizes of predators and prey. Changes in relative body sizes of predator and prey may be an important factor in understanding nutrients, materials, and energy flows in aquatic and terrestrial linkages in the context of flow regime.     

Impact of extreme flooding on the annual survival of a riparian predator,the Brown Dipper Cinclus pallasii

The predicted effects of global climate change include altered patterns of precipitation and more extreme weather events, leading to an increase in the severity and frequency of episodic disturbances such as floods. These changes may affect lotic prey communities, which could indirectly affect aquatic and riparian predators through trophic linkages. We assessed whether extreme flooding affected the apparent survival of Brown Dippers Cinclus pallasii in Taiwan using mark–resighting data and climate data. The probability of survival was negatively correlated with the degree of flooding, and survival of first‐year birds was lower than that of adults. Previous analyses of this system suggest that the main, indirect mechanism driving such patterns is the impact of flood disturbance on the aquatic invertebrate prey of Brown Dippers. Our results show that changes in prey communities induced by flooding have the potential to affect predators in aquatic and adjacent riparian habitats. This highlights the importance of considering cross‐ecosystem linkages when identifying conservation and management goals in the face of future climate uncertainty.     

Species replacement by a nonnative salmonid alters ecosystem function by reducing prey subsidies that support riparian spiders

Replacement of a native species by a nonnative can have strong effects on ecosystem function, such as altering nutrient cycling or disturbance frequency. Replacements may cause shifts in ecosystem function because nonnatives establish at different biomass, or because they differ from native species in traits like foraging behavior. However, no studies have compared effects of wholesale replacement of a native by a nonnative species on subsidies that support consumers in adjacent habitats, nor quantified the magnitude of these effects. We examined whether streams invaded by nonnative brook trout (Salvelinus fontinalis) in two regions of the Rocky Mountains, USA, produced fewer emerging adult aquatic insects compared to paired streams with native cutthroat trout (Oncorhynchus clarkii), and whether riparian spiders that depend on these prey were less abundant along streams with lower total insect emergence. As predicted, emergence density was 36% lower from streams with the nonnative fish. Biomass of brook trout was higher than the cutthroat trout they replaced, but even after accounting for this difference, emergence was 24% lower from brook trout streams. More riparian spiders were counted along streams with greater total emergence across the water surface. Based on these results, we predicted that brook trout replacement would result in 6–20% fewer spiders in the two regions. When brook trout replace cutthroat trout, they reduce cross-habitat resource subsidies and alter ecosystem function in stream-riparian food webs, not only owing to increased biomass but also because traits apparently differ from native cutthroat trout.     

Growth and development rates in a riparian spider are altered by asynchrony between the timing and amount of a resource subsidy

Rapid growth in response to increased prey abundance may be induced by environmental variability associated with resource subsidies. Spiders living in riparian areas are subject to frequent, episodic bursts of aquatic prey (subsidies). These periods of high resource abundance may occur at different points in recipient consumers’ development through variation in emergence patterns of prey between years or across a landscape. We examine how variable timing of subsidy abundance intersects with life history scheduling to produce different growth and development outcomes for individuals within a population. Through a series of controlled feeding experiments, we tested the hypotheses that the spider Tetragnatha versicolor: (1) exhibits compensatory growth in response to subsidy variability, (2) that rapid increases in mass may result in a greater risk of mortality, and (3) that the timing of subsidy resources relative to the development schedule of this spider may produce different outcomes for individual growth patterns and adult condition. Spiders fed at very high rates grew fastest but also showed evidence of increased mortality risk during moulting. T. versicolor is capable of exhibiting strong growth compensation—individuals suffering initial growth restriction were able to catch up completely with animals on a constant diet utilising the same amount of food. Spiders that received an early pulse of resources (simulating an early arrival of an aquatic insect subsidy to riparian forests) did worse on all measures of development and fitness than spiders that received either a constant supply of food or a late pulse of resources. Importantly, receiving large amounts of food early in life appears to actually confer relative disadvantages in terms of later performance compared with receiving subsidies later in development. Subsidies may provide greater benefits to individuals or age cohorts encountering this resource abundance closer to the onset of reproductive efforts than subsidies arriving early in development.     

