Local‐ to continental‐scale variation in the richness and composition of an aquatic food web

Aim We investigated patterns of species richness and composition of the aquatic food web found in the liquid‐filled leaves of the North American purple pitcher plant, Sarracenia purpurea (Sarraceniaceae), from local to continental scales. Location We sampled 20 pitcher‐plant communities at each of 39 sites spanning the geographic range of S. purpurea– from northern Florida to Newfoundland and westward to eastern British Columbia. Methods Environmental predictors of variation in species composition and species richness were measured at two different spatial scales: among pitchers within sites and among sites. Hierarchical Bayesian models were used to examine correlates and similarities of species richness and abundance within and among sites. Results Ninety‐two taxa of arthropods, protozoa and bacteria were identified in the 780 pitcher samples. The variation in the species composition of this multi‐trophic level community across the broad geographic range of the host plant was lower than the variation among pitchers within host‐plant populations. Variation among food webs in richness and composition was related to climate, pore‐water chemistry, pitcher‐plant morphology and leaf age. Variation in the abundance of the five most common invertebrates was also strongly related to pitcher morphology and site‐specific climatic and other environmental variables. Main conclusions The surprising result that these communities are more variable within their host‐plant populations than across North America suggests that the food web in S. purpurea leaves consists of two groups of species: (1) a core group of mostly obligate pitcher‐plant residents that have evolved strong requirements for the host plant and that co‐occur consistently across North America, and (2) a larger set of relatively uncommon, generalist taxa that co‐occur patchily.     

Geographic variation in network structure of a nearctic aquatic food web

Aim The network structure of food webs plays an important role in the maintenance of diversity and ecosystem functioning in ecological communities. Previous research has found that ecosystem size, resource availability, assembly history and biotic interactions can potentially drive food web structure. However, the relative influence of climatic variables that drive broad‐scale biogeographic patterns of species richness and composition has not been explored for food web structure. In this study, we assess the influence of broad‐scale climatic variables in addition to known drivers of food web structure on replicate observations of a single aquatic food web, sampled from the leaves of the pitcher plant (Sarracenia purpurea), at different geographic sites across a broad latitudinal and climatic range. Location Using standardized sampling methods, we conducted an extensive ‘snapshot’ survey of 780 replicated aquatic food webs collected from the leaves of the pitcher plant S. purpurea at 39 sites from northern Florida to Newfoundland and westward to eastern British Columbia. Methods We examined correlations of 15 measures of food web structure at the pitcher and site scales with geographic variation in temperature and precipitation, concentrations of nutrients from atmospheric nitrogen deposition, resource availability, ecosystem size and the abundance of the pitcher plant mosquito (Wyeomyia smithii), a potential keystone species. Results At the scale of a single pitcher plant leaf, linkage density, species richness, measures of chain length and the proportion of omnivores in a web all increased with pitcher volume. Linkage density and species richness were greater at high‐latitude sites, which experience low mean temperatures and precipitation and high annual variation in both of these variables. At the site scale, variation in 8 of the 15 food web metrics decreased at higher latitudes, and variation in measures of chain length increased with the abundance of mosquitoes. Main conclusions Ecosystem size and climatic variables related to latitude were most strongly correlated with network structure of the Sarracenia food web. However, in spite of large sample sizes, thorough standardized sampling and the large geographic extent of the survey, even the best‐fitting models explained less than 40% of the variation in food web structure. In contrast to biogeographic patterns of species richness, food web structure was largely independent of broad‐scale climatic variables. The large proportion of unexplained variance in our analyses suggests that stochastic assembly may be an important determinant of local food web structure.     

Predicting food‐web structure with metacommunity models

Synthesis Metacommunity theory aims to elucidate the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity research has focused largely on assemblages of competing organisms within a single trophic level. Here, we test the ability of metacommunity models to predict the network structure of the aquatic food web found in the leaves of the northern pitcher plant Sarracenia purpurea. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions play an important role in structuring Sarracenia food webs. Our approach can be applied to any well‐resolved food web for which data are available from multiple locations. The metacommunity framework explores the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity models and empirical studies have focused mostly on assemblages of competing organisms within a single trophic level. Studies of multi‐trophic metacommunities are predominantly restricted to simplified trophic motifs and rarely consider entire food webs. We tested the ability of the patch‐dynamics, species‐sorting, mass‐effects, and neutral metacommunity models, as well as three hybrid models, to reproduce empirical patterns of food web structure and composition in the complex aquatic food web found in the northern pitcher plant Sarracenia purpurea. We used empirical data to determine regional species pools and estimate dispersal probabilities, simulated local food‐web dynamics, dispersed species from regional pools into local food webs at rates based on the assumptions of each metacommunity model, and tested their relative fits to empirical data on food‐web structure. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions were important in structuring Sarracenia food webs. However, differences in dispersal abilities were also important in models that accurately reproduced empirical food web properties. Although the models were tested using pitcher‐plant food webs, the approach we have developed can be applied to any well‐resolved food web for which data are available from multiple locations.     

