High‐resolution food webs based on nitrogen isotopic composition of amino acids

Food webs are known to have myriad trophic links between resource and consumer species. While herbivores have well‐understood trophic tendencies, the difficulties associated with characterizing the trophic positions of higher‐order consumers have remained a major problem in food web ecology. To better understand trophic linkages in food webs, analysis of the stable nitrogen isotopic composition of amino acids has been introduced as a potential means of providing accurate trophic position estimates. In the present study, we employ this method to estimate the trophic positions of 200 free‐roaming organisms, representing 39 species in coastal marine (a stony shore) and 38 species in terrestrial (a fruit farm) environments. Based on the trophic positions from the isotopic composition of amino acids, we are able to resolve the trophic structure of these complex food webs. Our approach reveals a high degree of trophic omnivory (i.e., noninteger trophic positions) among carnivorous species such as marine fish and terrestrial hornets.This information not only clarifies the trophic tendencies of species within their respective communities, but also suggests that trophic omnivory may be common in these webs.     

Stable isotopic structure of aquatic ecosystems

Isotopic, biogeochemical and ecological structure can provide a new dimension for understanding material flows, and the simultaneous function and structure of an ecosystem. Distributions ofδ ¹³C andδ ¹⁵N for biogenic substances in the Nanakita river estuary involving Gamo lagoon in Japan were investigated to construct isotope biogeochemical and ecological structure for assessing fate and transfer of organic matter, and food web structure. The isotopic framework of the ecosystem was successfully described in aδ ¹⁵N–δ ¹³C map. In this estuary the variations of isotope ratios of biogenic substances were clearly explained by the mixing of land-derived organic matter, and marine-derived organic matter. A trophic-level effect of¹⁵N enrichment was clearly observed. Organisms were classified into three groups depending upon the contribution of land-derived organic matter in a food chain. Almost all biota except mollusca in the lagoon depend on organic matter of marine origin. The contributions of both land and marine organic matter were comparable for mollusca in the lagoon.     

Quantitative food web analysis supports the energy-limitation hypothesis in cave stream ecosystems

Energy limitation has long been the primary assumption underlying conceptual models of evolutionary and ecological processes in cave ecosystems. However, the prediction that cave communities are actually energy-limited in the sense that constituent populations are consuming all or most of their resource supply is untested. We assessed the energy-limitation hypothesis in three cave streams in northeastern Alabama (USA) by combining measurements of animal production, demand, and resource supplies (detritus, primarily decomposing wood particles). Comparisons of animal consumption and detritus supply rates in each cave showed that all, or nearly all, available detritus was required to support macroinvertebrate production. Furthermore, only a small amount of macroinvertebrate prey production remained to support other predatory taxa (i.e., cave fish and salamanders) after accounting for crayfish consumption. Placing the energy demands of a cave community within the context of resource supply rates provided quantitative support for the energy-limitation hypothesis, confirming the mechanism (limited energy surpluses) that likely influences the evolutionary processes and population dynamics that shape cave communities. Detritus-based surface ecosystems often have large detrital surpluses. Thus, cave ecosystems, which show minimal surpluses, occupy the extreme oligotrophic end of the spectrum of detritus-based food webs.     

Fish predation can induce mesohabitat-specific differences in food web structures in small stream ecosystems

Diverse benthic communities in streams include a wide variety of predators with different habitat preferences, e.g. for pools or riffles. We hypothesised that these preferences result in mesohabitat-specific predator community structures with quantitative differences concerning predation intensity by vertebrate and invertebrate predators, importance of intraguild predation, or top–down pressure. This hypothesis was evaluated for a small submontane stream by means of mesohabitat-specific quantification of prey consumption by two benthivorous fish species (Gobio gobio and Barbatula barbatula) and several invertebrate predators. The estimation was based on daily food rations and diet composition of predators and mesohabitat-specific predator biomass. We found clear differences between the two mesohabitat types. Predator food webs were less complex in pools than in riffles. Fish predation was more important than invertebrate predation in pools, and intraguild predation had a higher relative importance in these mesohabitats. These differences were probably caused by the mesohabitat use of G. gobio, the largest top predator, which preferred pools. Consequently, the predator food webs were more similar between the mesohabitats when fish were absent. Top–down pressure on primary consumers by all predators together was lowest in pools without fish, but the effect was not significant. Omnivory (including cannibalism) was intense, but its potentially destabilising effects were probably counterbalanced by mesohabitat connectivity. From the results of our experimental study, we conclude that even in small stream ecosystems, food web structures and predation pathways can differ between mesohabitats and that a mesohabitat-specific consideration will help to explain the variety of top–down effects on benthic communities.     