Substrate Availability May Be More Important than Aquatic Insect Abundance in the Distribution of Riparian Orb-web Spiders in the Tropics

Spiders that are abundant along streams may depend on energy subsidies across land–water ecotones, but the effects of season and habitat structure on this trophic linkage remain poorly understood in the tropics. We carried out surveys and a manipulative experiment to investigate the effects of season and substrate availability on the distribution of riparian orb-web spiders in Hong Kong, southern China. In the surveys, spider abundance, prey, substrate use, and web orientation were recorded. The experiment involved installation of in-stream artificial substrates (ropes and bamboo poles) to increase substrate availability for web attachment. We found no seasonal difference in web abundance, but seasonal differences were observed for the prey on webs: aquatic insects (mostly Ephemeroptera and chironomid midges) contributed 69 percent of total prey collected during the wet season, but only 38 percent during the dry season. Most webs (50–80%) were < 0.5 m above the water and 45–51 percent of them tended to be orientated horizontally to the water surface and supported by overhanging vegetation and boulders. The addition of artificial substrates resulted in a 23–34 percent increase in the number of webs at the four treatment sites compared to controls, indicating that availability of web-building substrates is a critical determinant of the spider distribution. Our results suggest that riparian zones are potential 'hotspots' of food availability for spiders, and that the aquatic insect subsidy allows this habitat to support increased densities of spiders when the constraint of substrate availability is relaxed.     

Seasonal variation in the trophic structure of a spatial prey subsidy linking aquatic and terrestrial food webs: adult aquatic insects

Research over the past decade has established spatial resource subsidies as important determinants of food web dynamics. However, most empirical studies have considered the role of subsidies only in terms of magnitude, ignoring an important property of subsidies that may affect their impact in recipient food webs: the trophic structure of the subsidy relative to in situ resources. This may be especially important when subsidies are composed of organisms, as opposed to nutrient subsidies, because the trophic position of subsidy organisms may differ from in situ prey. I explored the relative magnitude and trophic structure of a cross-habitat prey subsidy, adult aquatic insects, in terrestrial habitats along three streams in the south–central United States. Overall, adult aquatic insects contributed more than one-third of potential insect prey abundance and biomass to the terrestrial habitat. This contribution peaked along a permanent spring stream, reaching as high as 94% of abundance and 86% of biomass in winter. Trophic structure of adult aquatic and terrestrial insects differed. Nearly all adult aquatic insects were non-consumers as adults, whereas all but one taxon of terrestrial insects were consumers. Such a difference created a strong relationship between the relative contribution of the prey subsidy and the trophic structure of the prey assemblage: as the proportion of adult aquatic insects increased, the proportion of consumers in the prey assemblage declined. Specific effects varied seasonally and with distance from the stream as the taxonomic composition of the subsidy changed, but general patterns were consistent. These findings show that adult aquatic insect subsidies to riparian food webs not only elevate prey availability, but also alter the trophic structure of the entire winged insect prey assemblage.     

Does the strength of cross‐ecosystem trophic cascades vary with ecosystem size? A test using a natural microcosm

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Increasing donor ecosystem productivity decreases terrestrial consumer reliance on a stream resource subsidy

Because nutrient enrichment can increase ecosystem productivity, it may enhance resource flows to adjacent ecosystems as organisms cross ecosystem boundaries and subsidize predators in recipient ecosystems. Here, we quantified the biomass and abundance of aquatic emergence and terrestrial spiders in a reference and treatment stream that had been continuously enriched with nitrogen and phosphorus for 5 years. Because we previously showed that enrichment increased secondary production of stream consumers, we predicted that aquatic emergence flux would be higher in the treatment stream, subsequently increasing the biomass and abundance of terrestrial spiders. Those increases were predicted to be greatest for spiders specializing on aquatic emergence subsidies (e.g., Tetragnathidae). By adding a ¹⁵N stable isotope tracer to both streams, we also quantified nitrogen flow from the stream into the riparian community. Emergence biomass, but not abundance, was higher in the treatment stream. The average body size of emerging adult insects and the relative dominance of Trichoptera adults were also greater in the treatment stream. However, spider biomass did not differ between streams. Spiders also exhibited substantially lower reliance on aquatic emergence nitrogen in the treatment stream. This reduced reliance likely resulted from shifts in the body size distributions and community composition of insect emergence that may have altered predator consumption efficiency in the treatment stream. Despite nutrient enrichment approximately doubling stream productivity and associated cross-ecosystem resource flows, the response of terrestrial predators depended more on the resource subsidy’s characteristics that affected the predator’s ability to capitalize on such increases.     

Responses to River Inundation Pressures Control Prey Selection of Riparian Beetles

Background

Riparian habitats are subjected to frequent inundation (flooding) and are characterised by food webs that exhibit variability in aquatic/terrestrial subsidies across the ecotone. The strength of this subsidy in active riparian floodplains is thought to underpin local biodiversity. Terrestrial invertebrates dominate the fauna, exhibiting traits that allow exploitation of variable aquatic subsidies while reducing inundation pressures, leading to inter-species micro-spatial positioning. The effect these strategies have on prey selection is not known. This study hypothesised that plasticity in prey choice from either aquatic or terrestrial sources is an important trait linked to inundation tolerance and avoidance.