Relative importance of top-down and bottom-up forces in food webs of Sarracenia pitcher communities at a northern and a southern site

The relative importance of resources (bottom-up forces) and natural enemies (top-down forces) for regulating food web dynamics has been debated, and both forces have been found to be critical for determining food web structure. How the relative importance of top-down and bottom-up forces varies between sites with different abiotic conditions is not well understood. Using the pitcher plant inquiline community as a model system, I examine how the relative importance of top-down and bottom-up effects differs between two disparate sites. Resources (ant carcasses) and top predators (mosquito larvae) were manipulated in two identical 4 × 4 factorial press experiments, conducted at two geographically distant sites (Michigan and Florida) within the range of the purple pitcher plant, Sarracenia purpurea, and the aquatic community that resides in its leaves. Overall, top predators reduced the density of prey populations while additional resources bolstered them, and the relative importance of top-down and bottom-up forces varied between sites and for different trophic levels. Specifically, top-down effects on protozoa were stronger in Florida than in Michigan, while the opposite pattern was found for rotifers. These findings experimentally demonstrate that the strength of predator–prey interactions, even those involving the same species, vary across space. While only two sites are compared in this study, I hypothesize that site differences in temperature, which influences metabolic rate, may be responsible for variation in consumer–resource interactions. These findings warrant further investigation into the specific factors that modify the relative importance of top-down and bottom-up effects.     

Reverse latitudinal trends in species richness of pitcher-plant food webs

Latitudinal patterns in species richness have been well documented for guilds and individual trophic groups, but comparable patterns for entire, multitrophic communities have not been described. We studied the entire food web that inhabits the water‐filled leaves of the pitcher plant Sarracenia purpurea across North America at two spatial scales: among sites and among leaves within sites. Contrary to the expectation, total species richness at both scales increased with latitude, because of increasing species richness at the lower trophic levels. This latitudinal pattern may be driven by a top‐down effect. The abundance of the mosquito Wyeomyia smithii, a ubiquitous top predator in this system, decreases from south to north and may permit greater species richness of prey trophic levels at higher latitudes.     

Food-web models predict species abundances in response to habitat change

Plant and animal population sizes inevitably change following habitat loss, but the mechanisms underlying these changes are poorly understood. We experimentally altered habitat volume and eliminated top trophic levels of the food web of invertebrates that inhabit rain-filled leaves of the carnivorous pitcher plant Sarracenia purpurea. Path models that incorporated food-web structure better predicted population sizes of food-web constituents than did simple keystone species models, models that included only autecological responses to habitat volume, or models including both food-web structure and habitat volume. These results provide the first experimental confirmation that trophic structure can determine species abundances in the face of habitat loss.     

Effects of temperature variability on community structure in a natural microbial food web

Climate change research has demonstrated that changing temperatures will have an effect on community‐level dynamics by altering species survival rates, shifting species distributions, and ultimately, creating mismatches in community interactions. However, most of this work has focused on increasing temperature, and still little is known about how the variation in temperature extremes will affect community dynamics. We used the model aquatic community held within the leaves of the carnivorous plant, Sarracenia purpurea, to test how food web dynamics will be affected by high temperature variation. We tested the community response of the first (bacterial density), second (protist diversity and composition), and third trophic level (predator mortality), and measured community respiration. We collected early and late successional stage inquiline communities from S. purpurea from two North American and two European sites with similar average July temperature. We then created a common garden experiment in which replicates of these communities underwent either high or normal daily temperature variation, with the average temperature equal among treatments. We found an impact of temperature variation on the first two, but not on the third trophic level. For bacteria in the high‐variation treatment, density experienced an initial boost in growth but then decreased quickly through time. For protists in the high‐variation treatment, alpha‐diversity decreased faster than in the normal‐variation treatment, beta‐diversity increased only in the European sites, and protist community composition tended to diverge more in the late successional stage. The mortality of the predatory mosquito larvae was unaffected by temperature variation. Community respiration was lower in the high‐variation treatment, indicating a lower ecosystem functioning. Our results highlight clear impacts of temperature variation. A more mechanistic understanding of the effects that temperature, and especially temperature variation, will have on community dynamics is still greatly needed.     

