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.     

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.     

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.     

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.     

Riparian vegetation loss, stream channelization, and web-weaving spiders in northern Japan

Removal of riparian vegetation and straightening of stream channels (channelization) are the most prevalent forms of habitat degradation in streams and their riparian zones. Both have direct effects on organisms in the habitats where they occur, but also have potential to cause indirect effects by interrupting the flux of invertebrate prey between the two adjacent ecosystems. We measured abundance of web-building riparian spiders along four types of streams in Hokkaido, Japan: relatively undisturbed streams, streams where riparian vegetation had been removed, previously channelized streams where the banks had revegetated, and streams that had been both channelized and had the vegetation removed. Spider abundance was reduced by 70% or more by either habitat disturbance alone, or both combined, and the number of spider families was also reduced. Spiders of the family Tetragnathidae, which specialize in capturing adult insects emerging from streams, were strongly reduced by either form of habitat degradation alone, or in combination. In contrast, abundance of spiders in other families that capture prey from both terrestrial and aquatic sources was reduced more strongly by vegetation loss than channelization. These results indicate that riparian vegetation loss has strong direct effects on spiders by reducing habitat for web sites. They also suggest that channelization can have strong indirect effects on riparian-specialist tetragnathid spiders, probably by reducing the flux of adult aquatic insects from the stream to the riparian zone.     

Land‐use effects on terrestrial consumers through changed size structure of aquatic insects

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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.     

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.     

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.     

Stream meanders increase insectivorous bird abundance in riparian deciduous forests

Adult aquatic insects emerging from streams are a fundamental resource sustaining riparian bird communities in broad-leaved deciduous forests. We investigated how stream geomorphology affects the aquatic insect flux and insectivorous bird abundance in 26 riparian-forest plots during spring season in northern Japan. Lateral dispersal of emergent aquatic insects into the riparian forest exponentially decreased with distance from the stream. Similar to aquatic insect distribution, flycatchers and gleaners concentrated their foraging attacks around the stream channel, preying intensively upon emergent aquatic insects. In contrast, bark probers consumed fewer emergent aquatic insects. The abundance of flycatchers and gleaners was closely related to stream geomorphology, whereas that of bark probers was associated with snag density in the study plots. A path analysis showed that the study plots with longer stream channels had greater aquatic insect abundance. This can be interpreted as a consequence of the increased amount of both stream edge and stream surface, where emergent aquatic insects readily penetrate. The increased flux of aquatic insects by stream meanders elevated gleaner abundance in the study plots. In addition, their abundance was directly affected by stream length per se. On the other hand, flycatcher abundance was only directly affected by stream length. Flycatchers, which mainly consumed emergent aquatic insects in the air, may have increased in response to the increase in suitable foraging sites (i.e., open spaces adjacent to perches) accompanying longer stream channels. Although the causal links affecting bird abundance differed among guilds, meandering streams apparently support abundant insectivorous birds in riparian forests. Therefore, to conserve riparian bird communities, it will be necessary to maintain the functions of stream geomorphology that affect the magnitude of energy transfer across the forest-stream interface.     

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.     

Dynamics of aquatic insect flux affects distribution of riparian web-building spiders

The flux of emerging aquatic insects from streams can provide a significant energy subsidy to riparian web-building spiders. However, despite the high temporality of aquatic insect emergence, the effects of such aquatic insect dynamics on spider distribution are poorly understood. To examine the relationship, the aquatic insect flux from a headwater stream in a northern Japanese deciduous forest was experimentally manipulated by using a greenhouse-type covering, during May to July. Under natural conditions, the aquatic and terrestrial insect abundances dramatically decreased and increased from May through July, respectively. The experimental reduction of aquatic insect flux depressed the density of horizontal orb weavers (Tetragnathidae) in both May and June, but not in July when aquatic insects were scarce, indicating a temporal limitation on spider distribution by aquatic insect flux. In contrast, the densities of both vertical orb weavers (Araneidae) and sheet weavers (Linyphiidae) were unaffected by the manipulation throughout the study period. These various responses, differing among months or spider guilds, may be attributed to the degree of specialization for aquatic prey in the spiders and their mobility in response to aquatic insect flux. The experimental results provided direct evidence that the temporal dynamics of aquatic insect flux, as well as spider characteristics, were primary factors determining the distributional patterns of riparian web-building spiders.     

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

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 reciprocal connections is helping to define a more holistic perspective of catchments, and has the potential to shape new directions for ecology in general.     

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.     

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.     

