Weak Response of Animal Allochthony and Production to Enhanced Supply of Terrestrial Leaf Litter in Nutrient-Rich Lakes

Ecosystems are generally linked via fluxes of nutrients and energy across their boundaries. For example, freshwater ecosystems in temperate regions may receive significant inputs of terrestrially derived carbon via autumnal leaf litter. This terrestrial particulate organic carbon (POC) is hypothesized to subsidize animal production in lakes, but direct evidence is still lacking. We divided two small eutrophic lakes each into two sections and added isotopically distinct maize litter to the treatment sections to simulate increased terrestrial POC inputs via leaf litter in autumn. We quantified the reliance of aquatic consumers on terrestrial resources (allochthony) in the year subsequent to POC additions by applying mixing models of stable isotopes. We also estimated lake-wide carbon (C) balances to calculate the C flow to the production of the major aquatic consumer groups: benthic macroinvertebrates, crustacean zooplankton, and fish. The sum of secondary production of crustaceans and benthic macroinvertebrates supported by terrestrial POC was higher in the treatment sections of both lakes. In contrast, total secondary and tertiary production (supported by both autochthonous and allochthonous C) was higher in the reference than in the treatment sections of both lakes. Average aquatic consumer allochthony per lake section was 27–40%, although terrestrial POC contributed less than about 10% to total organic C supply to the lakes. The production of aquatic consumers incorporated less than 5% of the total organic C supply in both lakes, indicating a low ecological efficiency. We suggest that the consumption of terrestrial POC by aquatic consumers facilitates a strong coupling with the terrestrial environment. However, the high autochthonous production and the large pool of autochthonous detritus in these nutrient-rich lakes make terrestrial POC quantitatively unimportant for the C flows within food webs.     

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.     

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.     

Prey use by web-building spiders: stable isotope analyses of trophic flow at a forest-stream ecotone

A forest-stream trophic link was examined by stable carbon isotope analyses which evaluated the relationship of aquatic insects emerging from a stream to the diets of web-building spiders. Spiders, aquatic and terrestrial prey, and basal resources of forest and stream food webs were collected in a deciduous forest along a Japanese headwater stream during May and July 2001. The ¹³C analyses suggested that riparian tetragnathid spiders relied on aquatic insects and that the monthly variation of such dependence is partly associated with the seasonal dynamics of aquatic insect abundance in the riparian forest. Similarly, linyphiid spiders in the riparian forest exhibited ¹³C values similar to aquatic prey in May. However, their ¹³C values were close to terrestrial prey in both riparian and upland (150m away from the stream) forests during June to July, suggesting the seasonal incorporation of stream-derived carbon into their tissue. In contrast, araneid spiders relied on terrestrial prey in both riparian and upland forests throughout the study period. These isotopic results were consistent with a previous study that reported seasonal variation in the aquatic prey contribution to total web contents for each spider group in this forest, implying that spiders assimilate trapped prey and that aquatic insect flux indeed contributes to the energetics of riparian tetragnathid and linyphiid spiders.     

A stable isotope study of linkages between stream and terrestrial food webs through spider predation

SUMMARY 1. Transfer of carbon from freshwater to terrestrial ecosystems can occur through predation on adult aquatic insects, but the significance of this trophic pathway to the energetics of riparian communities is poorly understood. We used stable isotopes of carbon and nitrogen to explore linkages between aquatic insect production and the nutrition of web‐building and free‐living spiders alongside two streams in the North Island of New Zealand. 2. δ¹³C values for riparian tree leaves (means for each site = ?32.2 and ?30.3‰) were distinct from those of lichens collected from stream channel rocks and instream algae, both of which were similar (?23.4 to ?22.4‰). δ¹⁵N values for leaves were similar at both sites (?3.4 and ?2.7‰), but algae were considerably more depleted in δ¹⁵N atonesite suggesting significant differences in instream nitrogen sources between the twostreams. 3. Isotope values for potential aquatic prey of spiders indicated that aquatic algal production was their primary carbon source at both sites. Terrestrial invertebrates collected and assumed to be potential prey reflected a range of carbon sources and represented several trophic levels. 4. At one site, δ¹³C values indicated a primarily algae‐aquatic insect pathway of carbon transfer to both web‐building and free‐living spider guilds. The other site appeared to have a primarily terrestrial carbon pathway for the free‐living spider guild, and a mixed aquatic‐terrestrial pathway for the web‐building guild. 5. Overall, web‐building spiders were estimated to obtain around 61% of their body carbon from aquatic production compared with 55% for free‐living spiders. Our findings suggest that consumption of prey derived from aquatic sources can provide significant nutrition for spiders living along some stream channels. This pathway may represent an important feedback mechanism contributing to the energetics of riparian communities at sites where aquatic insect production is high.     

