The Diet of the Brown Trout Salmo trutta (L.) during the Reproductive Period: Size‐Related and Sexual Effects

We investigated the autumnal diet of the brown trout Salmo trutta, in a Prepyrenean stream (NW Iberian Peninsula) focusing on intraspecific dietary differences related to size and sex. The diet of trout included 18 types of prey, with Plecoptera and Ephemeroptera nymphs and Diptera larvae as the most consumed taxa. Large trout ate larger prey, than did small trout, and also increased the consumption of terrestrial‐surface prey with respect to aquatic‐benthic prey. As terrestrial‐surface preys were larger than aquatic‐benthic prey, the size‐related differences in the diet of trout were related to gape‐limitations. Although male and female trout did not differ in size, we found that males foraged on a more diverse type of prey than females, probably owing to male territoriality during the reproductive period. This study provides new evidence of dietary plasticity in the brown trout and confirms the importance of local dietary studies to better understand factors which drive trophic ecology of predators. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Effects of riparian canopy cover on salmonid diet and prey selectivity in low nutrient streams

This study focuses on stream sections within a relatively low nutrient catchment in south‐east Ireland in an attempt to characterize the probable effects of riparian canopy on salmonid diet and prey selectivity within two size classes of nursery stream. Sampling found that brown trout Salmo trutta diet changed significantly in response to riparian canopy regardless of stream size. The observation that S. trutta within unshaded stream sites did not feed on drifting terrestrial prey items to the same extent as those within shaded streams was not due to a lack of availability of this food source. There was no evidence to suggest that S. trutta selectively choose particular prey items.     

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.     

Diel feeding periodicity, daily ration and prey selection of a riverine population of juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

The diel feeding periodicity, daily ration and prey selection of juvenile chinook salmon, Oncorhynchus tshawytscha , were studied in relation to the available prey. Maximum dry weight of food intake occurred about dawn, when mayflies were the major prey, but the greatest number of freshly eaten prey occurred during the afternoon, when chironomids and terrestrial dipterans predominated. Feeding activity at night was low, with smaller mayflies comprising up to 50% of the prey. During the day the young salmon fed selectively on chironomids and the larger mayflies, while trichopterans and terrestrial taxa were under-represented in the diet. Food consumption over the 24-h period averaged 8.3% of the fish dry body weight. Prey abundance in the drift explained about 50% of the composition of the diet. Although the fish selected larger mayflies, size apparently was not a main criterion for selection because chironomids, although smaller than mayflies, were also frequently eaten. Previous dietary experience of the fish and the diel pattern of prey abundance appear to best explain the selective feeding of juvenile chinook salmon.     

Variation of diet and food consumption of the scaldfish Arnoglossus laterna(Walbaum, 1792)

Although Arnoglossus laterna (Walbaum, 1792) is a common benthic fish in Portugal, several aspects of its feeding ecology remain incomplete. In this study, diet was examined and the food consumption estimated on inshore waters in the central coast of Portugal. The diet of the scaldfish included mainly mysids, amphipods and polychaetes, while the decapod Philocheras bispinosus Hailstone was the most frequent prey. Variation in the diet according to fish size (two size‐classes), sex and spawning/non‐spawning seasons was examined. Significant differences in diet composition were found between seasons, whereby the most frequently consumed prey in the non‐spawning season were crustaceans, while polychaetes and nemerteneans comprised most of its diet in the spawning season. The average value of the vacuity index was very similar between seasons (24.7% non‐spawning season and 25.0% spawning season). Diet overlap between size‐classes and sexes was high, but low between the two seasons. Food consumption of A. laterna was estimated based on diet characterization and evacuation rate. The estimate of the daily food consumption was 1.15 mg dry weight for the non‐spawning season and 1.67 mg dry weight for the spawning season. Significant differences in consumption rates were found between size‐classes and sexes. Larger individuals showed a higher consumption rate than smaller individuals. Females consumed more food than males in the non‐spawning season, while in the spawning season males consumed more than females.     

