Marine resource flows to terrestrial arthropod predators on a temperate island: the role of subsidies between systems of similar productivity

Marine-terrestrial resource flows can subsidies recipient consumers at various trophic levels. Theory suggests that the importance of such spatial subsidies depends on the productivity gradient between adjacent systems; however, the empirical data required to test this assumption are scarce. Most studies of marine-terrestrial subsidies have been performed in arid coastal habitats of low productivity surrounded by productive ocean waters. We examined the importance of marine resource inputs for terrestrial consumers on a temperate, productive forest island surrounded by a marine system of similar productivity. The importance of marine resources for the dominant arthropod consumers was estimated using stable isotopes and linear mixing models. We compared isotopic signatures of spiders and ants captured along a gradient from shore to inland to estimate how far marine-derived energy penetrates the island. We evaluated the distribution of ground-dwelling arthropods using pitfall-trap transects extending from the supratidal-forest boundary to the middle of the island. The contribution of marine-derived energy assimilated by arthropod consumers differed both among taxa and location. Marine-derived resources contributed >80% to the assimilated C of intertidal spiders and 5-10% for spiders at the forest edge and further inland. Ants assimilated 20% of their C from marine-derived resources and this proportion was not affected by distance from shore. Spiders, ants, and all arthropods combined exhibited no spatial aggregation towards the shore. Our results indicate that on temperate islands marine-terrestrial subsidies might be predominantly an edge effect, confined to intertidal consumers. Mobile consumers that opportunistically forage in intertidal habitats play an important role in transferring marine-derived energy further inland. This suggests that the importance of the productivity gradient for spatial subsidies can be modified by the mobility traits of the recipient consumers and their degree of specialization on the interface habitat.     

Shared patterns of species turnover between seaweeds and seed plants break down at increasing distances from the sea

We tested for correlations in the degree of spatial similarity between algal and terrestrial plants communities along 5500 km of temperate Australian coastline and whether the strength of correlation weakens with increasing distance from the coast. We identified strong correlations between macroalgal and terrestrial plant communities within the first 100 km from shore, where the strength of these marine–terrestrial correlations indeed weakens with increasing distance inland. As such, our results suggest that marine‐driven community homogenization processes decompose with increasing distance from the shore toward inland. We speculate that the proximity to the marine environment produces lower levels of community turnover on land, and this effect decreases progressively farther inland. Our analysis suggests underlying ecological and evolutionary processes that give rise to continental‐scale biogeographic influence from sea to land.     

Geographical variation in the use of intertidal rocky shores by the lizard Microlophus atacamensis in relation to changes in terrestrial productivity along the Atacama Desert coast

1. The movement of materials and organisms between ecosystems is a common process in nature. 2. In the present study we investigate the hypothesis that the movement of consumers between ecosystems depends not only on the differences in productivity between ecosystems and prey availability, but also on these animals' biological characteristics. 3. To address this hypothesis we investigated the changes in abundance, habitat utilization and diet of the lizard Microlophus atacamensis along its geographical range on the coast of the Atacama Desert. Within this range, intertidal rocky shore communities do not show important variations in their species composition and abundance, but terrestrial communities show a steep gradient of productivity associated with the increase in rainfall from north to south. 4. Our results show that the use of intertidal habitats and the consumption of intertidal prey by M. atacamensis change within its geographical range: in the North, the species uses intertidal areas and behaves as an herbivore consuming mostly algae, whereas in the South it expends most of the time in terrestrial habitats as a carnivore mainly of arthropods. 5. Our study gives new evidence for cross-ecosystem connections created by consumer movement between habitats of contrasting but variable productivity levels.     

Responses of Dune Plant Communities to Continental Uplift from a Major Earthquake: Sudden Releases from Coastal Squeeze