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.     

Food source of riparian spiders analyzed by using stable isotope ratios

We analyzed the food source of riparian spiders in a middle reach of the Chikuma River, Japan, by using stable isotope ratios of carbon and nitrogen. The carbon and nitrogen isotope ratios of attached algae were higher than those of terrestrial plants, reflecting a large carbon isotope fractionation in terrestrial plants and a difference in nitrogen sources. The carbon isotope ratios of terrestrial insects were similar to those of the terrestrial plants, and the ratios of aquatic insects were scattered between those of the terrestrial plants and the attached algae. The carbon and nitrogen isotope ratios of spiders were intermediate between those of the terrestrial and aquatic insects. The two-source mixing model using the carbon isotope ratio showed that the web-building spiders utilized both the terrestrial and aquatic insects, with large contribution by the aquatic insects (54% on average with a maximum of 92% among spiders taxa collected in each zone), in the riparian area in a middle reach of the Chikuma River. The large contribution of the aquatic insects was often observed for the spiders collected near river channel (<5m) and for the horizontal web-building spiders collected across the riparian area. The relative contribution of the aquatic insects might be related with food availability (distance from river channel) and spiders food preference reflected in their web types (horizontal vs. vertical). Our results showed that organic materials produced in the river channel, in the riparian area, and in the terrestrial area surrounding the riparian area were mixed at the carnivorous trophic level of riparian spiders.     

Habitat structure determines resource use by zooplankton in temperate lakes

While the importance of terrestrial linkages to aquatic ecosystems is well appreciated, the degree of terrestrial support of aquatic consumers remains debated. Estimates of terrestrial contributions to lake zooplankton have omitted a key food source, phytoplankton produced below the mixed layer. We used carbon and nitrogen stable isotope data from 25 Pacific Northwest lakes to assess the relative importance of particulate organic matter (POM) from the mixed layer, below the mixed layer and terrestrial detritus to zooplankton. Zooplankton and deep POM were depleted in 13C relative to mixed layer POM in lakes that can support deep primary production. A Bayesian stable isotope mixing model estimated that terrestrial detritus contributed <5% to zooplankton production, and confirms the role of lake optical and thermal properties; deep POM accounted for up to 80% of zooplankton production in the clearest lakes. These results suggest terrestrial support of lake zooplankton production is trivial.     

Dietary Dependence of Predatory Arthropods on Volant Aquatic Insects in Tropical Stream Riparia

The dietary dependence on volant aquatic insects of eight species of predatory arthropods from three different orders was determined by stable isotope analyses in combination with three‐source, two‐isotope (C and N) Bayesian mixing models. The predators were collected from riparian zones along three streams in tropical Hong Kong during both the wet and dry seasons. Dietary importance of aquatic insects varied according to predator hunting modes, and showed a consistent pattern across all sites during the wet season. The web‐building tetragnathid spider (Orsinome diporusa) had the greatest reliance (~40–55%) on this water‐to‐land subsidy, followed by two species of damselflies (40–50%), three cursorial spiders (Lycosidae, Pisauridae, and Sparassidae: 32–51%) and two neustic gerrids (17–36%). Such reliance also varied according to the microhabitat preferences of different cursorial spiders. Four species of predators (gerrids and cursorial spiders) that were active year‐round showed generally consistent reliance on aquatic insects between seasons, which probably reflected the observed lack of seasonal variability in the relative proportions of aquatic and terrestrial prey. There was a marked overlap in isotopic signatures of aquatic and terrestrial prey at all sites which, combined with the absence of data on the extent to which isotopic fractionations may vary among individual species of prey and predators, contributes some uncertainty to the estimates of dietary compositions derived by mixing models. The findings of the present study are thus likely to be indicative rather than definitive.     