Hypotheses and trends on how body size affects trophic interactions in a guild of South American killifishes

A chief structuring force in food webs is the hierarchy of trophic interactions, where bigger animals feed on smaller ones. The anatomic and physiological explanations of why body size determines this hierarchy are embodied within the concept of gape limitation. The relaxation of gape limitation and an increase in energetic demands due to predators' larger body size determine the size and diversity of prey species. However, these patterns may be related to further trends in trophic interactions with body size, which have been less considered. Specifically, the passive incorporation of prey should involve a nested distribution of prey among predator size classes. However, predators avoid smaller resources because of their low energy return, with a clumped distribution of prey potentially generating modular organization with qualitative changes in prey identity (e.g. zooplankton, macroinvertebrates and fishes). Finally, size‐mediated interactions (such as direct and indirect competition) may cause predators of similar body size to differentiate among prey organisms, resulting in a checkerboard distribution of prey identity. Consequently, nestedness, modularity and checkerboard distributions of prey among predators of different size classes should form emergent network structures that are directly related to clear ecological mechanisms. We analyse these predictions in a killifish guild, where trends in trophic positions, prey richness, evenness and the number of energy sources systematically scale with body size. We found significant nestedness and segregation in diet among different size classes, supporting the progressive incorporation of prey items coupled with prey differentiation among similar classes. However, we also detected an ‘anti‐modular’ trend, which contradicts theoretical expectations and previous results. We hypothesize that this anti‐modularity is determined by the high biodiversity of the system and the continuous representation of prey size classes. These results reinforce the concept of size‐mediated interactions and its connection with community biodiversity as a main structuring force of food webs.     

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.     

Feeding ecology of the Iberian cyprinid Barbus bocagei Steindachner, 1865 in a lowland river

Patterns of food resource utilization by Barbus bocagei in a lowland river in the central part of the Iberian Peninsula are presented and discussed. Focal points include feeding activity, diet composition and seasonal and ontogenetic diet shifts. Striking features of overall feeding pattern of barbel are its continuous feeding activity, bottom habits, high consumption of detritus and dipteran larvae and narrow diet breadth. Seasonal changes in food resource availability elicited changes in food utilization patterns, revealing a trend to more generalist feeding during spring. Barbel increase consumption of plant material and large prey during ontogeny, but overlap in prey resource utilization between size classes is high. The versatility of barbel foraging behaviour may be an important feature in its exploitation of Iberian lotic systems.     

Feeding ecology and prey preferences of a piscivorous fish in the Lagoa do Peixe National Park, a Biosphere Reserve in Southern Brazil

We investigated the diet, feeding strategy, size-related dietary shifts and prey preferences of South American Hoplias aff. malabaricus in an internationally recognized but poorly investigated Biosphere Reserve in southern Brazil. Fish were caught between April 2008 and March 2009 using a variety of fishing gear. The analysis of 113 individuals revealed a diet essentially composed of fish (16 species), particularly characid species (9). The diet became more diverse and contained larger fish prey with increasing predator size. Feeding strategy analysis revealed a clear specialization towards the consumption of fish. However, individuals did not prey upon particular prey species, instead opportunistically consuming many different fish species, which could be a strategy to avoid intraspecific competition. Characid species were the most important prey, followed by poecillids. A multi-gear sampling of the ichthyofauna revealed that these prey species were the most abundant (Characidae: 61.3%, Poeciliidae 18.8%) of the 14 fish families occurring at the study site, suggesting that the predator exploits the most abundant fish resources available rather than the rarer fish prey. These findings suggest that potential top-down controls exerted by H. aff. malabaricus in this system follow specific food web pathways that seem to be mediated by the abundance of prey resources.     