Vegetated dunes are recognized as important natural barriers that shelter inland ecosystems and coastlines suffering daily erosive impacts of the sea and extreme events, such as tsunamis. However, societal responses to erosion and shoreline retreat often result in man-made coastal defence structures that cover part of the intertidal and upper shore zones causing coastal squeeze and habitat loss, especially for upper shore biota, such as dune plants. Coseismic uplift of up to 2.0 m on the Peninsula de Arauco (South central Chile, ca. 37.5º S) caused by the 2010 Maule earthquake drastically modified the coastal landscape, including major increases in the width of uplifted beaches and the immediate conversion of mid to low sandy intertidal habitat to supralittoral sandy habitat above the reach of average tides and waves. To investigate the early stage responses in species richness, cover and across-shore distribution of the hitherto absent dune plants, we surveyed two formerly intertidal armoured sites and a nearby intertidal unarmoured site on a sandy beach located on the uplifted coast of Llico (Peninsula de Arauco) over two years. Almost 2 years after the 2010 earthquake, dune plants began to recruit, then rapidly grew and produced dune hummocks in the new upper beach habitats created by uplift at the three sites. Initial vegetation responses were very similar among sites. However, over the course of the study, the emerging vegetated dunes of the armoured sites suffered a slowdown in the development of the spatial distribution process, and remained impoverished in species richness and cover compared to the unarmoured site. Our results suggest that when released from the effects of coastal squeeze, vegetated dunes can recover without restoration actions. However, subsequent human activities and management of newly created beach and dune habitats can significantly alter the trajectory of vegetated dune development. Management that integrates the effects of natural and human induced disturbances, and promotes the development of dune vegetation as natural barriers can provide societal and conservation benefits in coastal ecosystems.     

The influence of freshwater-lake subsidies on invertebrates occupying terrestrial vegetation

Studies investigating effects of aquatic-derived resource subsidies have often found large effects on terrestrial systems. Those studies have mostly been performed on effects of subsidies derived from oceanic and riverine systems, and very few have considered effects of subsidies from freshwater lakes. However, since lakes can produce large quantities of emergent aquatic insects that end up on nearby land, it is likely that also freshwater-lake subsidies influence terrestrial systems. We performed sweep-net collections of aquatic and terrestrial invertebrates at varying distances from the shore on vegetation of islands of varying size, in two freshwater lakes in northern Sweden, as well as on the surrounding mainland. We found that the amounts of aquatic insects on terrestrial vegetation decreased with distance from the shore, and that they were the most abundant on small islands, presumably because small islands have a higher perimeter-to-area ratio. Web-building spiders responded positively to the aquatic subsidy by being the most abundant on small islands and by showing a positive relationship with aquatic insect biomass. However, distance from the shore showed no effects on the spiders. Our results strongly support the view that terrestrial systems are subsidized by lakes, and indicate that freshwater-lake subsidies are important for terrestrial invertebrate community structure on adjacent land. Further, our study shows that ecosystems should be treated as interdependent, not as self-contained units, and may as such be important for an increased understanding of the nature and importance of resource flows across ecosystem boundaries.     

Marine resources subsidize insular rodent populations in the Gulf of California,Mexico

Inputs of energy and nutrients from one ecosystem may subsidize consumers in adjacent ones, with significant consequences for local communities and food webs. We used stable isotope and faecal pellet analysis to quantify use of ocean-derived resources by small mammals on islands in the Gulf of California, Mexico. Rodents were live-trapped on grids originating near shore and extending 125-200 m inland to evaluate the extent to which rodents transport marine nutrients inland, and to determine whether marine foods subsidize island populations, permitting higher densities than would be possible based on terrestrial resources alone. Both faeces and stable carbon and nitrogen isotopes revealed that omnivorous mice (Peromyscus maniculatus) consume ocean-derived prey, including littoral and supralittoral invertebrates, and that their diets differed markedly from those of granivorous rodents (Chaetodipus rudinoris). On a small, seabird roosting island, marine prey were important in the diet of mice regardless of their proximity to shore, underscoring the pervasive influence of the ocean on small islands with relatively large coastline area. On a large island, however, consumption of marine foods declined sharply > or =50 m from shore, which suggests that mice are poor conduits of inland movement of energy and nutrients from the sea. Marine resources seemed to act as subsidies for omnivorous rodents: more P. maniculatus were captured near shore than farther inland and there was an inverse relationship between island area and rodent abundance, suggesting that small islands with large amounts of marine inputs support the highest population densities. Patterns of local and island-wide abundance of P. maniculatus are likely the result of several interacting factors, including frustrated dispersal, competition with C. rudinoris, and the absence of predators. We speculate, however, that the availability of marine resources allows P. maniculatus to reach high densities and to persist on small islands in the Gulf despite low and unpredictable terrestrial productivity. Spatial trophic subsidies thus provide a possible mechanistic explanation for the widely reported inverse relationship between population density and island or habitat area.     