Lake‐derived midges increase abundance of shoreline terrestrial arthropods via multiple trophic pathways

Aquatic insects link adjacent ecosystems by transporting nutrients, energy, and material as they move from bodies of water into terrestrial habitats. Insects emerging from streams and rivers are known to benefit arthropod predators such as spiders, but their influence may extend to other arthropod feeding groups as well. We conducted a terrestrial arthropod survey at a series of lakes spanning a strong gradient of midge (Chironomidae, Diptera) emergence. These small, short‐lived insects reach high densities in some areas such that their carcasses litter the ground, and serve as a potential resource for non‐predatory arthropods. Our study revealed that arthropod assemblages in areas of high midge density were significantly different from those with few midges, the result of an increase of all taxa rather than changes in taxonomic composition. Eight of nine terrestrial arthropod taxa sampled showed a strong positive response to the presence of midges including detritivores and herbivores in addition to predators. Taxa that could consume living or dead midges directly responded especially strongly to midge gradients. Our results strongly suggest that midges enter the terrestrial arthropod food web through multiple pathways, increasing numbers of a wide range of arthropods. Furthermore, they emphasize the importance of lakes as sources of aquatic insects that significantly alter processes in the neighboring terrestrial environment.     

Ecosystem linkages revealed by experimental lake-derived isotope signal in heathland food webs

Cross-ecosystem movement of nutrients and biomass can have important effects on recipient systems. Emerging aquatic insects are subsidies to terrestrial ecosystems and can influence foodweb interactions in riparian systems. In a 2-year field experiment, we simulated aquatic insect deposition by adding adult midge carcasses (150?g dry mass m^?2 year^?1) to 1-m² heathland plots at a site with low natural midge deposition. We established four levels of midge-addition treatments and measured stable isotopes (??¹³C and ??¹⁵N) in plants and arthropods within each treatment. We used a multiple-source isotope Bayesian mixing model to estimate the terrestrial versus aquatic contribution to the diets of arthropods. Aquatic resources were incorporated into plant, detritivore, and predator biomass. Detritivorous Collembola showed the greatest difference in isotope values (+3??? ??¹⁵N and +4??? ??¹³C) between midge-addition and reference treatments. Isotope values of small spiders followed the same trend of enrichment as Collembola while other arthropods (mites and large spiders) were only enriched after 2?years of midge addition. Although predator diets did not change, they became isotopically enriched via their likely prey (Collembola). Plants also had elevated ??¹⁵N (+1???) in midge-addition treatments. The time required and amount of midge-derived C and N detected varied and depended on trophic position. Midge-derived nutrients were no longer present in arthropod biomass in the year following midge addition. Aquatic insect carcasses can be rapidly incorporated into terrestrial food webs in nearshore habitats, and repeated inputs can be detected at multiple trophic levels, thus highlighting the importance of the detrital pathway for aquatic to terrestrial cross-ecosystem subsidies.     

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.     

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.     

Shoreline urbanization interrupts allochthonous subsidies to a benthic consumer over a gradient of lake size

The role of resource subsidies across ecosystem boundaries has emerged as an important concept in contemporary ecology. For lake ecosystems, this has led to interest in quantifying the contribution of terrestrial allochthonous carbon to aquatic secondary production. An inverse relationship between habitat area and the role of allochthonous subsidies has been documented on marine islands and assumed for lakes, yet there have been no tests of this pattern among benthic (lake bottom) consumers. Here, we used carbon stable isotopes to trace terrestrial allochthonous and benthic autochthonous carbon use by the crayfish Pacifastacus leniusculus over a gradient of lake area, productivity and urbanization. Consistent with findings from terrestrial islands, habitat size dictated the importance of allochthonous subsidies, as P. leniusculus transitioned from using predominantly terrestrial carbon in small lakes to an increased reliance on autochthonous production in larger lakes. However, shoreline urbanization interacted with this pattern, particularly for small lakes where greater urbanization resulted in reduced use of allochthonous resources. As such, we provide, to our knowledge, the first confirmation of the predicted relationship between habitat size and importance of allochthonous subsidies to lake benthic consumers, but found that urbanization can interfere with this pattern.     