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

1. Aquatic resource fluxes from streams can provide significant subsidies for riparian consumers. Because aquatic resource fluxes can be highly variable in space and time, the subsidy efficiency (i.e. transfer to the recipient food web) is controlled by the short‐term aggregative response of riparian consumers. 2. Field manipulations of stream‐derived invertebrate prey subsidies were used to examine specific aggregative responses of ground‐dwelling arthropods to riverine subsidy pulses in a braided‐river (Tagliamento River, NE Italy). Subsidy manipulation comprised short‐term reductions of natural stream‐derived subsidies and increased subsidies of stream‐derived invertebrate prey during four seasons. 3. We hypothesised that specific aggregative responses of riparian arthropods depend on their specialisation on aquatic insects which was inferred from stable isotope analysis. Natural riverine subsidy sources including aquatic insect emergence and surface‐drifting organisms were quantified. 4. Arthropods responded significantly with a reduction in abundance by 51%, at reduced subsidies and an increase by 110% at increased subsidies, when averaged over all seasons. Different arthropod taxa responded differently to subsidy manipulations in relation to their specialisation on aquatic subsidies: ground beetles with a diet consisting predominantly of aquatic insects responded only to subsidy reductions, indicating that their local abundance was not limited by natural stream‐derived subsidies; lycosid spiders with a partly aquatic diet showed no significant response; and ants, although relying on a terrestrial diet, responded positively to added stream‐derived invertebrate prey, indicating that stranding of surface‐drifting terrestrial invertebrates represented an important subsidy pathway. 5. Ground beetles and lycosid spiders were seasonally separated in their use of aquatic subsidies. Results indicate that the life‐history characteristics of riparian consumers can control the subsidy efficiency for the recipient community. By the effective uptake of pulsed riverine‐derived subsidies, riparian arthropods can enhance the transfer of riverine food sources to the riparian food web.     

Breeding Yellow-legged Gulls increase consumption of terrestrial prey after landfill closure

The ecology of opportunistic foragers can be highly dependent on anthropogenic food sources, such as landfills, resulting in changes in several ecological and demographic aspects. The total closure of several landfill sites and the use of deterrence systems to prevent access to the remaining open landfill sites in a region in the northern Iberian Peninsula provided an excellent opportunity to evaluate the consequences of landfills on the trophic ecology of an opportunistic forager, the Yellow-legged Gull Larus michahellis, using these resources. Based on stable isotope analyses, we used mixing models to estimate changes in diet before and after the closure of the majority of landfills in the breeding and the non-breeding season. We found a decrease in the consumption of food from landfills after their closure, which was accompanied by an increase in feeding on terrestrial prey (mostly earthworms), but only in the breeding season. Interestingly, we observed no increase in marine prey consumption after the landfill closures. In winter there was a decrease in terrestrial prey consumption, whereas the consumption of marine and, despite their reduce availability, landfill resources increased. Thus, and unlike when all the landfills were open, we detected significant trophic differences between breeding and non-breeding seasons. Additionally, we found significant trophic differences among colonies that were quite close to each other, but not between breeding adults and chicks. In conclusion, landfill closure or the use of deterrence systems had a direct impact on the trophic ecology of Yellow-legged Gulls; loss of refuse was mainly compensated for by prey of terrestrial origin when breeding, but not in winter. Thus, this species may experience foraging constraints in winter with potential effects on other life-history aspects including their dispersal, breeding and survival that needs further evaluation.     

Dietary changes in predators and scavengers in a nocturnally illuminated riparian ecosystem

