Emerging aquatic insects as predators in terrestrial systems across a gradient of stream temperature in North and South America

1. Empirical and theoretical research over the past decade has demonstrated the widespread importance of aquatic subsidies to terrestrial food webs. In particular, adult aquatic insects that emerge from streams and lakes are prey for terrestrial predators. While variation in the magnitude of this subsidy is clearly important, the potential top‐down effects of the predatory adults of some aquatic insects in terrestrial food webs are largely unknown. 2. I used published data on benthic insect density (as a proxy for emergence) in North and South America to explore how the proportion of benthic insects that are predatory as adults varies across a gradient of mean annual stream temperature. 3. The proportion of benthic insects that are predatory as adults varied widely across sites (0–12% by abundance; 0–86% by biomass). There was a positive relationship between mean annual stream temperature and the proportion of predatory adults across all sites, driven largely by the greater abundance/biomass of predatory taxa (e.g. odonates), relative to non‐predators (e.g. midges, mayflies, caddisflies), in tropical than in temperate streams. 4. The ‘trophic structure’ (i.e. the proportion of predators) of emerging adult aquatic insects is an understudied source of variation in aquatic–terrestrial interactions. Incorporation of trophic structure in future studies is needed to understand how future modification of fresh waters may affect adjacent terrestrial food webs through both bottom‐up and top‐down effects.     

Seasonal and spatial variation in zooplankton community structure and their relation to possible controlling variables in Lake Okeechobee

1. We examined the behavioural response of stream macroinvertebrates to real and simulated predatory activity by a river bird, the Eurasian dipper, Cinclus cinclus L.
2. In the field, we assessed whether invertebrate drift changed in response to live dippers in enclosures; we found no effects on the drift of any of the five families for which individual analyses were possible, either because it was infrequent, or involved distances too short to be detected (< 0.5 m).
3. In a laboratory stream, we observed prey during encounters with a model dipper which simulated flight, swimming, bill contact with the prey, and stone turning. Invertebrate families varied in their response. Simuliids and hydropsychid caddis lacked effective escape behaviour, consistent with heavy losses through predation by dippers in the wild. Other families either drifted (Baetidae, Gammaridae) or moved away (Heptageniidae, Ephemerellidae, Leuctridae, Perlidae) from the model dipper, but responded only to bill contact or simulated stone turning. Such delayed responses would not protect individuals directly targeted by foraging dippers and partly explain the lack of detectable effects by dippers on drift in the field.
4. We suggest why invertebrates do not show more marked escape responses to this important predator.     

Seasonal and spatial variation in zooplankton community structure and their relation to possible controlling variables in Lake Okeechobee

1. We examined the behavioural response of stream macroinvertebrates to real and simulated predatory activity by a river bird, the Eurasian dipper, Cinclus cinclus L.
2. In the field, we assessed whether invertebrate drift changed in response to live dippers in enclosures; we found no effects on the drift of any of the five families for which individual analyses were possible, either because it was infrequent, or involved distances too short to be detected (< 0.5 m).
3. In a laboratory stream, we observed prey during encounters with a model dipper which simulated flight, swimming, bill contact with the prey, and stone turning. Invertebrate families varied in their response. Simuliids and hydropsychid caddis lacked effective escape behaviour, consistent with heavy losses through predation by dippers in the wild. Other families either drifted (Baetidae, Gammaridae) or moved away (Heptageniidae, Ephemerellidae, Leuctridae, Perlidae) from the model dipper, but responded only to bill contact or simulated stone turning. Such delayed responses would not protect individuals directly targeted by foraging dippers and partly explain the lack of detectable effects by dippers on drift in the field.
4. We suggest why invertebrates do not show more marked escape responses to this important predator.     