A keystone predator controls bacterial diversity in the pitcher-plant (Sarracenia purpurea) microecosystem

The community of organisms inhabiting the water-filled leaves of the carnivorous pitcher-plant Sarracenia purpurea includes arthropods, protozoa and bacteria, and serves as a model system for studies of food web dynamics. Despite the wealth of data collected by ecologists and zoologists on this food web, very little is known about the bacterial assemblage in this microecosystem. We used terminal restriction fragment length polymorphism (T-RFLP) analysis to quantify bacterial diversity within the pitchers as a function of pitcher size, pH of the pitcher fluid and the presence of the keystone predator in this food web, larvae of the pitcher-plant mosquito Wyeomyia smithii. Results were analysed at two spatial scales: within a single bog and across three isolated bogs. Pitchers were sterile before they opened and composition of the bacterial assemblage was more variable between different bogs than within bogs. Measures of bacterial richness and diversity were greater in the presence of W. smithii and increased with increasing pitcher size. Our results suggest that fundamental ecological concepts derived from macroscopic food webs can also be used to predict the bacterial assemblages in pitcher plants.     

Proteomic characterization of the major arthropod associates of the carnivorous pitcher plant Sarracenia purpurea

The array of biomolecules generated by a functioning ecosystem represents both a potential resource for sustainable harvest and a potential indicator of ecosystem health and function. The cupped leaves of the carnivorous pitcher plant, Sarracenia purpurea, harbor a dynamic food web of aquatic invertebrates in a fully functional miniature ecosystem. The energetic base of this food web consists of insect prey, which is shredded by aquatic invertebrates and decomposed by microbes. Biomolecules and metabolites produced by this food web are actively exchanged with the photosynthesizing plant. In this report, we provide the first proteomic characterization of the sacrophagid fly (Fletcherimyia fletcheri), the pitcher plant mosquito (Wyeomyia smithii), and the pitcher-plant midge (Metriocnemus knabi). These three arthropods act as predators, filter feeders, and shredders at distinct trophic levels within the S. purpurea food web. More than 50 proteins from each species were identified, ten of which were predominantly or uniquely found in one species. Furthermore, 19 peptides unique to one of the three species were identified using an assembled database of 100 metazoan myosin heavy chain orthologs. These molecular signatures may be useful in species monitoring within heterogeneous ecosystem biomass and may also serve as indicators of ecosystem state.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

The Pitcher Plant Sarracenia purpurea Can Directly Acquire Organic Nitrogen and Short-Circuit the Inorganic Nitrogen Cycle

Background

Despite the large stocks of organic nitrogen in soil, nitrogen availability limits plant growth in many terrestrial ecosystems because most plants take up only inorganic nitrogen, not organic nitrogen. Although some vascular plants can assimilate organic nitrogen directly, only recently has organic nitrogen been found to contribute significantly to the nutrient budget of any plant. Carnivorous plants grow in extremely nutrient-poor environments and carnivory has evolved in these plants as an alternative pathway for obtaining nutrients. We tested if the carnivorous pitcher plant Sarracenia purpurea could directly take up intact amino acids in the field and compared uptake of organic and inorganic forms of nitrogen across a gradient of nitrogen deposition. We hypothesized that the contribution of organic nitrogen to the nitrogen budget of the pitcher plant would decline with increasing nitrogen deposition.

Methodology and Principal Findings

At sites in Canada (low nitrogen deposition) and the United States (high nitrogen deposition), individual pitchers were fed two amino acids, glycine and phenylalanine, and inorganic nitrogen (as ammonium nitrate), individually and in mixture. Plants took up intact amino acids. Acquisition of each form of nitrogen provided in isolation exceeded uptake of the same form in mixture. At the high deposition site, uptake of organic nitrogen was higher than uptake of inorganic nitrogen. At the low deposition site, uptake of all three forms of nitrogen was similar. Completeness of the associated detritus-based food web that inhabits pitcher-plant leaves and breaks down captured prey had no effect on nitrogen uptake.

