Riparian Consumers Research Articles

Variation in riparian consumer diet composition and differential bioaccumulation by prey influence the risk of exposure to elements from a recently remediated fly ash spill.

Emerging aquatic insects play a key role in transporting aquatic nutrients and contaminants to riparian consumers. However, little is known about how within- and between-year variation in the diet or patterns of element bioaccumulation in emerging insect taxa may influence the risk of exposure to wildlife. During 2 breeding seasons, the composition of the diet of tree swallows (Tachycineta bicolor) was examined at several colonies that were exposed to element contamination from a remediated coal fly ash spill to determine how variation in the diet influenced the risk of exposure to elements. The proportion of the diet that consisted of insects with an aquatic larval stage was positively related to concentrations of As, Fe, Se, and Tl in the samples. The proportion of the diet that consisted of Chironomidae (midges) was positively related to exposure to these elements at most colonies within and between years. Ephemeroptera (mayflies) contained higher concentrations of Se than midges, including 17 samples with concentrations of Se above 5 µg/g dry mass, the threshold of toxicological concern for birds. This was even the case at colonies several kilometers downstream from the spill. The results indicate that greater consideration should be given to the pattern of element bioaccumulation among different prey taxa and their relative importance in the diet to better assess the risk of contaminant exposure.

Pathways and consequences of contaminant flux to Acadian flycatchers (Empidonax virescens) in urbanizing landscapes of Ohio, USA

A prevalent environmental contaminant, mercury (Hg) is mobile and persistent in aquatic systems, where it often occurs in its bioavailable form methylmercury. Because methylmercury can bioaccumulate in aquatic insects and then transfer to terrestrial food webs, riparian consumers reliant upon aquatic emergent insects, should be disproportionately affected. Using the aerial insectivore Acadian flycatcher (Empidonax virescens) as a focal species, we examined (1) the extent to which total Hg loads in breeding flycatchers affected body condition and reproductive output and (2) potential pathways of contaminant flux in 19 riparian forest fragments distributed across an urban-to-rural landscape gradient in Ohio, USA. From April–August 2011–2012, we collected blood samples from adult (n=76) and nestling (n=17 from 7 nests) flycatchers, monitored their annual reproductive success (i.e., total number of fledglings), and sampled water, sediment, and aquatic emergent insects at each site. Hg concentrations in adult flycatcher blood (47 to 584μg/kg, x¯=211.8, SD=95.5) were low relative to published advisory levels and not related to body condition. However, even at low concentrations, blood Hg was negatively related to reproductive success, with a 0.83 decline in the number of fledglings per μg/kg (loge) increase of blood Hg. Adult flycatchers had 11× greater concentrations of blood Hg than their offspring. Hg levels in flycatcher blood were not predicted by Hg concentrations in sediment, water, or aquatic emergent insects, with the exception of rural landscapes alone, in which flycatcher Hg was negatively related to sediment Hg. In addition to illustrating the difficulty of predicting exposure pathways that may vary among landscape contexts, our study provides evidence that even trace levels of contaminants may impair reproductive success of free-living songbirds.

Cross‐ecosystem impacts of stream pollution reduce resource and contaminant flux to riparian food webs

The effects of aquatic contaminants are propagated across ecosystem boundaries by aquatic insects that export resources and contaminants to terrestrial food webs; however, the mechanisms driving these effects are poorly understood. We examined how emergence, contaminant concentration, and total contaminant flux by adult aquatic insects changed over a gradient of bioavailable metals in streams and how these changes affected riparian web-building spiders. Insect emergence decreased 97% over the metal gradient, whereas metal concentrations in adult insects changed relatively little. As a result, total metal exported by insects (flux) was lowest at the most contaminated streams, declining 96% among sites. Spiders were affected by the decrease in prey biomass, but not by metal exposure or metal flux to land in aquatic prey. Aquatic insects are increasingly thought to increase exposure of terrestrial consumers to aquatic contaminants, but stream metals reduce contaminant flux to riparian consumers by strongly impacting the resource linkage. Our results demonstrate the importance of understanding the contaminant-specific effects of aquatic pollutants on adult insect emergence and contaminant accumulation in adults to predict impacts on terrestrial food webs.