The potential influence of Atlantic salmon Salmo salar and brown trout Salmo trutta on density and breeding of the white‐throated dipper Cinclus cinclus

Interactions between birds and fish are often overlooked in aquatic ecosystems. We studied the influence of Atlantic salmon and brown trout on the breeding population size and reproductive output of the white‐throated dipper in a Norwegian river. Acidic precipitation led to the extinction of salmon, but salmon recolonized after liming was initiated in 1991. We compared the dipper population size and reproductive output before (1978–1992) and after (1993–2014) salmon recolonization. Despite a rapid and substantial increase in juvenile salmon, the breeding dipper population size and reproductive output were not influenced by juvenile salmon, trout, or total salmonid density. This might be due to different feeding strategies in salmonids and dippers, where salmonids are mainly feeding on drift, while the dipper is a benthic feeder. The correlation between the size of the dipper population upstream and downstream of a salmonid migratory barrier was similar before and after recolonization, indicating that the downstream territories were not less attractive after the recolonization of salmon. Upstream dipper breeding success rates declined before the recolonization event and increased after, indicating improved water quality due to liming, and increasing invertebrate prey abundances and biodiversity. Surprisingly, upstream the migratory barrier, juvenile trout had a weak positive effect on the dipper population size, indicating that dippers may prey upon small trout. It is possible that wider downstream reaches might have higher abundances of alternative food, rending juvenile trout unimportant as prey. Abiotic factors such as winter temperatures and acidic precipitation with subsequent liming, potentially mediated by prey abundance, seem to play the most important role in the life history of the dipper.     

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.     

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.     

The distribution of dippers, Cinclus cinclus (L.), in the acid‐sensitive region of Wales 1984–95

1. Dippers were surveyed along seventy-four acid-sensitive streams in upland Wales in 1984 and again in 1995. At forty-eight of the sites, and in an additional twenty-three to twenty-nine adjacent catchments, changes in acid-base status and macroinvertebrate communities were assessed. River habitat surveys (RHS) and GIS (Geographical Information System) provided information on habitat features. 2. pH means across all the sites were higher on average by 0.12 pH units in 1995 than 1984, accompanied by significantly increased total hardness and reduced sulphate. However, geometric mean aluminium concentrations increased significantly, while the abundances of important dipper prey either remained constant (Ephemeroptera) or declined (Trichoptera; Plecoptera). 3. As in 1984, dipper distribution in 1995 was related to acid-base status: aluminium concentrations were significantly higher, and pH significantly lower, at sites where dippers were absent. In both surveys, there were significantly more bankside broadleaves and fewer conifers where dippers were present. 4. Although dippers occurred in 1995 on a similar number of streams as in 1984, there were both gains and losses, and an overall significant reduction in the number of visits on which birds were recorded (= registrations) per survey reach. Gains and losses were not related to habitat structure or acid-base status, and might be stochastic. Mean aluminium concentrations increased more (P < 0.06) at sites where dipper registrations fell, than where they increased, but changes were large enough to explain altered occupancy at only five sites. Plecopteran abundances declined most at sites losing birds. 5. We conclude that recovery from acidification has not yet been large or sustained enough to allow widescale increase in the Welsh dipper population, and continued decline cannot be excluded. There is a need for better understanding of how recovery processes will permeate foodwebs to reach top predators in acidified streams, and of the geographical scale of recovery required to increase populations in dispersed organisms such as birds. Such uncertainties, together with the mismatch in trends revealed by stream chemistry and dippers, illustrate reasons why chemical data alone are inadequate for the assessment of changing river quality.     