Aquatic and terrestrial ecosystems are linked by fluxes of carbon and nutrients in riparian areas. Processes that alter these fluxes may therefore change the diet and composition of consumer communities. We used stable carbon isotope (δ¹³C) analyses to test whether the increased abundance of aquatic prey observed in a previous study led to a dietary shift in riparian consumers in areas illuminated by artificial light at night (ALAN). We measured the contribution of aquatic‐derived carbon to diets in riparian arthropods in experimentally lit and unlit sites along an agricultural drainage ditch in northern Germany. The δ¹³C signature of the spider Pachygnatha clercki (Tetragnathidae) was 0.7‰ lower in the ALAN‐illuminated site in summer, indicating a greater assimilation of aquatic prey. Bayesian mixing models also supported higher intake of aquatic prey under ALAN in spring (34% versus 21%). In contrast, isotopic signatures for P. clercki (0.3‰) and Pardosa prativaga (0.7‰) indicated a preference for terrestrial prey in the illuminated site in spring. Terrestrial prey intake increased in spring for P. clercki under ALAN (from 70% to 74%) and in spring and autumn for P. prativaga (from 68% to 77% and from 67% to 72%) and Opiliones (from 68% to 72%; 68% to 75%). This was despite most of the available prey (up to 80%) being aquatic in origin. We conclude that ALAN changed the diet of riparian secondary consumers by increasing the density of both aquatic and terrestrial prey. Dietary changes were species‐ and season‐specific, indicating that the effects of ALAN may interact with phenology and feeding strategy. Because streetlights can occur in high density near freshwaters, ALAN may have widespread effects on aquatic–terrestrial ecosystem linkages.     

Partitioning seasonal time: interactions among size, foraging activity and diet in leaf-litter frogs

This study investigates hypotheses about partitioning of food resources among all species and several size classes in an assemblage of diurnal leaf-litter frogs in central Amazonia. All species in this assemblage change the type and size of prey as they grow. An ordination of diet composition was significantly associated with frog size and species-specific behaviour. However, a partial Mantel analysis indicated that species explained about 1.5 times more of the variation in diet overlap between individuals than frog size. Diet and foraging activity are correlated in juveniles, but not in adults, and this result holds whether species are considered as statistically independent observations or whether relationships are analysed using phylogenetically independent contrasts. This study showed that the partitioning of food resources between species changes with the population size structures. Thus, intraspecific and interspecific changes in diet, coupled with different patterns of juvenile recruitment, cause diet segregation among species due to temporal segregation of equivalent size classes. Received: 25 August 1997 / Accepted: 1 April 1998     

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     

Sexual and seasonal variation in the diet and foraging behaviour of a sexually dimorphic carnivore, the honey badger (Mellivora capensis)

The honey badger, or ratel, Mellivora capensis has not been well studied despite its extensive distribution. As part of the first detailed study, visual observations of nine habituated free-living individuals (five females, four males) were used to investigate seasonal, annual and sexual differences in diet and foraging behaviour. Theory predicts that generalist predators 'switch' between alternative prey species depending on which prey species are currently most abundant, and diet breadth expands in response to decreased availability of preferred food types. There were significant seasonal differences in the consumption of eight prey categories related to changes in prey availability but no seasonal differences in food intake per kg of body mass. As predicted, the cold-dry season diet was characterized by low species richness and low foraging yield but high dietary diversity, while the reverse was true in the hot-dry and hot-wet seasons. In accordance with these predictions, results suggest that the honey badger maintains its intake level by food switching and by varying dietary breadth. Despite marked sexual size dimorphism, male and female honey badgers showed no intersexual differences in prey size, digging success, daily food intake per unit body weight or foraging behaviour. Results do not support the hypothesis that size dimorphism is primarily an adaptation to reduce intersexual competition for food.     

Effects of woody debris and the supply of terrestrial invertebrates on the diet and growth of brown trout (Salmo trutta) in a boreal stream

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Comparisons of the food niches of three native and two introduced fish species in an Australian river