Emerging aquatic insects as predators in terrestrial systems across a gradient of stream temperature in North and South America

1. Empirical and theoretical research over the past decade has demonstrated the widespread importance of aquatic subsidies to terrestrial food webs. In particular, adult aquatic insects that emerge from streams and lakes are prey for terrestrial predators. While variation in the magnitude of this subsidy is clearly important, the potential top‐down effects of the predatory adults of some aquatic insects in terrestrial food webs are largely unknown. 2. I used published data on benthic insect density (as a proxy for emergence) in North and South America to explore how the proportion of benthic insects that are predatory as adults varies across a gradient of mean annual stream temperature. 3. The proportion of benthic insects that are predatory as adults varied widely across sites (0–12% by abundance; 0–86% by biomass). There was a positive relationship between mean annual stream temperature and the proportion of predatory adults across all sites, driven largely by the greater abundance/biomass of predatory taxa (e.g. odonates), relative to non‐predators (e.g. midges, mayflies, caddisflies), in tropical than in temperate streams. 4. The ‘trophic structure’ (i.e. the proportion of predators) of emerging adult aquatic insects is an understudied source of variation in aquatic–terrestrial interactions. Incorporation of trophic structure in future studies is needed to understand how future modification of fresh waters may affect adjacent terrestrial food webs through both bottom‐up and top‐down effects.     

Strong land‐use effects on the dispersal patterns of adult stream insects: implications for transfers of aquatic subsidies to terrestrial consumers

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Linking stream habitats and spider distribution: spatial variations in trophic transfer across a forest–stream boundary

In headwater streams, many aquatic insects rely on terrestrial detritus, while their emergence from streams often subsidizes riparian generalist predators. However, spatial variations in such reciprocal trophic linkages remain poorly understood. The present study, conducted in a northern Japanese stream and the surrounding forest, showed that pool–riffle structure brought about heterogeneous distributions of detritus deposits and benthic aquatic insects. The resulting variations in aquatic insect emergence influenced the distributions of riparian web-building spiders. Pools with slow current stored greater amounts of detritus than riffles, allowing more benthic aquatic insects to develop in pools. The greater larval biomass in pools and greater tendency for riffle insects to drift into pools at metamorphosis resulted in an emergence rate of aquatic insects from pools that was some four to five times greater than from riffles. In the riparian forest, web-building spiders (Tetragnathidae and Linyphiidae) were distributed in accordance with the emergence rates of aquatic insects, upon which both spider groups heavily depended. Consequently, the riparian strips bordering pools had a density of tetragnathid spiders that was twice as high as that of the riparian strips adjacent to riffles. Moreover, although limitations of vegetation structure prevented the aggregation of linyphiid spiders around pools, linyphiid density normalized by shrub density was higher in habitats adjacent to pools than those adjacent to riffles. The results indicated that stream geomorphology, which affects the storage of terrestrial organic material and the export of such material to riparian forests via aquatic insect emergence, plays a role in determining the strength of terrestrial–aquatic linkages in headwater ecosystems.     

What predicts the use by brook trout (Salvelinus fontinalis) of terrestrial invertebrate subsidies in headwater streams?

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Leaf litter quality affects aquatic insect emergence: contrasting patterns from two foundation trees

Reciprocal subsidies between rivers and terrestrial habitats are common where terrestrial leaf litter provides energy to aquatic invertebrates while emerging aquatic insects provide energy to terrestrial predators (e.g., birds, lizards, spiders). We examined how aquatic insect emergence changed seasonally with litter from two foundation riparian trees, whose litter often dominates riparian streams of the southwestern United States: Fremont (Populus fremontii) and narrowleaf (Populus angustifolia) cottonwood. P. fremontii litter is fast-decomposing and lower in defensive phytochemicals (i.e., condensed tannins, lignin) relative to P. angustifolia. We experimentally manipulated leaf litter from these two species by placing them in leaf enclosures with emergence traps attached in order to determine how leaf type influenced insect emergence. Contrary to our initial predictions, we found that packs with slow-decomposing leaves tended to support more emergent insects relative to packs with fast-decomposing leaves. Three findings emerged. Firstly, abundance (number of emerging insects m^?2 day^?1) was 25 % higher on narrowleaf compared to Fremont leaves for the spring but did not differ in the fall, demonstrating that leaf quality from two dominant trees of the same genus yielded different emergence patterns and that these patterns changed seasonally. Secondly, functional feeding groups of emerging insects differed between treatments and seasons. Specifically, in the spring collector-gatherer abundance and biomass were higher on narrowleaf leaves, whereas collector-filterer abundance and biomass were higher on Fremont leaves. Shredder abundance and biomass were higher on narrowleaf leaves in the fall. Thirdly, diversity (Shannon’s H′) was higher on Fremont leaves in the spring, but no differences were found in the fall, showing that fast-decomposing leaves can support a more diverse, complex emergent insect assemblage during certain times of the year. Collectively, these results challenge the notion that leaf quality is a simple function of decomposition, suggesting instead that aquatic insects benefit differentially from different leaf types, such that some use slow-decomposing litter for habitat and its temporal longevity and others utilize fast-decomposing litter with more immediate nutrient release.