Cross‐ecosystem disturbance influences on the life history and population size structure of a riparian predator

Cross‐ecosystem transfers of resources could alter the life history traits of consumers in adjacent systems by changing the nature and availability of prey. However, large‐scale influences, such as natural disturbances, that control the magnitude of prey subsidies are likely to modify these effects. To investigate impacts of cross‐ecosystem subsidies on the life history traits of a riparian predator we measured the size, sex and condition of riparian fishing spiders (Dolomedes aquaticus) across a gradient of flooding frequency and intensity. These spiders rely on adult aquatic insects for a large proportion of their diet and previous research demonstrated that increased flooding decreased the abundance of aquatic insect prey. In this study, laboratory experiments indicated that increased prey availability hastened the first moult of the spiders after winter and decreased the propensity for cannibalistic interactions of individuals of the same size. However, despite the likely positive influences of increased food supply, in the field the highest abundance and proportion of large, potentially reproductive females occurred at the most flood‐prone rivers, where aquatic prey availability was the lowest. It is likely that other factors modified by the disturbance regime, such as habitat availability, flood‐related mortality and intra‐specific interaction rates, altered the influence of cross‐ecosystem subsidies on the life history traits of these spiders. Thus, our results indicate that disturbance‐related effects can flow across ecosystem boundaries and alter the life history traits of predators relying on allochthonous resources.     

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.     

Effects of prey dispersal on predator–prey interactions in streams

1. We studied the effect of mesh size (6 and 3 mm) on interactions between brown trout ( Salmo trutta ) and benthic invertebrates in enclosures placed in a stream in southern Sweden. We also compared how different prey exchange rates affected interactions between trout and invertebrates.
2. Trout had strong impacts on some benthic taxa, and different mesh sizes produced different patterns. Trout affected the abundance of 10 of the 21 taxa examined, six in enclosures with 3 mm mesh and six in enclosures with 6 mm mesh. The abundance of nine of the prey taxa was lower in the presence of trout, only leptocerids were more numerous in the presence of trout.
3. Our measurements of prey immigration/emigration, together with trout diet data, suggest that direct consumption by trout, rather than avoidance behaviour by prey, explains most decreases in prey abundance. There was avoidance behaviour by only two of the twenty-one prey taxa, with trout inducing emigration of the mayflies Baetis rhodani and Paraleptophlebia sp.
4. Trout indirectly increased periphyton biomass in both 3 and 6 mm enclosures. The effect of trout on periphyton was probably due to strong effects of trout on the grazer, Baetis rhodani , Heptagenia sp. and Paralepthoplebia sp.
5. Our results suggest that mesh size, through its effects on exchange rates of prey, may affect interactions between predators and prey in running waters, but that the effects of dispersal and predation on invertebrates are taxon specific.     

Response of American dippers (Cinclus mexicanus) to variation in stream water quality

1. Semi‐aquatic birds may be sensitive to altered water quality. While avian species are not used in the bioassessment of streams, they may complement the more common use of benthic macroinvertebrates and fish. We estimated the extent to which water quality can predict attributes of the populations of one common semi‐aquatic bird, the American dipper (Cinclus mexicanus). 2. First, we estimated dipper presence/absence in relation to water quality as measured by a multimetric assessment index and individual bioassessment metrics. Second, we estimated dipper territory area and reproductive success in response to variation in water quality. We studied the diet, territory area and fecundity of dippers and sampled benthic macroinvertebrates, water chemistry and physical variables at 32 sites with and 17 sites without nesting dippers. 3. Dipper presence was only weakly related to chemical, physical and commonly recorded bioassessment metrics such as per cent Ephemeroptera, Plecoptera and Trichoptera (%EPT). Dippers were strongly related to the abundance of their common prey, Drunella and Heptageniidae, which are only a small component of the commonly recorded bioassessment metrics. The variances in territory area and reproductive success were weakly predicted by water quality variables. 4. Dipper presence reflected disturbance as measured by their common prey, showing that lower abundance of these stream invertebrates affected this semi‐aquatic bird. We suggest dipper presence/absence might be used in multimetric indices of biotic integrity for the bioassessment of streams.     