Carbon sources for lake food webs in the Canadian High Arctic and other regions of Arctic North America

We investigated the role of autochthonous and terrestrial carbon in supporting aquatic food webs in the Canadian High Arctic by determining the diet of the dominant primary consumer, aquatic chironomids. These organisms were studied in fresh waters on 3 islands of the Arctic Archipelago (~74–76°N) including barren polar desert watersheds and a polar oasis with lush meadows. Stomach content analysis of 578 larvae indicated that chironomids primarily ingested diatoms and sediment detritus with little variation among most genera. Carbon and nitrogen stable isotope mixing models applied to 2 lakes indicated that benthic algae contributed 68–95% to chironomid diet at a polar desert site and 70–78% at a polar oasis site. Detritus, originating from either phytoplankton or terrestrial sources, also contributed minor amounts to chironomid diet (0–32%). Radiocarbon measurements for the 2 lakes showed that old terrestrial carbon did not support chironomid production. Carbon stable isotope ratios of chironomids in other High Arctic lakes provided further dietary evidence that was consistent with mixing model results. These findings indicate that, in the Canadian High Arctic, chironomids (and fish that consume them) are supported primarily by benthic algae in both polar desert and oasis lakes. In contrast, our review of carbon flow studies for lakes in other Arctic regions of North America shows that terrestrial carbon and phytoplankton can be important energy sources for consumers. This study provides a baseline to detect future climate-related impacts on carbon pathways in High Arctic lakes.     

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

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Climate change modifies the size structure of assemblages of emerging aquatic insects

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The influence of freshwater-lake subsidies on invertebrates occupying terrestrial vegetation

Studies investigating effects of aquatic-derived resource subsidies have often found large effects on terrestrial systems. Those studies have mostly been performed on effects of subsidies derived from oceanic and riverine systems, and very few have considered effects of subsidies from freshwater lakes. However, since lakes can produce large quantities of emergent aquatic insects that end up on nearby land, it is likely that also freshwater-lake subsidies influence terrestrial systems. We performed sweep-net collections of aquatic and terrestrial invertebrates at varying distances from the shore on vegetation of islands of varying size, in two freshwater lakes in northern Sweden, as well as on the surrounding mainland. We found that the amounts of aquatic insects on terrestrial vegetation decreased with distance from the shore, and that they were the most abundant on small islands, presumably because small islands have a higher perimeter-to-area ratio. Web-building spiders responded positively to the aquatic subsidy by being the most abundant on small islands and by showing a positive relationship with aquatic insect biomass. However, distance from the shore showed no effects on the spiders. Our results strongly support the view that terrestrial systems are subsidized by lakes, and indicate that freshwater-lake subsidies are important for terrestrial invertebrate community structure on adjacent land. Further, our study shows that ecosystems should be treated as interdependent, not as self-contained units, and may as such be important for an increased understanding of the nature and importance of resource flows across ecosystem boundaries.     

Indirect effects of introduced trout on Cascades frogs (Rana cascadae) via shared aquatic prey

1. The introduction of trout to montane lakes has negatively affected amphibian populations across the western United States. In northern California’s Klamath–Siskiyou Mountains, introduced trout have diminished the distribution and abundance of a native ranid frog, Rana (=Lithobates) cascadae. This is primarily thought to be the result of predation on frog larvae. However, if trout feed on larval aquatic insects that are available to R. cascadae only after emergence, then resource competition may also affect this declining native amphibian. 2. Stomach contents of R. cascadae were compared between lakes that contained trout and those from which introduced trout were removed. Total prey mass in stomach contents relative to frog body mass was not significantly different between lakes with fish and fish‐removal lakes, but in the former R. cascadae consumed a smaller proportion of adult aquatic insects. The stomach contents of fish included larvae of aquatic insects that are, as adults, eaten by R. cascadae. 3. Rana cascadae consumed fewer caddisflies (Trichoptera) and more grasshoppers (Orthoptera) at lakes with higher densities of fish. At lakes with greater aquatic habitat complexity, R. cascadae consumed more water striders (Hemiptera: Gerridae) and terrestrial spiders (Araneae). 4. We suggest that reductions in the availability of emerging aquatic insects cause R. cascadae to consume more terrestrial prey where trout are present. Thus, introduced trout may influence native amphibians directly through predation and, indirectly, through pre‐emptive resource competition.