Conclusions and Significance

By taking up intact amino acids, Sarracenia purpurea can short-circuit the inorganic nitrogen cycle, thus minimizing potential bottlenecks in nitrogen availability that result from the plant''s reliance for nitrogen mineralization on a seasonally reconstructed food web operating on infrequent and irregular prey capture.     

Motifs in the assembly of food web networks

The assembly of local communities from regional pools is a multifaceted process that involves the confluence of interactions and environmental conditions at the local scale and biogeographic and evolutionary history at the regional scale. Understanding the relative influence of these factors on community structure has remained a challenge and mechanisms driving community assembly are often inferred from patterns of taxonomic, functional, and phylogenetic diversity. Moreover, community assembly is often viewed through the lens of competition and rarely includes trophic interactions or entire food webs. Here, we use motifs – subgraphs of nodes (e.g. species) and links (e.g. predation) whose abundance within a network deviates significantly as compared to a random network topology – to explore the assembly of food web networks found in the leaves of the northern pitcher plant Sarracenia purpurea. We compared counts of three‐node motifs across a hierarchy of scales to a suite of null models to determine if motifs are over‐, under‐, or randomly represented. We then assessed if the pattern of representation of a motif in a given network matched that of the network it was assembled from. We found that motif representation in over 70% of site networks matched the continental network they were assembled from and over 75% of local networks matched the site networks they were assembled from for the majority of null models. This suggests that the same processes are shaping networks across scales. To generalize our results and effectively use a motif perspective to study community assembly, a theoretical framework detailing potential mechanisms for all possible combinations of motif representation is necessary.     

Nutrient limitation in detritus-based microcosms in Sarracenia purpurea

Most prior work on the role of top-down and bottom-up effects in aquatic communities has ignored the significant detrital component that occurs in natural systems. We investigated the effects of specific nutrients (carbon, phosphorus, and nitrogen), as well as a top predator (the mosquito Wyeomyia smithii), on the structure of the detritivore community found in the water-filled leaves of the pitcher plant Sarracenia purpurea. The concentrations of three nutrients and the presence of the predators were manipulated in a factorial design, while the response of the remaining community was quantified. Bacterial growth was found to be strongly carbon-limited and somewhat less limited by phosphorus and there was an interaction between the effects of the two nutrients. Neither carbon or phosphorus addition affected protozoan or rotifer abundance, and nitrogen had only a minor effect. The presence of the predator, however, significantly reduced the abundance of the four numerically dominant bacteriovores. There were no interactions between top-down and bottom-up effects; the strong direct reciprocal effects between adjacent trophic levels seem to be greatly attenuated as they are propagated farther up or down the food chain.     

Intermediate-consumer identity and resources alter a food web with omnivory

1. Omnivory is an important interaction that has been the centre of numerous theoretical and empirical studies in recent years. Most of these studies examine the conditions necessary for coexistence between an omnivore and an intermediate consumer. Trait variation in ecological interactions (competition and predator tolerance) among intermediate consumers has not been considered in previous empirical studies despite the evidence that variation in species-specific traits can have important community-level effects. 2. I conducted a multifactorial microcosm experiment using species from the Sarracenia purpurea phytotelmata community, organisms that inhabit the water collected within its modified leaves. The basal trophic level consisted of bacterial decomposers, the second trophic level (intermediate consumers) consisted of protozoa and rotifers, and the third trophic level (omnivore) were larvae of the pitcher plant mosquito Wyeomyia smithii. Trophic level number (1, 2 and 3), resources (low and high), omnivore density (low and high) and intermediate consumer (monoculture of five protozoa and rotifers) identity were manipulated. Abundance of the basal trophic level, intermediate consumers, and growth of the omnivore were measured, as well as time to extinction (intermediate consumers) and time to pupation (mosquito larvae). 3. The presence of different intermediate consumers affected both bacteria abundance and omnivore growth. At high resource levels, Poteriochromonas, Colpidium and Habrotrocha rosa reduced bacteria densities greater than omnivore reduction of bacteria. Mosquito larvae did not pupate at low resource levels except when Poteriochromonas and Colopoda were present as intermediate consumers. Communities with H. rosa were the only ones consistent with the prediction that omnivores should exclude intermediate consumers at high resources. 4. These results had mixed support for predictions from omnivory food web theory. Intermediate consumers responded and affected this community differently under different community structures and resource levels. Consequently, variation in species-specific traits can have important population- and community-level effects and needs to be considered in food webs with omnivory.     