Increases in disturbance and reductions in habitat size interact to suppress predator body size

Food webs are strongly size‐structured so will be vulnerable to changes in environmental factors that affect large predators. However, mechanistic understanding of environmental controls of top predator size is poorly developed. We used streams to investigate how predator body size is altered by three fundamental climate change stressors: reductions in habitat size, increases in disturbance and warmer temperatures. Using new survey data from 74 streams, we showed that habitat size and disturbance were the most important stressors influencing predator body size. A synergistic interaction between that habitat size and disturbance due to flooding meant the sizes of predatory fishes peaked in large, benign habitats and their body size decreased as habitats became either smaller or harsher. These patterns were supported by experiments indicating that habitat‐size reductions and increased flood disturbance decreased both the abundance and biomass of large predators. This research indicates that interacting climate change stressors can influence predator body size, resulting in smaller predators than would be predicted from examining an environmental factor in isolation. Thus, climate‐induced changes to key interacting environmental factors are likely to have synergistic impacts on predator body size which, because of their influence on the strength of biological interactions, will have far‐reaching effects on food‐web responses to global environmental change.     

Ecological effects of invasive alien species on native communities,with particular emphasis on the interactions between aphids and ladybirds

The ecological effects of introduced species on native organisms can sometimes, but not always be significant. The risks associated with invasive alien pests are difficult to quantify. This paper concentrates on the ecological effects of invasive insect predators that feed on pest insects, because the former may potentially affect the biological control of the latter. The literature indicates that invasive predatory insects generally are resistant to changes in environmental conditions, long-lived and voracious with a high reproductive rate, high dispersal ability, able to spread very rapidly across landscapes and exhibit phenotypic plasticity. Their colonization of patches of prey may induce native predators to leave, but the evidence that invaders negatively affect the abundance of the native species is scarce and not persuasive. Insect predators do not substantially affect the abundance of their prey, if the ratio of generation time of the predator to that of the prey is large (the generation time ratio hypothesis), therefore the effect of an invasion by long-lived alien predators on systems consisting of long-lived native predators and short-lived prey on the abundance of the prey is hard to detect.     

Predictable changes in trophic community structure along a spatial disturbance gradient in streams

1. We assessed insect and resource standing stocks along a spatial gradient of flood disturbance in 19 sub-alpine Swedish streams to test the prediction that change in trophic structure arises from the joint action of disturbance, which affect basal resources, and resource-control, which ties the response of the consumers to the response of the resources.
2. Trophic structure, quantified as scores of non-metric multidimensional scaling based on the biomass of insect trophic groups, changed predictably along the disturbance gradient. In early summer, predators and algae feeders decreased relative to suspension feeders with increasing disturbance; in autumn, algae feeders decreased relative to leaf feeders with increasing disturbance.
3. Across the disturbance gradient, the biomass of algae-, deposit- and leaf-feeders was principally controlled by the availability of the respective resource (algae, fine detritus and coarse detritus), while disturbance only had subsidiary effects on algae feeders in early summer.
4. Overall, patterns in trophic-group biomass along the disturbance gradient were more likely to reflect indirect effects of disturbance via impact on the resources, which reverberated to the consumers because of resource-control, rather than direct effects. In contrast with the view that stream communities are the result of stochastic colonization following disturbance events, in the study streams the trophic structure of insect assemblages is predictable and partly organized by resource-control across a broad range of disturbance conditions.     

Evaluating the effects of trophic complexity on a keystone predator by disassembling a partial intraguild predation food web