Distribution of seagrasses along the Indian coast

Seagrass environments, from the main coast of India, Lakshadweep and Andaman Islands, were surveyed for seagrass and marine algal composition. Extensive seagrass meadows and the maximum number of species (seven genera and 12 species) occurred along the Tamil Nadu coast. Seagrasses were observed from intertidal to subtidal regions down to 8 m depth. Thalassia hemprichii (Ehrenberg) Aschers. and Cymodocea serrulata (R. Brown) Aschers. & Magnus were the dominant seagrasses in the subtidal zones. Halophila beccarii Aschers. was restricted to the intertidal mudflats in association with mangroves. The rich growth of seagrasses along the Tamil Nadu coast and Lakshadweep can be attributed mainly to high salinity, clarity of the water and sandy substratum. One hundred species of marine algae were recorded from the seagrass environments of India.     

The loss of seagrasses in Cockburn Sound, Western Australia. I. The time course and magnitude of seagrass decline in relation to industrial development

The areas of seagrass meadows in Cockburn Sound, a marine embayment in Western Australia, were estimated from historical aerial photographs supplemented by ground surveys, studies on meadows in adjoining areas, and coring for rhizome remains. Ten species of seagrasses with different habitat tolerances are recorded for the area, with Posidonia sinuosa Cambridge et Kuo forming the most extensive meadows. It is estimated that from 1954 to 1978 the meadow area was reduced from some 4200 to 900 ha. Based on measurements of aboveground productivity at several sites, this represents a reduction of leaf detritus production from 23 000 to 4000 t (dry wt.) y⁻¹. The major loss of seagrass occurred during a period of industrial development on the shore, and the discharge of effluents rich in plant nutrients.     

The Ulva connection: marine algae subsidize terrestrial predators in coastal Peru

How can terrestrial animals survive in a desert with scant primary productivity? The Peruvian coastal desert is hyper‐arid, but faces one of the world's most productive marine ecosystems, the Peru–Chile cold current. Given the stark difference in productivity between these adjacent ecosystems, we expected to find strong linkages connecting the terrestrial and marine food web. We investigated how marine resources are incorporated in the diet, and influence the distribution of terrestrial consumers (geckos, scorpions, solifuges and darkling beetles). Stomach contents from geckos, and δ¹³C and δ¹⁵N values of geckos and other terrestrial consumers suggest that marine green algae of the genus Ulva provide energy and nutrients to the terrestrial food web. Isotopic values suggest that amphipods, which feed on stranded Ulva, make marine resources available to terrestrial predators by moving between the intertidal and supratidal zones. The relative contribution of terrestrial and algal carbon sources varied among terrestrial predators, because scorpions assimilated a lower proportion of energy from Ulva than did geckos and solifuges. These δ¹³C patterns reflected differences in the spatial distribution of consumers. Our study supports the idea that in places where ecosystems with contrasting productivity levels are spatially juxtaposed, it is not possible to understand the structure and dynamics of food webs without taking into account the effects of energy and nutrients flowing from adjacent ecosystems. In contrast to other studied systems, especially those in Baja California, our site in Peru receives very little rainfall and the amount of precipitation is not affected by El Niño events. The near absence of rainfall promotes an extreme dependence of terrestrial consumers on marine resources, and causes permanent indirect food‐web effects that are affected by temporal variability in marine productivity, rather than temporal patterns of plant growth.     

Heterozoan carbonate-enriched beach sand and coastal dunes—with particular reference to rhodoliths,Dirk Hartog Island,Shark Bay,Western Australia

This is a preliminary interdisciplinary study on the enrichment of heterozoan carbonates on Dirk Hartog Island, Shark Bay, Western Australia, with particular reference to rhodolith (free-living non-geniculate) coralline algae. The current study aims to investigate the geological impact of shallow-water rhodoliths in Shark Bay, as well as fill critical information gaps on the biogeographical distribution of rhodoliths in Australia. We analyzed the composition of sand from eight sites (totaling 21 beach and sand dune samples) on the eastern (windward) shore of the island, and investigated the origin of the coralline algal grains. Heterozoan carbonates (shell, geniculate coralline algae grains, and rhodolith grains) together comprised 3–84% of the carbonate-enriched beach and dune sand samples. While shell fragments often comprised the highest percentage (up to 73%), rhodolith grains (up to 27%) were found in 12 of 21 samples, with rhodolith grains also occurring in two dune samples. Geologically, the study has shown that rhodoliths and rhodolith beds are important shallow-marine habitats in Shark Bay, with a proven capacity to enrich beach/dune sands in Shark Bay and potentially other areas along the Australian coast. Biogeographically, the study confirmed the presence of a previously undescribed shallow rhodolith bed in Shark Bay (the first bed documented on the Western shore) with the possibility of a third bed near Sandy Point on Dirk Hartog Island. It also confirmed the presence of rhodolith forming Neogoniolithon brassica-florida and Lithophyllum sp. in Shark Bay, and is the first record of Hydrolithon reinboldii rhodoliths in Australia.     