Local and Regional-Scale Food Web Structure in Nepenthes alata Pitchers1

Tropical Nepenthes pitcher plants provide small, isolated aquatic habitats. We examined inter-pitcher variation in the community structure of the inhabitants of Nepenthes alata Blanco in West Sumatra, focusing on the conditions of the pitchers, bacterial density in the pitcher fluid, density and biomass of metazoan inhabitants, and the frequencies of interspecific encounters. Older pitchers contained more insect carcasses. The bacterial density increased with the age of the pitchers, but decreased in withered pitchers that contained finely decomposed detritus. In live pitchers, the bacterial density, the density, mass and species richness of metazoa, and the number of trophic levels per pitcher were positively correlated with detrital mass, which was correlated with volume of pitcher fluid. The metazoan fauna from N. alata consisted of 4 predators and 12 saprophages, among the richest known for Nepenthes species. However, each individual pitcher harbored a limited numbers of species, owing to (1) the low incidence of many species, and (2) the aggregated distribution and different temporal colonization pattern of major species. Six dipteran taxa (one predator and five saprophages) accounted for the bulk of metazoan inhabitant biomass. Of 48 combinations of predator-prey encountered, only four occurred frequently (in > 30% of pitchers), which included two predators and three saprophages. Thus, individual pitchers harbored relatively simple communities despite the regional species richness, and only limited kinds of predator-prey encounters seemed to occur frequently in the regional food web. The local-scale properties of the subdivided communities presented here provide the basic information for understanding the maintenance of regional species richness and food web complexity.     

Variance in composition of inquiline communities in leaves of Sarracenia purpurea L. on multiple spatial scales

A survey of the abundances of species that inhabit the water-bearing leaves of the pitcher plant Sarracenia purpurea was conducted at several different spatial scales in northern Florida. Individual leaves are hosts to communities of inquiline species, including mosquitoes, midges, mites, copepods, cladocerans, and a diverse bacterial assemblage. Inquiline communities were quantified from four pitchers per plant, three plants per subpopulation, two subpopulations per population, and three populations. Species varied in abundance at different spatial scales. Variation in the abundances of mosquitoes and copepods was not significantly associated with any spatial scale. Midges varied in abundance at the level of populations; one population contained significantly more midges than the other two. Cladocerans varied at the level of the subpopulation, whereas mites varied at the level of the individual plants. Bacterial communities were described by means of Biolog plates, which quantify the types of carbon media used by the bacteria in each pitcher. Bacterial communities were found to vary significantly in composition among individual plants but not among populations or subpopulations. These results suggest that independent factors determining the abundances of individual species are important in determining community patterns in pitcher-plant inquilines.     

Soil Food Web Changes during Spontaneous Succession at Post Mining Sites: A Possible Ecosystem Engineering Effect on Food Web Organization?

Parameters characterizing the structure of the decomposer food web, biomass of the soil microflora (bacteria and fungi) and soil micro-, meso- and macrofauna were studied at 14 non-reclaimed 1– 41-year-old post-mining sites near the town of Sokolov (Czech Republic). These observations on the decomposer food webs were compared with knowledge of vegetation and soil microstructure development from previous studies. The amount of carbon entering the food web increased with succession age in a similar way as the total amount of C in food web biomass and the number of functional groups in the food web. Connectance did not show any significant changes with succession age, however. In early stages of the succession, the bacterial channel dominated the food web. Later on, in shrub-dominated stands, the fungal channel took over. Even later, in the forest stage, the bacterial channel prevailed again. The best predictor of fungal bacterial ratio is thickness of fermentation layer. We argue that these changes correspond with changes in topsoil microstructure driven by a combination of plant organic matter input and engineering effects of earthworms. In early stages, soil is alkaline, and a discontinuous litter layer on the soil surface promotes bacterial biomass growth, so the bacterial food web channel can dominate. Litter accumulation on the soil surface supports the development of the fungal channel. In older stages, earthworms arrive, mix litter into the mineral soil and form an organo-mineral topsoil, which is beneficial for bacteria and enhances the bacterial food web channel.     