Method/Principal Findings

We used hydrological, isotopic and habitat analyses to investigate the diet of riparian Coleoptera in relation to inundation risk and relative spatial positioning in the floodplain. The study examined patch scale and longitudinal changes in utilisation of the aquatic subsidy according to species traits. Prey sourced from terrestrial or emerging/stranded aquatic invertebrates varied in relation to traits for inundation avoidance or tolerance strategies. Traits that favoured rapid dispersal corresponded with highest proportions of aquatic prey, with behavioural traits further predicting uptake. Less able dispersers showed minimal use of aquatic subsidy and switched to a terrestrial diet under moderate inundation pressures. All trait groups showed a seasonal shift in diet towards terrestrial prey in the early spring. Prey selection became exaggerated towards aquatic prey in downstream samples.

Conclusions/Significance

Our results suggest that partitioning of resources and habitat creates overlapping niches that increase the processing of external subsidies in riparian habitats. By demonstrating functional complexity, this work advances understanding of floodplain ecosystem processes and highlights the importance of hydrological variability. With an increasing interest in reconnecting rivers to their floodplains, these invertebrates represent a key functional element in ensuring that such reconnections have demonstrable ecological value.     

Carbon and nitrogen transfer from a desert stream to riparian predators

Adult aquatic insects emerging from streams may be a significant source of energy for terrestrial predators inhabiting riparian zones. In this study, we use natural abundance delta(13)C and delta(15)N values and an isotopic (15)N tracer addition to quantify the flow of carbon and nitrogen from aquatic to terrestrial food webs via emerging aquatic insects. We continuously dripped labeled (15)N-NH(4) for 6 weeks into Sycamore Creek, a Sonoran desert stream in the Tonto National Forest (central Arizona) and traced the flow of tracer (15)N from the stream into spiders living in the riparian zone. After correcting for natural abundance delta(15)N, we used isotopic mixing models to calculate the proportion of (15)N from emerging aquatic insects incorporated into spider biomass. Natural abundance delta(13)C values indicate that orb-web weaving spiders inhabiting riparian vegetation along the stream channel obtain almost 100% of their carbon from instream sources, whereas ground-dwelling hunting spiders obtain on average 68% of their carbon from instream sources. During the 6-week period of the (15)N tracer addition, orb-web weaving spiders obtained on average 39% of their nitrogen from emerging aquatic insects, whereas spider species hunting on the ground obtained on average 25% of their nitrogen from emerging aquatic insects. To determine if stream subsidies might be influencing the spatial distribution of terrestrial predators, we measured the biomass, abundance and diversity of spiders along a gradient from the active stream channel to a distance of 50 m into the upland using pitfall traps and timed sweep net samples. Spider abundance, biomass and richness were highest within the active stream channel but decreased more than three-fold 25 m from the wetted stream margin. Changes in structural complexity of vegetation, ground cover or terrestrial prey abundance could not account for patterns in spider distributions, however nutrient and energy subsidies from the stream could explain elevated spider numbers and richness within the active stream channel and riparian zone of Sycamore Creek.     

Prothonotary warbler nestling growth and condition in response to variation in aquatic and terrestrial prey availability

Aquatic prey subsidies entering terrestrial habitats are well documented, but little is known about the degree to which these resources provide fitness benefits to riparian consumers. Riparian species take advantage of seasonal pulses of both terrestrial and aquatic prey, although aquatic resources are often overlooked in studies of how diet influences the reproductive ecology of these organisms. Ideally, the timing of resource pulses should occur at the time of highest reproductive demand. This study investigates the availability of aquatic (mayfly) and terrestrial (caterpillar) prey resources as well as the nestling diet of the prothonotary warbler (Protonotaria citrea) at two sites along the lower James River in Virginia during the 2014 breeding season. We found large differences in availability of prey items between the two sites, with one having significantly higher mayfly availability. Nestling diet was generally reflective of prey availability, and nestlings had faster mean growth rates at the site with higher aquatic prey availability. Terrestrial prey were fed more readily at the site with lower aquatic prey availability, and at this site, nestlings fed mayflies had higher mean growth rates than nestlings fed only terrestrial prey. Our results suggest that aquatic subsidies are an important resource for nestling birds and are crucial to understanding the breeding ecology of riparian species.     

The lateral extent of the subsidy from an upland stream to riparian lycosid spiders

Adult aquatic insects emerging from streams can subsidize riparian food webs, but little is known of the spatial extent of these subsidies. Stable isotope (¹⁵N) enrichment of aquatic insects, principally a species of stonefly (Plecoptera: Leuctridae), emerging from an upland stream was used to trace the subsidy from the stream ecosystem to riparian spiders (Lycosidae). The downstream profile of spider δ¹⁵N correlated closely with that of adult stoneflies, indicating that they were deriving nutrition from aquatic sources. The contribution of adult aquatic insects to spider diets was determined using a two-source mixing model. Adult aquatic insects made up over 40% of spider diets adjacent to the stream, but <1% at 20 m from the stream. Enrichment of riparian spiders declined exponentially with distance from the stream channel. Aquatic-terrestrial subsidies were spatially restricted, but locally important, to riparian lycosid spiders at the study site.     