Synopsis The breadth, correspondence and overlap of the diets of the small and large size classes of the three native species (Galaxias occidentalis, Bostockia porosa and Edelia vittata) and two introduced species (Gambusia holbrooki and Perca fluviatilis) of fishes found in the shallows of the main channel and in the tributary creeks of a south-western Australian river have been compared in each season. Classification and ordination were used to examine the overall interrelationships of the diets across species, size groups, seasons and the location where the fish were caught (channel or creek). The smaller fish had a narrower dietary breadth than larger fish in the spring and summer, presumably reflecting the size limit imposed on prey size by their possession of a relatively small mouth in these seasons. Intraspecific dietary overlap between large and small size classes was usually high in G. occidentalis, but generally low in G. holbrooki and P. fluviatilis, and also in B. porosa and E. vittata when the difference between the lengths of the two size groups was greatest. Dietary overlap was least in autumn when the main prey taxa were most abundant. During winter, the diets of the three native species in the tributaries converged, probably reflecting a relatively low faunal diversity in these highly seasonal water bodies. The only relatively consistent interspecific overlap in diet was between B. porosa and E. vittata. Classification and ordination of the dietary samples separated G. occidentalis (which fed extensively on terrestrial organisms from the water surface) from the smaller P. fluviatilis (that concentrated on copepods in the plankton) and from B. porosa and E. vittata (which ingested primarily benthic organisms). Furthermore, B. porosa tended to ingest larger prey taxa than E. vittata. The diet of Gambusia holbrooki is sometimes dominated by terrestrial insects and at other times by benthic organisms, demonstrating that this species is an opportunistic carnivore. It is concluded that food partitioning by the three native and two introduced fish species in the Collie River is likely to be one of the principal factors facilitating the coexistence of substantial populations of these species in this system.     

Dietary and Temporal Niche Differentiation in Tropical Ants—Can They Explain Local Ant Coexistence?

How species with similar ecological requirements avoid competitive exclusion remains contentious, especially in the species‐rich tropics. Niche differentiation has been proposed as a major mechanism for species coexistence. However, different niche dimensions must be studied simultaneously to assess their combined effects on diversity and composition of a community. In most terrestrial ecosystems, ants are among the most abundant and ubiquitous animals. Since they display direct, aggressive competition and often competitively displace subordinate species from resources, niche differentiation may be especially relevant among ants. We studied temporal and trophic niche differentiation in a ground ant community in a forest fragment in French Guiana. Different baits were presented during day and night to assess the temporal and dietary niches of the local species. They represented natural food resources such as sugars, carrion, excrements, seeds, and live prey. In addition, pitfalls provided a background measure of ant diversity. The communities attracted to the different baits significantly differed from each other, and even less attractive baits yielded additional species. We detected species specialized on living grasshoppers, sucrose, seeds, or dead insects. Community‐level differences between day and night were larger than those between baits, and many species were temporally specialized. In contrast to commonness, foraging efficiency of species was correlated to food specialization. We conclude that many ant species occupy different temporal or dietary niches. However, for many generalized species, the dietary, and temporal niche differentiation brought forward through our sampling effort, cannot alone explain their coexistence.     

Food web structure and body size: trophic position and resource acquisition

Ulrich Brose Body size is recognized as an important determinant of trophic structure as it affects individual energetic demands, population density, and the interaction between potential prey and predators. However, its relationship with trophic position remains unclear. It has been hypothesized that a positive relationship between body size and trophic position would be associated to some particular trophic structures, which would allow larger organisms to satisfy their energetic demand and sustain viable population sizes at higher trophic positions, where fewer resources are available. To test this hypothesis, we analyzed the diet of 619 killifishes from four species (Austrolebias cheradophilus, A. luteoflammulatus, A. viarius and Cynopoecilus melanotaenia), collected in temporary ponds occurring in the grasslands of Rocha, Uruguay. Trophic position, diet richness, number of energy sources, and evenness were estimated for 20 size classes, formed by consecutive groups of 31 individuals. Gape limitation and preference for the larger available prey were evaluated as explanations for observed patterns with an individual based model (IBM). In agreement with the hypothesis, killifishes presented a strong positive relationship between trophic position and body size (R²=0.86), associated with a trophic structure that could allow larger organisms to have access to more energy from the environment. This was reflected in a positive relationship between body size and 1) prey richness, 2) number of basal energy sources (i.e. plants, detritus, phytoplankton and terrestrial prey), and 3) evenness in prey use. IBM results showed that changes in trophic structure with body size are well explained by gape limitation, but not by size preferences. Our results suggest that the fulfilment of the greater energetic demands of larger organism will depend on community diversity, which typically increases with ecosystem size, indicating a novel connection between area, diversity, body size, and food chain length.     

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.