1. Many taxa can be found in food webs that differ in trophic complexity, but it is unclear how trophic complexity affects the performance of particular taxa. In pond food webs, larvae of the salamander Ambystoma opacum occupy the intermediate predator trophic position in a partial intraguild predation (IGP) food web and can function as keystone predators. Larval A. opacum are also found in simpler food webs lacking either top predators or shared prey. 2. We conducted an experiment where a partial IGP food web was simplified, and we measured the growth and survival of larval A. opacum in each set of food webs. Partial IGP food webs that had either a low abundance or high abundance of total prey were also simplified by independently removing top predators and/or shared prey. 3. Removing top predators always increased A. opacum survival, but removal of shared prey had no effect on A. opacum survival, regardless of total prey abundance. 4. Surprisingly, food web simplification had no effect on the growth of A. opacum when present in food webs with a low abundance of prey but had important effects on A. opacum growth in food webs with a high abundance of prey. Simplifying a partial IGP food web with a high abundance of prey reduced A. opacum growth when either top predators or shared prey were removed from the food web and the loss of top predators and shared prey influenced A. opacum growth in a non-additive fashion. 5. The non-additive response in A. opacum growth appears to be the result of supplemental prey availability augmenting the beneficial effects of top predators. Top predators had a beneficial effect on A. opacum populations by reducing the abundance of A. opacum present and thereby reducing the intensity of intraspecific competition. 6. Our study indicates that the effects of food web simplification on the performance of A. opacum are complex and depend on both how a partial IGP food web is simplified and how abundant prey are in the food web. These findings are important because they demonstrate how trophic complexity can create variation in the performance of intermediate predators that play important roles in temporary pond food webs.     

Changes in aquatic insect emergence in response to whole-lake experimental manipulations of introduced trout

1. Insects emerging from mountain lakes provide an important food source for many terrestrial predators. The amount of insect subsidy that emerges from lakes is influenced by predator composition, but predator effects could be ameliorated by greater habitat complexity. We conducted a replicated whole‐lake experiment to test the effects of introduced fish predators on the abundance and composition of aquatic insects within and emerging from the littoral zone of 16 mountain lakes in the Trinity Alps Wilderness in northwestern California. 2. Study treatments matched the fisheries management options being implemented in California’s wilderness areas: (i) continued stocking with non‐native trout, (ii) suspension of stocking, and (iii) suspension of stocking and removal of fish. We also included four naturally fishless ‘reference’ lakes. We compared abundances and biomass of emerging aquatic insects before treatments were initiated and for 3 years following their establishment. Abundances of benthic insects were also compared in the third year post‐treatment. 3. Trout removal rapidly increased abundances of mayflies, caddisflies, and insect predators, and overall insect biomass emerging from lakes. No significant differences were found between the suspension of stocking lakes and continued stocking lakes. Fish density was a more important predictor of aquatic insect emergence than habitat complexity. 4. Mayfly larvae responded positively to fish removal and caddisfly larvae tended to be more abundant in lakes without fish, but we did not detect effects on abundance of predatory insects. However, we found large insect predators in shallower water in lakes with fish compared to fish removal or fish‐free reference lakes. 5. These results provide insights into the continuing effects of past and current fish stocking practices on the flow of insect prey from mountain lakes into the neighbouring terrestrial environment. We also show that these consequences can rapidly be reversed by removing non‐native fishes.     

The ecology of dippers Cinclus cinclus (L.) in relation to stream acidity in upland Wales: time-activity budgets and energy expenditure

Summary The time-activity budget and energy expenditure of a riverine bird, the dipper Cinclus cinclus, was studied from March 1988 to July 1989, across a range of streams of contrasting acidity in upland Wales. Differences in time-activity budgets of birds on acidic and circumneutral streams were consistent with documented differences in prey availability and diet. Birds spent a significantly greater proportion of their active day foraging, swimming and flying, and less time resting, on acidic streams. Activity measurements varied significantly through the year, through the day, and with river flow. Despite differences in time budgets, mean Daily Energy Expenditure (DEE) on acidic streams was only 4.5–7.0% greater than on circumneutral streams. However, rates of energy gain were greater for dippers on circumneutral streams for every month of the year, a pattern confirmed by differences in body condition. By spending more time feeding, dippers on acidic streams will have less time for other activities such as self-maintenance and predator surveillance; they may also be less able to meet the additional demands accompanying the initiation of breeding. These findings are discussed in relation to the feeding ecology and breeding performance of dippers on streams of contrasting water quality in upland Wales.     

