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.     

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.     

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

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

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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.     

Prey Vulnerability Limits Top-Down Control and Alters Reciprocal Feedbacks in a Subsidized Model Food Web

Resource subsidies increase the productivity of recipient food webs and can affect ecosystem dynamics. Subsidies of prey often support elevated predator biomass which may intensify top-down control and reduce the flow of reciprocal subsidies into adjacent ecosystems. However, top-down control in subsidized food webs may be limited if primary consumers posses morphological or behavioral traits that limit vulnerability to predation. In forested streams, terrestrial prey support high predator biomass creating the potential for strong top-down control, however armored primary consumers often dominate the invertebrate assemblage. Using empirically based simulation models, we tested the response of stream food webs to variations in subsidy magnitude, prey vulnerability, and the presence of two top predators. While terrestrial prey inputs increased predator biomass (+12%), the presence of armored primary consumers inhibited top-down control, and diverted most aquatic energy (∼75%) into the riparian forest through aquatic insect emergence. Food webs without armored invertebrates experienced strong trophic cascades, resulting in higher algal (∼50%) and detrital (∼1600%) biomass, and reduced insect emergence (−90%). These results suggest prey vulnerability can mediate food web responses to subsidies, and that top-down control can be arrested even when predator-invulnerable consumers are uncommon (20%) regardless of the level of subsidy.     

Warming,eutrophication, and predator loss amplify subsidies between aquatic and terrestrial ecosystems

The exchange of organisms and energy among ecosystems has major impacts on food web structure and dynamics, yet little is known about how climate warming combines with other pervasive anthropogenic perturbations to affect such exchanges. We used an outdoor freshwater mesocosm experiment to investigate the interactive effects of warming, eutrophication, and changes in top predators on the flux of biomass between aquatic and terrestrial ecosystems. We demonstrated that predatory fish decoupled aquatic and terrestrial ecosystems by reducing the emergence of aquatic organisms and suppressing the decomposition of terrestrial plant detritus. In contrast, warming and nutrients enhanced cross‐ecosystem exchanges by increasing emergence and decomposition, and these effects were strongest in the absence of predators. Furthermore, we found that warming advanced while predators delayed the phenology of insect emergence. Our results demonstrate that anthropogenic perturbations may extend well beyond ecosystem boundaries by influencing cross‐ecosystem subsidies. We find that these changes are sufficient to substantially impact recipient communities and potentially alter the carbon balance between aquatic and terrestrial ecosystems and the atmosphere.     

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 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.     

Predator diversity effects cascade across an ecosystem boundary

Food webs in different ecosystems are often connected through spatial resource subsidies. As a result, biodiversity effects in one ecosystem may cascade to adjacent ecosystems. I tested the hypothesis that aquatic predator diversity effects cascade to terrestrial food webs by altering a prey subsidy (biomass and trophic structure of emerging aquatic insects) entering terrestrial food webs, in turn altering the distribution of a terrestrial consumer (spider) that feeds on emerging aquatic insects. Fish presence, but not diversity, altered the trophic structure of emerging aquatic insects by strongly reducing the biomass of emerging predators (dragonflies) relative to non‐feeding taxa (chironomid midges). Fish diversity reduced emerging insect biomass through enhanced effects on the most common prey taxa: predatory dragonflies Pantala flavescens and non‐feeding chironomids. Terrestrial spiders (Tetragnathidae) primarily captured emerging chironomids, which were reduced in the high richness (3 spp.) treatment relative to the 1 and 2 species treatments. As a result, terrestrial spider abundance was lower above pools with high fish richness (3 species) than pools with 1 and 2 species. Synergistic predation effects were mostly limited to the high richness treatment, in which fish occupied each level of vertical microhabitat in the water‐column (benthic, middle, surface). This study demonstrates that predator diversity effects are not limited to the habitat of the predator, but can propagate to adjacent ecosystems, and demonstrates the utility of using simple predator functional traits (foraging domain) to more accurately predict the direction of predator diversity effects.     

Low river flow alters the biomass and population structure of a riparian predatory invertebrate

1. Low flows in rivers are predicted to increase in extent and severity in many areas in the future, yet the consequent impacts of river drying on terrestrial communities via (i) changes to riparian microclimatic conditions and (ii) the identity and abundance of emerging aquatic insects available to riparian predators have not been quantified. 2. We investigated the influence of low river flow on a riparian fishing spider, Dolomedes aquaticus, in five New Zealand rivers containing permanently flowing and drying reaches and, in one river, along a longitudinal drying gradient. 3. The biomass of aquatic insects, potential prey for D. aquaticus, declined with low river flows while the abundance of potential terrestrial prey remained similar at all sites. In the replicate rivers, and along the longitudinal drying gradient, spider biomass was lower, and size classes were skewed towards more small and fewer large spiders, in drying sites. A desiccation experiment in the laboratory indicated high sensitivity of the spiders, with prey presence increasing spider survival. 4. Differences in the spatial distribution, biomass and population size structure of spiders were observed along the longitudinal drying gradient and disappeared within 16 days of the water returning to all sites. 5. In total, low river flow affected the biomass of D. aquaticus, as well as their size class structure and spatial distribution. This indicates that low river flows have the potential to affect adjacent terrestrial ecosystems.     

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.     

