Influences of river regulation and environmental variables on upland bird assemblages in northern Sweden

Most large rivers in Sweden are regulated to produce hydropower. This transformation from free-flowing rivers to chains of elongate run-of-river impoundments has been shown to have consequences for aquatic, riparian and adjacent upland environments, and for the emergence patterns of aquatic insects that are important for terrestrial consumers. In this study, we investigated bird assemblages in upland-forest environments along seven large rivers (three heavily impounded and four free flowing) in northern Sweden. Bird densities were assessed by point counts in the breeding and post-breeding seasons. While we observed no significant differences in bird species richness between regulated and free-flowing rivers, cumulative densities of two feeding groups of birds (those feeding on seeds and/or large insects and those feeding on small insects) were higher along free-flowing rivers than along regulated rivers in the breeding season, consistent with known differences in aquatic-insect emergence. Further, ordination analyses showed seasonal shifts in bird assemblage structure, and that these shifts differed between regulated and free-flowing rivers and between the two feeding groups. However, the variables explaining the most variance (11?C28?%) in bird assemblage structure were related to a gradient of agricultural-to-forest land use. River regulation contributed to the model in the post-breeding season, but was of relatively low importance. Nevertheless, the observed contrasting seasonal shifts in upland-forest bird assemblage structure between regulated and free-flowing rivers suggest that regulation-induced modifications of aquatic-insect emergence and subsequent changes in prey availability to the birds are also important considerations.     

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.     

Seasonal variation in the trophic structure of a spatial prey subsidy linking aquatic and terrestrial food webs: adult aquatic insects

Research over the past decade has established spatial resource subsidies as important determinants of food web dynamics. However, most empirical studies have considered the role of subsidies only in terms of magnitude, ignoring an important property of subsidies that may affect their impact in recipient food webs: the trophic structure of the subsidy relative to in situ resources. This may be especially important when subsidies are composed of organisms, as opposed to nutrient subsidies, because the trophic position of subsidy organisms may differ from in situ prey. I explored the relative magnitude and trophic structure of a cross-habitat prey subsidy, adult aquatic insects, in terrestrial habitats along three streams in the south–central United States. Overall, adult aquatic insects contributed more than one-third of potential insect prey abundance and biomass to the terrestrial habitat. This contribution peaked along a permanent spring stream, reaching as high as 94% of abundance and 86% of biomass in winter. Trophic structure of adult aquatic and terrestrial insects differed. Nearly all adult aquatic insects were non-consumers as adults, whereas all but one taxon of terrestrial insects were consumers. Such a difference created a strong relationship between the relative contribution of the prey subsidy and the trophic structure of the prey assemblage: as the proportion of adult aquatic insects increased, the proportion of consumers in the prey assemblage declined. Specific effects varied seasonally and with distance from the stream as the taxonomic composition of the subsidy changed, but general patterns were consistent. These findings show that adult aquatic insect subsidies to riparian food webs not only elevate prey availability, but also alter the trophic structure of the entire winged insect prey assemblage.     

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.     

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.     

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

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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.     

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.     

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.     

Positive indirect effect of aquatic insects on terrestrial prey is not offset by increased predator density

1. Changes in one prey species' density can indirectly affect the abundance of another prey species if a shared predator eats both species. Sometimes, indirect effects occur when prey straddle habitats, including when riparian predator populations grow in response to emergent aquatic insects and increase predation on terrestrial prey. However, predators may largely switch to aquatic insects or become satiated, reducing predation on terrestrial prey. 2. To determine the net indirect effect of aquatic insects on terrestrial arthropods via generalist spider predators, a field experiment was conducted mimicking midge influx and a wolf spider numerical response inside enclosures near an Icelandic lake. Lab mesocosms were also used to assess per capita rates of spider predation u nder differing levels of midge abundance. 3. Midges always decreased sentinel prey predation, but this effect increased with predator density. When midges were absent, predation increased 30% at a high spider density, but predation was equal between spider treatments when midges were present. In situ arthropods showed no effect of midge or spider treatments, although non‐significant abundance patterns were observed congruent with sentinel prey results. 4. In lab mesocosms, prey survivorship increased ≥50% where midges were present and rapidly saturated; the addition of 5, 20, 50, and 100 midges equivalently reduced spider predation, supporting predator distraction rather than satiation as the root cause. 5. The present results demonstrate a strong positive indirect effect of midges and broadly support the concept that predator responses to alternative prey are a major influence on the magnitude and direction of predator‐mediated indirect effects.     