Effects of native and exotic range‐expanding plant species on taxonomic and functional composition of nematodes in the soil food web

Due to climate warming, many plant species shift ranges towards higher latitudes. Plants can disperse faster than most soil biota, however, little is known about how range‐expanding plants in the new range will establish interactions with the resident soil food web. In this paper we examine how the soil nematode community from the new range responds to range‐expanding plant species compared to related natives. We focused on nematodes, because they are important components in various trophic levels of the soil food web, some feeding on plant roots, others on microbes or on invertebrates. We expected that range expanding plant species have fewer root‐feeding nematodes, as predicted by enemy release hypothesis. We therefore expected that range expanders affect the taxonomic and functional composition of the nematode community, but that these effects would diminish with increasing trophic position of nematodes in the soil food web. We exposed six range expanders (including three intercontinental exotics) and nine related native plant species to soil from the invaded range and show that range expanders on average had fewer root‐feeding nematodes per unit root biomass than related natives. The range expanders showed resistance against rather than tolerance for root‐feeding nematodes from the new range. On the other hand, the overall taxonomic and functional nematode community composition was influenced by plant species rather than by plant origin. The plant identity effects declined with trophic position of nematodes in the soil food web, as plant feeders were influenced more than other feeding guilds. We conclude that range‐expanding plant species can have fewer root‐feeding nematodes per unit root biomass than related natives, but that the taxonomic and functional nematode community composition is determined more by plant identity than by plant origin. Plant species identity effects decreased with trophic position of nematodes in the soil food web.     

Resolving the predator first paradox: Arthropod predator food webs in pioneer sites of glacier forelands

Primary succession on bare ground surrounded by intact ecosystems is, during its first stages, characterized by predator‐dominated arthropod communities. However, little is known on what prey sustains these predators at the start of succession and which factors drive the structure of these food webs. As prey availability can be extremely patchy and episodic in pioneer stages, trophic networks might be highly variable. Moreover, the importance of allochthonous versus autochthonous food sources for these pioneer predators is mostly unknown. To answer these questions, the gut content of 1,832 arthropod predators, including four species of carabid beetles, two lycosid and several linyphiid spider species caught in early and late pioneer stages of three glacier forelands, was screened molecularly to track intraguild and extraguild trophic interactions among all major prey groups occurring in these systems. Two‐thirds of the 2,310 identified food detections were collembolans and intraguild prey, while one‐third were allochthonous flying insects. Predator identity and not successional stage or valley had by far the strongest impact on the trophic interaction patterns. Still, the variability of prey spectra increased significantly from early to late pioneer stage, as did the niche width of the predators. As such the structure of pioneer arthropod food webs in recently deglaciated Alpine habitats seems to be driven foremost by predator identity while site and early successional effects contribute to a lesser extent to food web variability. Our findings also suggest that in these pioneer sites, predatory arthropods depend less on allochthonous aeolian prey but are mainly sustained by prey of local production.     

A stable isotope study of linkages between stream and terrestrial food webs through spider predation

SUMMARY 1. Transfer of carbon from freshwater to terrestrial ecosystems can occur through predation on adult aquatic insects, but the significance of this trophic pathway to the energetics of riparian communities is poorly understood. We used stable isotopes of carbon and nitrogen to explore linkages between aquatic insect production and the nutrition of web‐building and free‐living spiders alongside two streams in the North Island of New Zealand. 2. δ¹³C values for riparian tree leaves (means for each site = ?32.2 and ?30.3‰) were distinct from those of lichens collected from stream channel rocks and instream algae, both of which were similar (?23.4 to ?22.4‰). δ¹⁵N values for leaves were similar at both sites (?3.4 and ?2.7‰), but algae were considerably more depleted in δ¹⁵N atonesite suggesting significant differences in instream nitrogen sources between the twostreams. 3. Isotope values for potential aquatic prey of spiders indicated that aquatic algal production was their primary carbon source at both sites. Terrestrial invertebrates collected and assumed to be potential prey reflected a range of carbon sources and represented several trophic levels. 4. At one site, δ¹³C values indicated a primarily algae‐aquatic insect pathway of carbon transfer to both web‐building and free‐living spider guilds. The other site appeared to have a primarily terrestrial carbon pathway for the free‐living spider guild, and a mixed aquatic‐terrestrial pathway for the web‐building guild. 5. Overall, web‐building spiders were estimated to obtain around 61% of their body carbon from aquatic production compared with 55% for free‐living spiders. Our findings suggest that consumption of prey derived from aquatic sources can provide significant nutrition for spiders living along some stream channels. This pathway may represent an important feedback mechanism contributing to the energetics of riparian communities at sites where aquatic insect production is high.