The importance of aquatic vegetation in beaver diets and the seasonal and habitat specificity of aquatic‐terrestrial ecosystem linkages in a subarctic environment

The extent and ecological significance of trophic linkages across ecosystem boundaries have been the subject of considerable recent research attention. North American beavers Castor canadensis engineer terrestrial influences in aquatic ecosystems by constructing terrestrial food caches near their lodges and aquatic influences in terrestrial ecosystems by building dams and flooding low lying areas. However, it is poorly resolved to what extent beavers rely on aquatic food sources and whether this reliance is greater during winter when ice cover physically confines beavers to aquatic habitats or during summer when warm, ice free water promotes the growth and accessibility of aquatic vegetation. Working in a subarctic region, we surveyed the abundance of aquatic and terrestrial food sources in and around lotic and lentic environments and estimated their contributions to beaver diets during open water and ice covered periods using carbon and nitrogen stable isotope analysis of hair samples. Ponds had four times more aquatic vegetation than streams, but terrestrial habitats around ponds had less than half as much shrub cover as habitats adjacent to streams. Beaver diets in this subarctic environment are estimated to be comprised of 60 to 80% aquatic vegetation, with beavers occupying ponds consuming more aquatic vegetation in winter than beavers occupying streams, which rely more on terrestrial shrubs cached near their lodge. Collectively, these results show how the influence of physical barriers on ecosystem linkages can be modified by habitat‐ and season‐specific abundances of preferred resources and the potential for animals to consume food in ecosystems and seasons different from where and when the food was harvested.     

From bottom-up to top-down control of invertebrate herbivores in a retrogressive chronosequence

In the long-term absence of disturbance, ecosystems often enter a decline or retrogressive phase which leads to reductions in primary productivity, plant biomass, nutrient cycling and foliar quality. However, the consequences of ecosystem retrogression for higher trophic levels such as herbivores and predators, are less clear. Using a post-fire forested island-chronosequence across which retrogression occurs, we provide evidence that nutrient availability strongly controls invertebrate herbivore biomass when predators are few, but that there is a switch from bottom-up to top-down control when predators are common. This trophic flip in herbivore control probably arises because invertebrate predators respond to alternative energy channels from the adjacent aquatic matrix, which were independent of terrestrial plant biomass. Our results suggest that effects of nutrient limitation resulting from ecosystem retrogression on trophic cascades are modified by nutrient-independent variation in predator abundance, and this calls for a more holistic approach to trophic ecology to better understand herbivore effects on plant communities.     

Potential alteration of cross‐ecosystem resource subsidies by an invasive aquatic macroinvertebrate: implications for the terrestrial food web

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Boomerang ecosystem fluxes: organic carbon inputs from land to lakes are returned to terrestrial food webs via aquatic insects

Many ecosystems are linked to their adjacent ecosystems by movements of organisms. For instance, aquatic and terrestrial ecosystems are linked via emerging aquatic insects that serve as prey for terrestrial consumers. However, the role of these organisms in returning recycled carbon to the ecosystem from which it originated is not well known. This is due to the fact that values of carbon isotope signatures from terrestrial leaves and aquatic resources are usually similar and hence results of isotope mixing models need to be considered with caution. We overcame this problem by adding isotopically distinct terrestrial particulate organic carbon (tPOC) as a tracer to the experimental sides of two lakes that were divided in two equal halves with plastic curtains. We focused on aquatic insect larvae (Chironomidae) that fed on maize Zea mays leaves experimentally added to the lakes, and subsequently became prey for terrestrial predators (spiders) after emergence. The carbon isotope values of Chironomidae and spiders were significantly elevated in the lake treatment sides as compared to reference sides, whereas the values of all autochthonous resources were not affected by maize additions. Estimates from stable isotope mixing models indicated a low but demonstrable contribution of maize leaves to the diet of Chironomidae. Overlap between the isotope values of alder leaves, the major natural tPOC source, and autochthonous resources prevented a reliable quantification of allochthony of Chironomidae. However, we qualitatively demonstrated the flow of terrestrial particulate organic carbon to lakes, as leaf fall, and back to terrestrial surroundings via emerging insects. This ‘boomerang’ carbon flux between land and lakes blurs the distinction between autochthonous and allochthonous carbon sources.     

Trophic‐level dependent effects on CO2 emissions from experimental stream ecosystems

Concern over accelerating rates of species invasions and losses have initiated investigations into how local and global changes to predator abundance mediate trophic cascades that influence CO₂ fluxes of aquatic ecosystems. However, to date, no studies have investigated how species additions or losses at other consumer trophic levels influence the CO₂ flux of aquatic ecosystems. In this study, we added a large predatory stonefly, detritivorous stonefly, or grazer tadpole to experimental stream food webs and over a 70‐day period quantified their effects on community composition, leaf litter decomposition, chlorophyll‐a concentrations, and stream CO₂ emissions. In general, streams where the large grazer or large detritivore were added showed no change in total invertebrate biomass, leaf litter loss, chlorophyll‐a concentrations, or stream CO₂ emissions compared with controls; although we did observe a spike in CO₂ emissions in the large grazer treatment following a substantial reduction in chlorophyll‐a concentrations on day 28. However, the large grazer and large detritivore altered the community composition of streams by reducing the densities of other grazer and detritivore taxa, respectively, compared with controls. Conversely, the addition of the large predator created trophic cascades that reduced total invertebrate biomass and increased primary producer biomass. The cascading effects of the predator additions on the food web ultimately led to decreased CO₂ emissions from stream channels by up to 95%. Our results suggest that stream ecosystem processes were more influenced by changes in large predator abundance than large grazer or detritivore abundance, because of a lack of functionally similar large predators. Our study demonstrates that the presence/absence of species with unique functional roles may have consequences for the exchange of CO₂ between the ecosystem and the atmosphere.     

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.     

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.     

Importance of terrestrial subsidies for native brook trout in Appalachian intermittent streams

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