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.     

Riparian ground beetles (Coeloptera, Carabidae) preying on aquatic invertebrates: a feeding strategy in alpine floodplains

The food and feeding habits of riparian ground beetles were studied in four alpine floodplains (Bavaria, Germany): a 5th-order stream (the Isar) and three 3rd-order streams. The riparian fauna along the streams mainly consists of predaceous species. Riparian ground beetle densities were much higher along the Isar than along the small streams. Aquatic invertebrates composed 89% of the potential prey for carnivorous terrestrial insects along the Isar. Besides aquatic organisms washed ashore, stoneflies emerging on land are of considerable importance as potential prey for terrestrial predators. In contrast, only 34% of the potential prey organisms collected along the small streams were of aquatic origin. Food abundance was 9 times higher in the shore region of the Isar compared to the small streams. Surface drift in the Isar, a potentially important food source for riparian organisms, was about 10⁶ organisms and exuviae per meter stream width in 24 h. The drift density in the Isar was 59 times higher than that in a small stream. Terrestrial organisms provided only 3% of the drifting particles in the Isar, but 50% in the small stream. Gut content analysis reveals, that riparian ground beetles in the Isar floodplain mainly feed on aquatic organisms washed ashore or emerging on land. While small Bembidion species prefer chironomids (larvae and adults) the larger species Nebria picicornis feeds on emerging stoneflies, terrestrial riparian organisms and aquatic organisms accumulating along the shoreline. The prey of riparian ground beetles in the floodplain of the three small streams mainly consists of terrestrial species some of which may have been washed ashore. Received: 2 September 1996 / Accepted: 26 February 1997     

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.     

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.     

Diet partitioning in sympatric Atlantic salmon and brown trout in streams with contrasting riparian vegetation

Prey intake by Atlantic salmon Salmo salar and brown trout Salmo trutta was measured across different riparian vegetation types: grassland, open canopy deciduous and closed canopy deciduous, in upland streams in County Mayo, Western Ireland. Fishes were collected by electrofishing while invertebrates were sampled from the benthos using a Surber sampler and drifting invertebrates collected in drift traps. Aquatic invertebrates dominated prey numbers in the diets of 0+ year Atlantic salmon and brown trout and 1+ year Atlantic salmon, whereas terrestrial invertebrates were of greater importance for diets of 1+ and 2+ year brown trout. Terrestrial prey biomass was generally greater than aquatic prey for 1+ and 2+ year brown trout across seasons and riparian types. Prey intake was greatest in spring and summer and least in autumn apart from 2+ year brown trout that sustained feeding into autumn. Total prey numbers captured tended to be greater for all age classes in streams with deciduous riparian canopy. Atlantic salmon consumed more aquatic prey and brown trout more terrestrial prey with an ontogenetic increase in prey species richness and diversity. Atlantic salmon and brown trout diets were most similar in summer. Terrestrial invertebrates provided an important energy subsidy particularly for brown trout. In grassland streams, each fish age class was strongly associated with aquatic, mainly benthic invertebrates. In streams with deciduous riparian canopy cover, diet composition partitioned between conspecifics with older brown trout associated with surface drifting terrestrial invertebrates and older Atlantic salmon associated with aquatic invertebrates with a high drift propensity in the water column and 0+ year fish feeding on benthic aquatic invertebrates. Deciduous riparian canopy cover may therefore facilitate vertical partitioning of feeding position within the water column between sympatric Atlantic salmon and brown trout. Implications for riparian management are discussed.     

Differential reliance on aquatic prey subsidies influences mercury exposure in riparian arachnids and songbirds

Cross‐ecosystem subsidies move substantial amounts of nutrients between ecosystems. Emergent aquatic insects are a particularly important prey source for riparian songbirds but may also move aquatic contaminants, such as mercury (Hg), to riparian food webs. While many studies focus on species that eat primarily emergent aquatic insects, we instead study riparian songbirds with flexible foraging strategies, exploiting both aquatic and terrestrial prey sources. The goal in this study is to trace reliance on aquatic prey sources and correlate it to Hg concentrations in common riparian arachnids (Families Tetragnathidae, Opiliones, and Salticidae) and songbirds (Common Yellowthroat Geothlypis trichas, Spotted Towhee Pipilo maculatus, Swainson''s Thrush Catharus ustulatus, Song Sparrow Melospiza melodia, and Yellow Warbler Setophaga petechia). We used stable isotopes of δ¹³C and δ¹⁵N and Bayesian mixing models in MixSIAR to determine the reliance of riparian predators on aquatic prey sources. Using mixed effects models, we found that arachnid families varied in their reliance on aquatic prey sources. While songbird species varied in their reliance on aquatic prey sources, songbirds sampled earlier in the season consistently relied more on aquatic prey sources than those sampled later in the season. For both arachnids and songbirds, we found a positive correlation between the amount of the aquatic prey source in their diet and their Hg concentrations. While the seasonal pulse of aquatic prey to terrestrial ecosystems is an important source of nutrients to riparian species, our results show that aquatic prey sources are linked with higher Hg exposure. For songbirds, reliance on aquatic prey sources early in the breeding season (and subsequent higher Hg exposure) coincides with timing of egg laying and development, both of which may be impacted by Hg exposure.     