Stream meanders increase insectivorous bird abundance in riparian deciduous forests

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

The predators of insects

Abstract. 1. Insect–insectivore trophic relations were reviewed using presence–absence data from sixty-one invertebrate-dominated food webs and fifteen food webs from Briand's (1983) original forty web collection. From counts of prey links in higher taxa (orders, classes, phyla), six phyla and thirteen classes of non-insect insectivores and fourteen orders of insect predators and prey were found. 2. Detritus-based habitats (phytotelmata, felled logs, carcasses, dungpads) harboured fewer orders of insects, that interact with other insects, than webs from grazer-based (host plants, some galls) and mixed-based systems (aquatic webs). Consumer–resource networks of higher insect taxa in these webs shared several features found in some species-level biological networks: the trend was towards few pairs of strong asymmetrical links, several weak links and many null interactions. 3. From counts of insect predator–insect prey links, hymenopterans as terrestrial predators and parasitoids interacted with the most number of higher insect taxa. Hymenopterans were also linked as prey more often than other terrestrial insects. In freshwater habitats, plecopterans were linked as predators more often than other aquatic taxa, whereas dipterans were listed as prey more often than other insects. 4. Dipterans were linked in the diets of non-insect insectivores from seven of eight common taxonomic classes. Arachnids were identified as insect predators by food web researchers in the largest number of webs, followed by passerine birds and cyprinodont fishes. From analysis of prey links at the ordinal level, predaceous insects were less polyphagous than other predators (other ectotherms and endotherms). 5. Analysis of chain lengths, as expected, showed that insect prey occupied mostly lowermost trophic levels, non-insect insectivores were found mostly at uppermost trophic levels, and predaceous insects were found mostly at intermediate trophic levels across most habitats. 6. This analysis offers evidence that insects are not just occupying intermediate trophic levels in some communities. Indeed, some taxa feed at the upper ends of long food chains, for example eupelmids in galls, staphylinids in carcasses, and perlid plecopterans in streams.     

Habitat requirements of the Seychelles Black Paradise Flycatcher Terpsiphone corvina: a re-evaluation of translocation priorities

The critically endangered Seychelles Black Paradise Flycatcher Terpsiphone corvina was once found on at least five of the inner (granitic) islands in the Seychelles archipelago, western Indian Ocean. Currently, it is only found on two islands, with c. 98% of the world population (c. 150–200 individuals) occurring on the 10‐km² island of La Digue. Creation of additional island populations is therefore considered crucial in improving its conservation status. The presence of native broad‐leaved plateau forest in proximity to wetland areas is proposed as an important selection criterion when considering the suitability of other islands for translocation, due to the presumed importance of insect prey (dependent on water) in the diet. We quantified habitat use, territory composition, the effect of water on invertebrate abundance, and foraging and breeding success to determine the importance of native broad‐leaved woodland and wetland areas for Flycatchers. Flycatcher territories contained significantly more native broad‐leaved woodland (88%) than its availability on the plateau (43%); Flycatchers used native tree species significantly more for both foraging (81%) and nesting (95%) than their availability within territories (71%); and territory size varied inversely with the density of native broad‐leaved tree species. Native broad‐leaved forest was associated with semipermanent and permanent water bodies and availability of native forest habitat was a good predictor of territory distribution. The number of aerial insects trapped was higher close to water, but there was no effect of the proximity to water on the number of insects counted on foliage. The majority of identified prey species in adult diet were not dependent on water: Orthoptera and Lepidoptera were the most common prey groups, comprising 66% of identified prey items. There was no effect of proximity to water on foraging or breeding success (c. 35%): depredation was the major factor determining breeding success, and accounted for the majority of nest failures. The importance of wetland areas to Flycatchers therefore appears to have been over‐emphasized. The implications for translocation and conservation of the Flycatcher are discussed.     

Nestbox orientation: a species-specific influence on occupation and breeding success in woodland passerines

Capsule Nestbox orientation has species-specific influences on nestbox occupation and breeding success for woodland passerines.

Aims To determine if nestbox orientation had an influence upon nestbox selection or breeding success for three co-occurring woodland passerines.

Methods We analysed 15 consecutive years of breeding data (1990–2004) from 295 nestboxes in the UK using circular statistical analyses to examine the influence of orientation upon nestbox occupation and breeding success for three species, Blue Tit Cyanistes caeruleus, Great Tit Parus major and Pied Flycatcher Ficedula hypoleuca.