Riparian swallows as integrators of landscape change in a multiuse river system: Implications for aquatic-to-terrestrial transfers of contaminants

Recent research has highlighted the transfer of contaminants from aquatic to terrestrial ecosystems via predation of aquatic emergent insects by riparian consumers. The influence of adjacent land use and land cover (LULC) on aquatic-to-terrestrial contaminant transfer, however, has received limited attention. From 2010 to 2012, at 11 river reaches in the Scioto River basin (OH, USA), we investigated the relationships between LULC and selenium (Se) and mercury (Hg) concentrations in four species of riparian swallows. Hg concentrations in swallows were significantly higher at rural reaches than at urban reaches (t=−3.58, P<0.001, df=30), whereas Se concentrations were positively associated with adjacent land cover characterized by mature tree cover (R2=0.49, P=0.006). To an extent, these relationships appear to be mediated by swallow reliance on aquatic emergent insects. For example, tree swallows (Tachycineta bicolor) at urban reaches exhibited a higher proportion of aquatic prey in their diet, fed at a higher trophic level, and exhibited elevated Se levels. We also found that both Se and Hg concentrations in adult swallows were significantly higher than those observed in nestlings at both urban and rural reaches (Se: t=−2.83, P=0.033, df=3; Hg: t=−3.22, P=0.024, df=3). Collectively, our results indicate that riparian swallows integrate contaminant exposure in linked aquatic–terrestrial systems and that LULC may strongly regulate aquatic contaminant flux to terrestrial consumers.

The fire pulse: wildfire stimulates flux of aquatic prey to terrestrial habitats driving increases in riparian consumers

We investigated the midterm effects of wildfire (in this case, five years after the fire) of varying severity on periphyton, benthic invertebrates, emerging adult aquatic insects, spiders, and bats by comparing unburned sites with those exposed to low severity (riparian vegetation burned but canopy intact) and high severity (canopy completely removed) wildfire. We observed no difference in periphyton chlorophyll a or ash-free dry mass among different burn categories but did observe significantly greater biomass of benthic invertebrates in both high severity burned and unburned reaches versus low severity burned reaches. Moreover, a significantly greater flux of adult aquatic insect emergence occurred at sites that experienced high severity fire versus low severity burned and unburned sites. The greatest number of spiders and bat echolocation calls were also observed at sites of high severity fire. Our results suggest that fires of different severity may have very different affects on stream-riparian food webs and that high severity wildfire may lead to an extended “fire pulse” that stimulates aquatic productivity and flux of prey to terrestrial habitats, driving local increases in riparian consumers.

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.

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

Summary1. 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.

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

Summary1. 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.

A basal aquatic-terrestrial trophic link in rivers: algal subsidies via shore-dwelling grasshoppers.

Rivers provide important resources for riparian consumers, especially in arid or seasonally arid biomes. Pygmy grasshoppers (Paratettix aztecus and P. mexicanus; Tetrigidae) graze river algae stranded along shorelines of the South Fork Eel River in northern California (39°44'N, 123°39'W) as the river recedes during the summer drought. Densities of tetrigids during the mid to late summer were highest (1individual/m2 in July) within 1m of the river margin, and declined to near zero at 4m from the margin, especially during peak temperatures in the afternoon. These observations suggested that the distribution of tetrigids was determined by the availability of algae, water, or both. We manipulated the presence/absence of water and beached algae (Cladophora glomerata) in a 2×2 factorial design. All treatments were positioned 2m upslope from the river's edge (about 30cm above the water table), where the cobble bar was naturally dry and devoid of algae and densities of tetrigids were lower than at the river margin (0.4 individuals/m2 in July). Tetrigids responded only to the wet Cladophora treatment, which had 30× higher densities than other treatments. Stable isotopic signatures (δ13C) of tetrigids (-19.7‰) collected from the same cobble bars were more similar to those of epilithic algae (-20.4‰) than terrestrial plants (-28.2‰), and higher than those of acridid grasshoppers (-27.9‰) from the same habitats. Mixing models suggest that 88-100% of the C in tetrigid grasshoppers at our study site is derived from riverine algae. A preliminary analysis suggests that tetrigids ingested sufficient quantities of algae to easily meet their energetic demands during the summer. This study supports the idea that algae, produced in stream systems, can determine the distribution and relative abundance of a common terrestrial scavenger and provide an additional pathway for energy exchange between rivers and riparian food webs.