The linkage between riparian predators and aquatic insects across a stream-resource spectrum

1. Spatial subsidies, defined as the flow of energy, nutrients, organisms or pollutants from one habitat to another, have been shown to affect the food–web dynamics in a wide range of ecosystems. An important subsidy to riparian communities is the contribution of adult stream insects to terrestrial predators such as birds, bats and lizards, but also invertebrates including ground and web‐building spiders. 2. We surveyed 37 first‐ and second‐order forest streams across differing environmental gradients in the Central South Island, New Zealand, to investigate the relationship between potential aquatic prey subsidies and predatory riparian arachnids. We anticipated that stream‐insect biomass would be positively associated with riparian arachnids, as a result of emergent adult aquatic insect subsidies to the adjacent habitat. 3. We confirmed positive associations between stream‐insect biomass as a predictor variable and riparian arachnid biomass (R² = 0.42, F_1,34 = 25.2, P < 0.001) and web densities (R² = 0.45, F_1,14 = 11.5, P < 0.01) respectively as dependent variables after adjusting for the confounding effects of environmental variables. Hierarchical partitioning confirmed the importance of stream insect biomass as a statistically significant contributor to the total explained variance in analyses calculated for arachnid biomass, abundance and web density. 4. A concurrent survey of spider‐web density along 20‐m transects from the stream edge into the forest indicated a strong decline in web‐building spider density moving away from the stream (R² = 0.41, F_1,158 = 109, P < 0.001), with stream‐insect biomass as a significant covariate (F_1,149 = 17.7, P < 0.001). 5. Our results suggest that productivity gradients present in the donor system affect the magnitude of the interaction between adjacent habitats. Productivity gradients may lead to increased reciprocal subsidies through a positive feedback loop involving the predation of spiders and other predatory terrestrial invertebrates by aquatic predators. However, terrestrial insectivores such as birds, bats and lizards that are not readily used as prey by aquatic predators may circumvent the feedback cycle by consuming a large proportion of emergent aquatic‐insect biomass. This may lead to asymmetry in the strength of food–web linkages between aquatic and terrestrial habitats.     

Aquatic Terrestrial Linkages Along a Braided-River: Riparian Arthropods Feeding on Aquatic Insects

Rivers can provide important sources of energy for riparian biota. Stable isotope analysis (δ¹³C, δ¹⁵N) together with linear mixing models, were used to quantify the importance of aquatic insects as a food source for a riparian arthropod assemblage inhabiting the shore of the braided Tagliamento River (NE Italy). Proportional aquatic prey contributions to riparian arthropod diets differed considerable among taxa. Carabid beetles of the genus Bembidion and Nebria picicornis fed entirely on aquatic insects. Aquatic insects made up 80% of the diet of the dominant staphylinid beetle Paederidus rubrothoracicus. The diets of the dominant lycosid spiders Arctosa cinerea and Pardosa wagleri consisted of 56 and 48% aquatic insects, respectively. In contrast, the ant Manica rubida fed mainly on terrestrial sources. The proportion of aquatic insects in the diet of lycosid spiders changed seasonally, being related to the seasonal abundance of lycosid spiders along the stream edge. The degree of spatial and seasonal aggregation of riparian arthropods at the river edge coincided with their proportional use of aquatic subsidies. The results suggest that predation by riparian arthropods is a quantitatively important process in the transfer of aquatic secondary production to the riparian food web.     

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

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.     

The demographic and life‐history costs of fear: Trait‐mediated effects of threat of predation on Aedes triseriatus

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Relating body size to the role of aquatic subsidies for the riparian spider <Emphasis Type="Italic">Nephila clavata</Emphasis>

We examined the relationship between body size of the riparian spider Nephila clavata and the contribution of allochthonous (aquatic insects) and autochthonous (terrestrial insects) sources to its diet using stable isotope analysis. During the study period from July to September, the body size of the females increased remarkably (about 60-fold) but that of males remained small. The biomass of both aquatic and terrestrial insects trapped on the spider webs increased with spider size, with the biomass of the former ranging between 30 and 70% of that of the terrestrial insects. The average relative contribution of aquatic insects to the diet of the spiders, calculated from δ¹³C values, was 40–50% in spiders in the early juvenile and juvenile stages, 35% in adult males and 4% in adult females. There was a significant negative relationship between the relative contribution of aquatic insects and body size of the female spiders. We conclude that aquatic insects might be an important seasonal dietary subsidy for small spiders and that these allochthonous subsidies may facilitate the growth of riparian spiders, which may in turn enable the spiders to feed on larger prey.