Shellfish subsidies along the Pacific coast of North America

Spatial subsidies are associated with pronounced ecosystem responses, as nutrients cross ecological boundaries and cascade through food webs. While the importance of subsidies is known, the role of shellfish, specifically molluscs, as a marine subsidy has not been formally described. Focusing primarily on the Pacific coast of North America, we identify vectors that transport shellfish-derived nutrients into coastal terrestrial environments, including birds, mammals, and over 13 000 yr of marine resource use by people. Evidence from recipient ecosystems suggests shellfish drastically influence soil chemistry, forest productivity and the diversity of primary producers at the regional and landscape level. Responses in higher trophic levels have not yet been investigated, but given documented responses in lower trophic levels, this may be due to a lack of examination. To determine if the processes we describe within the northeast Pacific are pertinent to coastal environments worldwide, we also explore shellfish subsidies globally, with a specific focus on temperate and tropical islands. As shellfish are not as spatially or temporally constrained as other subsidies, our examination suggests our findings are applicable to many other geographical regions along the marine–terrestrial interface.     

The Impact of Cormorants on Plant–Arthropod Food Webs on Their Nesting Islands

This study investigated the effects of cormorant colonies on plant–arthropod island food webs, the consequences of nutrient-rich runoff on marine communities, and feedback loops from marine to terrestrial ecosystems. Terrestrial plant responses were as expected, with the highest plant biomass on islands with low nest density and the highest nitrogen (N) content on islands with high nest density. In contrast to our hypothesis, we found no uniform density response across guilds. Among herbivores, the variable responses may depend on the relative importance of plant quality or quantity. As expected, nutrient-rich runoff entered water bodies surrounding cormorant nesting islands, but only at high nest density, and increased the density of emerging insects. This created a potential feed-back loop to spiders (major terrestrial predators), where stable isotope analyses suggested great use of chironomids. Contrary to our expectation, this potential feed-back did not result in the highest spider density on islands with a high cormorant nest density. Web spiders showed no changes in density on active cormorant islands, and lycosids were actually less abundant on active cormorant islands compared to reference islands. The variable response of spiders despite increased dipteran densities, and also in other consumer groups, may be due to direct negative effects of cormorants on soil chemistry, vegetation cover, and other density regulating forces (for example, top–down forces) not studied here. This study highlights the importance of including processes in the surrounding marine ecosystem to understand the impacts of seabirds on the food web structures of their nesting islands.     

The importance of marine vs. human-induced subsidies in the maintenance of an expanding mesocarnivore in the arctic tundra

1. Most studies addressing the causes of the recent increases and expansions of mesopredators in many ecosystems have focused on the top-down, releasing effect of extinctions of large apex predators. However, in the case of the northward expansion of the red fox into the arctic tundra, a bottom-up effect of increased resource availability has been proposed, an effect that can counteract prey shortage in the low phase of the multi-annual rodent cycle. Resource subsidies both with marine and with terrestrial origins could potentially be involved. 2. During different phases of a multi-annual rodent cycle, we investigated the seasonal dynamics and spatial pattern of resource use by red foxes across a coast to inland low arctic tundra gradient, Varanger Peninsula, Norway. We employed two complementary methods of diet analyses: stomach contents and stable isotope analysis. 3. We found that inland red foxes primarily subsisted on reindeer carrions during the low phase of a small rodent population cycle. Lemmings became the most important food item towards the peak phase of the rodent cycle, despite being less abundant than sympatric voles. Isotopic signatures of tissue from both predator and prey also revealed that red foxes near the coast used marine-derived subsidies in the winter, but these allochthonous resources did not spillover to adult foxes living beyond 20-25 km from the coast. 4. Although more needs to be learned about the link between increasing primary productivity due to climatic warming and trophic dynamics in tundra ecosystems, we suggest that changes in reindeer management through a bottom-up effect, at least regionally, may have paved the way towards the establishment of a new mesopredator in the tundra biome.     