Stable isotope analysis indicates microalgae as the predominant food source of fauna in a coastal forest stream,south‐east Brazil

Abstract: Stable isotope studies of food webs in floodplains, large rivers, mangroves, and seagrasses have shown that, although a large proportion of the biomass may come from higher plants, microalgae provide a disproportionate amount of carbon assimilated by metazoan consumers. Evidence is building that this may also be the case for streams, especially those in the tropics. At the level of individual consumer species we also see that the apparent diet may not be reflected in the carbon assimilated. Tropical streams commonly have omnivore‐detritivore species that potentially show this phenomenon. We tested these concepts in four moderately shaded sites in a stream in well‐preserved Atlantic rainforest at Ilha Grande, Rio de Janeiro. We sampled aquatic insects, shrimps and fish as well as potential terrestrial and aquatic primary food sources. Carbon stocks from terrestrial sources predominated over carbon of algal origin (>99% of total). The primary sources of carbon showed distinctly different isotopic signatures: terrestrial sources had δ¹³C values close to ?30‰, microalgae were ?20‰ and macroalgae were ?25‰. All fauna had δ¹³C values consistent with a carbon source derived from microalgae. Baetid mayflies and atyid shrimps exert a strong grazing pressure on periphyton and organic sediments but appear to assimilate predominantly microalgae. The palaemonid shrimp Macrobrachium olfersi also ingests large amounts of detritus of terrestrial origin, but apparently assimilates animal prey with algal δ¹³C signatures. These results support the growing view that tropical stream food chains are primarily algal based.     

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

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

Opportunistic top predators partition food resources in a tropical freshwater ecosystem

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

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

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.     

Stable isotope evidence indicates the incorporation into Japanese catchments of marine‐derived nutrients transported by spawning Pacific Salmon

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Temporal shift in contribution of terrestrial organic matter to consumer production in a grassland river

1. We used stable isotopes to study the temporal (early summer versus autumn) pattern of use of terrestrial and aquatic sources of organic carbon by consumers in two bedrock‐confined reaches of a grassland river in New Zealand.
2. The major sources of organic carbon available to primary consumers were expected to be terrestrial leaf‐litter and biofilm from the stream channel. These putative carbon sources showed no significant change in mean δ¹³C between summer and autumn. Leaf litter (mean δ¹³C13C compared to biofilm (mean δ¹³C>?19.92).
3. In contrast to leaf litter and biofilm, the δ¹³C of consumers changed over time, being enriched in ¹³C in the autumn compared with early summer. Both the magnitude (>5‰ in some cases) and rapidity of this shift (< 3 months) was surprising.
4. A two‐source mixing model indicated that, during early summer, terrestrial carbon comprised> 50% of tissue carbon for 15 of the 17 taxa of aquatic consumers analysed. During autumn, terrestrial carbon comprised> 50% of the tissue carbon of only five of 25 taxa. Because the mean δ¹³C of putative food sources was consistent over time, the shift in δ¹³C values for consumers is attributed to a change in relative amounts of terrestrial and aquatic carbon available for consumption.
5. Because seston consists of a mixture of many particles of diverse origin, it may provide an integrated measure of catchment‐wide sources of organic matter entering a stream channel. Like the tissues of most consumers, mean δ¹³C values for seston showed a significant shift toward ¹³C enrichment. This indicated that the relative availability of terrestrial carbon decreased from summer to autumn.
6. The actual quantity of carbon contributed to the stream food‐web by this potential terrestrial–aquatic link is unknown. Although terrestrial carbon may comprise a high proportion of the tissue carbon of consumers prior to summer, the majority of secondary production (and carbon sequestration) probably occurs during early summer as a consequence of rising temperature and high quality food in the form of biofilm.     

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.     

Responses to River Inundation Pressures Control Prey Selection of Riparian Beetles

Background

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

Method/Principal Findings

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

Conclusions/Significance

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

Do seasonally fluctuating aquatic subsidies influence the distribution pattern of birds between riparian and upland forests?