Results The three species used nestboxes of all orientations during the 15-year period. The frequency of nestbox occupation by Great Tits correlated with orientation (the mean number of nests in boxes oriented south-southwest was lower than the mean number of nests in boxes facing other directions). There was no such relationship for Blue Tits or Pied Flycatchers. Nestbox orientation influenced the breeding success of Pied Flycatcher (the mean number of young to fledge from boxes oriented south-southwest was lower than from boxes facing other directions). There was no such relationship for Blue or Great Tits.

Conclusion Nestbox orientation can be an important influence on occupation and breeding success, but this differed between species. Intriguingly, although the directionality reduced nestbox occupation (Great Tit) and breeding success (Pied Flycatcher) was the same (south-southwest), there was a disparity in the influence of orientation for Great Tit (orientation influenced the frequency of occupation but not success) and Pied Flycatcher (orientation did not influence occupation but did affect success). We discuss these disparities, considering the possible influences of mating strategy, breeding phenology, nestbox microclimate and offspring quality.     

Using prey availability to evaluate Lower Colorado River riparian restoration

The Lower Colorado River Multi‐species Conservation Program (MSCP) is charged with restoring habitat for 26 species such as the southwestern willow flycatcher (Empidonax traillii extimus) impacted by water development projects on the river. As of 2015, the MSCP had spent $200 million to create 1,200 ha of habitat at nine sites, but the benefits to these insectivorous birds and other target species have not been quantified. Many MSCP projects emphasized riparian plantings of willow (Salix exigua, Salix gooddingii) and cottonwood (Populus fremontii) on high terraces disconnected from the river. We documented prey availability for insectivores in constructed habitats as an indicator of restoration effectiveness. Using sticky traps as a proxy to estimate aerial insect flux, we found the number of aquatic insects, proportion of aquatic insects, total number of insects, and number of insect orders were all significantly lower in MSCP plantation sites than at the river's edge. Riparian restoration sites over 100 m from the river had only 4% of the aquatic insects, 20% of the total insects, and only half as many insect orders as sites adjacent to the river. Thus, food availability and overall habitat quality for insectivores are likely low in restoration sites that are distant from the river.     

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

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Structural and functional responses of invertebrate communities to climate change and flow regulation in alpine catchments

Understanding and predicting how biological communities respond to climate change is critical for assessing biodiversity vulnerability and guiding conservation efforts. Glacier‐ and snow‐fed rivers are one of the most sensitive ecosystems to climate change, and can provide early warning of wider‐scale changes. These rivers are frequently used for hydropower production but there is minimal understanding of how biological communities are influenced by climate change in a context of flow regulation. This study sheds light on this issue by disentangling structural (water temperature preference, taxonomic composition, alpha, beta and gamma diversities) and functional (functional traits, diversity, richness, evenness, dispersion and redundancy) effects of climate change in interaction with flow regulation in the Alps. For this, we compared environmental and aquatic invertebrate data collected in the 1970s and 2010s in regulated and unregulated alpine catchments. We hypothesized a replacement of cold‐adapted species by warming‐tolerant ones, high temporal and spatial turnover in taxa and trait composition, along with reduced taxonomic and functional diversities in consequence of climate change. We expected communities in regulated rivers to respond more drastically due to additive or synergistic effects between flow regulation and climate change. We found divergent structural but convergent functional responses between free‐flowing and regulated catchments. Although cold‐adapted taxa decreased in both of them, greater colonization and spread of thermophilic species was found in the free‐flowing one, resulting in higher spatial and temporal turnover. Since the 1970s, taxonomic diversity increased in the free flowing but decreased in the regulated catchment due to biotic homogenization. Colonization by taxa with new functional strategies (i.e. multivoltine taxa with small body size, resistance forms, aerial dispersion and reproduction by clutches) increased functional diversity but decreased functional redundancy through time. These functional changes could jeopardize the ability of aquatic communities facing intensification of ongoing climate change or new anthropogenic disturbances.     

Effects of forest disturbances on aquatic insect assemblages

Natural and human‐made disasters such as floods and logging occur in and around rivers. Stream‐dwelling aquatic insects respond to these disturbances in various ways. Primary consumers among them rely greatly on algae and leaf litter from riparian vegetation as food. Therefore, once a disturbance such as a flood has occurred, insects may find it difficult to find food in a stream, and the aquatic insect assemblage can be impacted greatly as a result. Disturbances in riparian areas also increase fine sediment loads into streams, damaging habitat and altering the aquatic insect assemblage. Deforestation impacts not only terrestrial but also aquatic animals. In this review paper, aquatic insect assemblages are assessed according to alterations in land use in and around streams. Following this paper, it is expected that clarification of aquatic insect fauna and their life cycles will progress and that the distribution and habitat use of aquatic insects will be afforded greater attention in forest management.     

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.     

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

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