Stable Isotope Ratios as a Tool for Assessing Changes in Carbon and Nutrient Sources in Antarctic Terrestrial Ecosystems

Samples of an angiosperm species, nine lichen species and a terrestrial alga, were collected from a variety of Antarctic terrestrial habitats, and were analysed for C and N stable isotope composition. Collections were made along natural gradients, the marine gradient, running from the sea coast inland and the moisture gradient, determined by melt water and precipitation runoff, and running towards the sea coast. Considerable variation in stable isotope ratios was found; δ¹³C values ranged between −16 and −32‰ and δ¹⁵N values between −23 and +23‰ The variation in stable carbon isotope ratios could be attributed in part to species specific differences, but differences in water availability also played a role, as was shown for the terrestrial alga Prasiola crispa and the lichen species Usnea antarctica. The differences in the isotope ratios of nitrogen could be retraced to the origin of nitrogen: marine or terrestrial. The nitrogen stable isotope ratios were influenced by both the marine gradient from the sea inland and the melt water and precipitation flow running in the opposite direction, towards the sea. This was shown for the lichen species Turgidosculum complicatulum and the angiosperm species Deschampsia antarctica. The variation in the C and N stable isotope ratios can be used to determine sources and pathways of N and changes in the water availability in Antarctic terrestrial ecosystems. Contrary to earlier reports the use of stable N isotope ratios is possible in this case because of the relative simplicity of the structure of the Antarctic terrestrial ecosystems.     

上川岛潮间带不同生境底栖软体动物物种多样性初步研究

目的:研究上川岛潮间带不同生境底栖软体动物物种多样性。方法:对上川岛潮间带(沙滩、泥沙滩、红树林泥滩、岩石滩) 4种不同生境的7个样方的底栖软体动物进行了生物多样性的初步调查。结果:共鉴定底栖软体动物14科30种,上川岛潮间带底栖软体动物物种丰富度指数为岩石滩(平均为1.784 ) >泥沙滩(平均为1.5 2 1) >红树林泥滩(平均为1.2 35 ) >沙滩(平均为0 .6 92 ) ;多样性指数岩石滩(平均为1.5 5 1) >红树林泥滩(平均为1.30 4 ) >泥沙滩(平均为1.16 2 ) >沙滩(平均为0 .4 0 3)。结论:底栖软体动物的种类及数量分布受底质、浪击等因素的影响。     

Land–Ocean Coupling of Carbon and Nitrogen Fluxes on Sandy Beaches

Rivers link oceans with the land, creating global hot spots of carbon processing in coastal seas. Coastlines around the world are dominated by sandy beaches, but beaches are unusual in that they are thought to rely almost exclusively on marine imports for food. No significant connections to terrestrial production having been demonstrated. By contrast, we isotopically traced carbon and nitrogen pathways leading to clams (Donax deltoides) on beaches. Clams from areas influenced by river plumes had significantly different isotope signatures (δ¹³C: −18.5 to −20.2‰; δ¹⁵N: 8.3–10.0‰) compared with clams remote from plumes (δ¹³C: −17.5 to −19.5‰; δ¹⁵N: 7.6–8.7‰), showing that terrestrial carbon and sewage, both delivered in river plumes, penetrate beach food webs. This is a novel mechanism of trophic subsidy in marine intertidal systems, linking the world’s largest shore ecosystem to continental watersheds. The same clams also carry pollution signatures of sewage discharged into rivers, demonstrating that coastal rivers connect ecosystems in unexpected ways and transfer contaminants across the land–ocean boundary. The links we demonstrate between terrigenous matter and the largest of all marine intertidal ecosystems are significant given the immense social, cultural, and economic values of beaches to humans and the predicted consequences of altered river discharge to coastal seas caused by global climate change.     