Seasonal fluctuation of allochthonous subsidies influences food web structure and dynamics in recipient communities. This study investigated whether aquatic subsidies influence the dynamics of insectivorous birds in entire catchment. We estimated the prey biomass and bird density in riparian and upland habitats in three catchments in temperate deciduous forests in Hokkaido, Japan. Aquatic prey was found only in riparian forests and the biomass peaked in early spring, while terrestrial prey was equally distributed between habitats and increased in biomass in late spring. Bird density was higher in riparian than in upland forests before bud break, when the biomass of aquatic insects peaked, but was similar in both forests during the rest of the seasons. These results suggest that aquatic prey subsidies are used not only by birds inhabiting riparian forests, but also by birds associated with upland forests. Aquatic prey subsidies may be particularly important in the spring as a critical food resource for survival and the breeding activities of birds, thereby, influencing the population dynamics of bird communities.     

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.     

Aquatic and terrestrial invertebrate drift in southern Appalachian Mountain streams: implications for trout food resources

1. We characterised aquatic and terrestrial invertebrate drift in six south‐western North Carolina streams and their implications for trout production. Streams of this region typically have low standing stock and production of trout because of low benthic productivity. However, little is known about the contribution of terrestrial invertebrates entering drift, the factors that affect these inputs (including season, diel period and riparian cover type), or the energetic contribution of drift to trout. 2. Eight sites were sampled in streams with four riparian cover types. Drift samples were collected at sunrise, midday and sunset; and in spring, early summer, late summer and autumn. The importance of drift for trout production was assessed using literature estimates of annual benthic production in the southern Appalachians, ecotrophic coefficients and food conversion efficiencies. 3. Abundance and biomass of terrestrial invertebrate inputs and drifting aquatic larvae were typically highest in spring and early summer. Aquatic larval abundances were greater than terrestrial invertebrates during these seasons and terrestrial invertebrate biomass was greater than aquatic larval biomass in the autumn. Drift rates of aquatic larval abundance and biomass were greatest at sunset. Inputs of terrestrial invertebrate biomass were greater than aquatic larvae at midday. Terrestrial invertebrate abundances were highest in streams with open canopies and streams adjacent to pasture with limited forest canopy. 4. We estimate the combination of benthic invertebrate production and terrestrial invertebrate inputs can support 3.3–18.2 g (wet weight) m⁻² year⁻¹ of trout, which is generally lower than values considered productive [10.0–30.0 g (wet weight) m⁻² year⁻¹]. 5. Our results indicate terrestrial invertebrates can be an important energy source for trout in these streams, but trout production is still low. Any management activities that attempt to increase trout production should assess trout food resources and ensure their availability.     

Disentangling the multiple effects of stream drying and riparian canopy cover on the trophic ecology of a highly threatened fish

1. Understanding risks to aquatic systems posed by changing drought regimes is particularly important for the conservation of already threatened taxa. However, little is known about how local environmental conditions, especially those in heavily human‐influenced situations, interact with regional shifts such as droughts to alter realised impacts on aquatic communities, including threatened top predators.
2. Here, we investigated the combined effects of stream drying intensity and riparian canopy cover on the trophic interactions of critically endangered kōwaro, or Canterbury mudfish (Neochanna burrowsius) in an agricultural area of New Zealand. Fish populations and their potential prey, both terrestrial and aquatic, as well as environmental variables, including riparian canopy cover and drying measured with stage loggers, were sampled over eight visits to 24 sites spanning orthogonal drying and canopy gradients. Stable isotope ratios, ¹³C/¹²C and ¹⁵N/¹⁴N, were used to investigate trophic links between mudfish and their terrestrial and aquatic prey across these gradients.
3. When non‐native willows (predominantly Salix fragilis) dominated the riparian canopy, increased tree cover led to elevated drying intensity, probably driven by their relatively high water demands compared to other trees. However, in the absence of willows, canopy cover had no effect on drying intensity. Although this was the only direct link between these two environmental factors, they had opposing effects on kōwaro populations, which will be important for management under drought.
4. Increased drying intensity contributed to elevated abundance of microcrustacea and aquatic Diptera larvae, and an increase in the relative abundance of kōwaro juveniles. However, drying‐affected kōwaro populations also had fewer large reproductive adults and elevated δ¹⁵N values, probably driven by physiological limitations and an increase in kōwaro cannibalism, respectively.
5. By comparison, increased canopy cover enhanced input of terrestrial invertebrates, a food resource for larger kōwaro, leading to elevated kōwaro δ¹³C values, no effects on δ¹⁵N values, and higher relative abundance of large kōwaro in shaded streams compared to unshaded streams. Thus, the riparian canopy cover was able to offset some of the effects of drying.
6. Overall, we found no interactions between drying intensity and canopy cover affecting kōwaro. However, their opposing effect highlights the important role local conditions such as riparian canopies play on aquatic communities and their potential role as a restoration tool to mitigate the effects of large‐scale shifts such as drought.

     

Shifts in reciprocal river‐riparian arthropod fluxes along an urban‐rural landscape gradient

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