Top-down and bottom-up influences on populations of a stream detritivore

1. Knowledge of the influence of predatory fish in detritus‐based stream food webs is poor. We tested whether larval abundance of the New Zealand leaf‐shredding caddisfly, Zelandopsyche ingens (family Oeconesidae), was affected by the presence of predatory brown trout, Salmo trutta and the abundance of their primary detrital resource (Nothofagus leaves). 2. The density of Z. ingens and the biomass of leaves were determined in seven fishless streams and four trout streams in the Cass region, central South Island, on four occasions spanning 5 years. 3. Physicochemical conditions were similar in trout and fishless streams, but ancova indicated that Z. ingens numbers were positively related to leaf biomass and that caddisfly numbers were significantly greater in fishless streams than trout streams for any given biomass of leaf. The cases of trout stream larvae were also heavier per unit length than those in fishless streams. 4. Our results provide evidence for both top‐down and bottom‐up influences on a detritus‐based stream food web. Although stream detritivores may benefit from a habitat that provides both food and a degree of protection from predators, top‐down effects of predators on detritivore population abundance were still important. Thus, detrital resource availability may determine maximum attainable population size, whereas predation is likely to reduce the population to a level below that.     

Life history variation in a dobsonfly, Protohermes grandis (Megaloptera: Corydalidae): effects of prey availability and temperature

SUMMARY. 1. The life cycle of a predatory insect, Protohermes grandis (Megaloptera: Corydalidae), was compared in four streams in central Japan. The effects of annual temperature regime and prey availability on life history characteristics were also assessed.
2. The larval period was 2 years and small adults emerged in the Morito River, where summer water temperature was high and large prey scarce.
3. In the Nagura River, rich in large prey, the larval period was also 2 years in spite of slightly lower temperature, and the adult size was largest among the streams.
4. In Anado Fork with a low summer temperature, the larval development took 3 years, and large adults emerged. Large prey were abundant in this stream.
5. Seasonal abundance of large prey also affected the time large larvae left the stream to pupate. Larvae emigrated earlier in streams where the density of large prey sharply decreased after spring, than in streams where large prey were available throughout the year.     

Density dependence,prey accessibility and prey depletion by fisheries drive Peruvian seabird population dynamics

In marine ecosystems top predator populations are shaped by environmental factors affecting their prey abundance. Coupling top predators’ population studies with independent records of prey abundance suggests that prey fluctuations affect fecundity parameters and abundance of their predators. However, prey may be abundant but inaccessible to their predators and a major challenge is to determine the relative importance of prey accessibility in shaping seabird populations. In addition, disentangling the effects of prey abundance and accessibility from the effects of prey removal by fisheries, while accounting for density dependence, remains challenging for marine top predators. Here, we investigate how climate, population density, and the accessibility and removal of prey (the Peruvian anchovy Engraulis ringens) by fisheries influence the population dynamics of the largest sedentary seabird community (≈ 4 million individuals belonging to guanay cormorant Phalacrocorax bougainvillii, Peruvian booby Sula variegata and Peruvian pelican Pelecanus thagus) of the northern Humboldt Current System over the past half‐century. Using Gompertz state–space models we found strong evidence for density dependence in abundance for the three seabird species. After accounting for density dependence, sea surface temperature, prey accessibility (defined by the depth of the upper limit of the subsurface oxygen minimum zone) and prey removal by fisheries were retained as the best predictors of annual population size across species. These factors affected seabird abundance the current year and with year lags, suggesting effects on several demographic parameters including breeding propensity and adult survival. These findings highlight the effects of prey accessibility and fishery removals on seabird populations in marine ecosystems. This will help refine management objectives of marine ecosystems in order to ensure sufficient biomass of forage fish to avoid constraining seabird population dynamics, while taking into account of the effects of environmental variability.