Investigations on the feeding habits of the rocky-shore miteHyadesia fusca (Acari: Astigmata: Hyadesiidae): diet range, food preference, food quality, and the implications for distribution patterns

Within the food web of estuarine and marine rocky shore ecosystems phytophagous mites of terrestrial and marine origin constitute an important part as grazers on algae and as a food source for certain arthropods, especially zoophagous mites. This investigation deals with the feeding biology ofHyadesia fusca taking as an example a population located on an artificial rocky shore of the middle Weser estuary in Northern Germany. The species is characterized by a broad diet range; in feeding experiments diatoms, lichens, detritus as well as blue, red and green algae were accepted. Even analyses of faecal pellets produced by field specimen suggest a non-specific feeding habit. However, the influence of certain diets on mortality, offspring number and rearing success showed that the food quality differs significantly. The most suitable food, the UlvaceaeBlidingia, was clearly preferred in a series of pairwise choice tests. These findings correlate with the vertical zonation of the field population i.e.: higher population densities in the vegetation zone dominated byBlidingia. It can be concluded that in addition to abiotic factors food supply could play an important role for distribution patterns of phytophagous mites.     

Sandy beaches at the brink

Sandy beaches line most of the world's oceans and are highly valued by society: more people use sandy beaches than any other type of shore. While the economic and social values of beaches are generally regarded as paramount, sandy shores also have special ecological features and contain a distinctive biodiversity that is generally not recognized. These unique ecosystems are facing escalating anthropogenic pressures, chiefly from rapacious coastal development, direct human uses — mainly associated with recreation — and rising sea levels. Beaches are increasingly becoming trapped in a 'coastal squeeze' between burgeoning human populations from the land and the effects of global climate change from the sea. Society's interventions (e.g. shoreline armouring, beach nourishment) to combat changes in beach environments, such as erosion and shoreline retreat, can result in severe ecological impacts and loss of biodiversity at local scales, but are predicted also to have cumulative large-scale consequences worldwide. Because of the scale of this problem, the continued existence of beaches as functional ecosystems is likely to depend on direct conservation efforts. Conservation, in turn, will have to increasingly draw on a consolidated body of ecological theory for these ecosystems. Although this body of theory has yet to be fully developed, we identify here a number of critical research directions that are required to progress coastal management and conservation of sandy beach ecosystems.     

Lake‐derived midges increase abundance of shoreline terrestrial arthropods via multiple trophic pathways

Aquatic insects link adjacent ecosystems by transporting nutrients, energy, and material as they move from bodies of water into terrestrial habitats. Insects emerging from streams and rivers are known to benefit arthropod predators such as spiders, but their influence may extend to other arthropod feeding groups as well. We conducted a terrestrial arthropod survey at a series of lakes spanning a strong gradient of midge (Chironomidae, Diptera) emergence. These small, short‐lived insects reach high densities in some areas such that their carcasses litter the ground, and serve as a potential resource for non‐predatory arthropods. Our study revealed that arthropod assemblages in areas of high midge density were significantly different from those with few midges, the result of an increase of all taxa rather than changes in taxonomic composition. Eight of nine terrestrial arthropod taxa sampled showed a strong positive response to the presence of midges including detritivores and herbivores in addition to predators. Taxa that could consume living or dead midges directly responded especially strongly to midge gradients. Our results strongly suggest that midges enter the terrestrial arthropod food web through multiple pathways, increasing numbers of a wide range of arthropods. Furthermore, they emphasize the importance of lakes as sources of aquatic insects that significantly alter processes in the neighboring terrestrial environment.     

Seasonally varying marine influences on the coastal ecosystem detected through molecular gut analysis

Terrestrial predators on marine shores benefit from the inflow of organisms and matter from the marine ecosystem, often causing very high predator densities and indirectly affecting the abundance of other prey species on shores. This indirect effect may be particularly strong if predators shift diets between seasons. We therefore quantified the seasonal variation in diet of two wolf spider species that dominate the shoreline predator community, using molecular gut content analyses with general primers to detect the full prey range. Across the season, spider diets changed, with predominantly terrestrial prey from May until July and predominantly marine prey (mainly chironomids) from August until October. This pattern coincided with a change in the spider age and size structure, and prey abundance data and resource selection analyses suggest that the higher consumption of chironomids during autumn is due to an ontogenetic diet shift rather than to variation in prey abundance. The analyses suggested that small dipterans with a weak flight capacity, such as Chironomidae, Sphaeroceridae, Scatopsidae and Ephydridae, were overrepresented in the gut of small juvenile spiders during autumn, whereas larger, more robust prey, such as Lepidoptera, Anthomyidae and Dolichopodidae, were overrepresented in the diet of adult spiders during spring. The effect of the inflow may be that the survival and growth of juvenile spiders is higher in areas with high chironomid abundances, leading to higher densities of adult spiders and higher predation rates on the terrestrial prey next spring.