Diversity patterns and food web structure in a Mediterranean intermittent stream

In this work we present the results of a one‐year study on the macroinvertebrate community in an intermittent stream in southern Spain. We have studied the taxonomic composition, diversity and food web monthly in order to consider temporal variability in these parameters. More than 60 macroinvertebrate taxa have been recorded in the stream, but they do not cohabit at the same time. Many of them join the community at the beginning of the wet period. Afterwards, some new taxa incorporate but some others disappear. This leads to huge fluctuations in the diversity of the community and in the food web of different sampling dates. These variations are linked to environmental disturbances, mainly to flow peaks and minimums. From a relatively well structured, but simple, food web at the beginning of the wet period, it can be seen how several trophic levels disappear with time, and how several organisms change or extend their trophic function within it. We relate these variations to the intermediate disturbance hypothesis and discuss the resilience of the community of this intermittent stream (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

How habitat-modifying organisms structure the food web of two coastal ecosystems

The diversity and structure of ecosystems has been found to depend both on trophic interactions in food webs and on other species interactions such as habitat modification and mutualism that form non-trophic interaction networks. However, quantification of the dependencies between these two main interaction networks has remained elusive. In this study, we assessed how habitat-modifying organisms affect basic food web properties by conducting in-depth empirical investigations of two ecosystems: North American temperate fringing marshes and West African tropical seagrass meadows. Results reveal that habitat-modifying species, through non-trophic facilitation rather than their trophic role, enhance species richness across multiple trophic levels, increase the number of interactions per species (link density), but decrease the realized fraction of all possible links within the food web (connectance). Compared to the trophic role of the most highly connected species, we found this non-trophic effects to be more important for species richness and of more or similar importance for link density and connectance. Our findings demonstrate that food webs can be fundamentally shaped by interactions outside the trophic network, yet intrinsic to the species participating in it. Better integration of non-trophic interactions in food web analyses may therefore strongly contribute to their explanatory and predictive capacity.     

Biodiversity in model ecosystems, II: species assembly and food web structure

This is the second of two papers dedicated to the relationship between population models of competition and biodiversity. Here, we consider species assembly models where the population dynamics is kept far from fixed points through the continuous introduction of new species, and generalize to such models the coexistence condition derived for systems at the fixed point. The ecological overlap between species and shared preys, that we define here, provides a quantitative measure of the effective interspecies competition and of the trophic network topology. We obtain distributions of the overlap from simulations of a new model based both on immigration and speciation, and show that they are in good agreement with those measured for three large natural food webs. As discussed in the first paper, rapid environmental fluctuations, interacting with the condition for coexistence of competing species, limit the maximal biodiversity that a trophic level can host. This horizontal limitation to biodiversity is here combined with either dissipation of energy or growth of fluctuations, which in our model limit the length of food webs in the vertical direction. These ingredients yield an effective model of food webs that produce a biodiversity profile with a maximum at an intermediate trophic level, in agreement with field studies.     

Stable carbon and nitrogen isotopic composition of diet and hair of Gidra-speaking Papuans

The carbon and nitrogen stable isotopic composition of the scalp hair and diet of Gidra-speaking people in four villages in Papua New Guinea is presented. The isotopic composition of hair was measured, while that of the diet was estimated from food consumption survey data and the measured isotopic composition and protein and carbohydrate contents of food items. The average isotopic ratios of the hair samples and of the diet varied among the four study villages, which were selected because of their diverse ecological settings. Comparison was made between hair and calculated dietary isotopic compositions. Two of the four diet-hair enrichment values obtained for ¹³C (+1.8 and 2.2%‰) were similar to those previously reported (1.4–2.0%‰), but the other two values (3.7 and 4.8%) were greater than in earlier reports. ¹⁵N enrichment was systematically greater (by 1%‰) than reported values (∼4.3%‰) except for one village, where a much greater enrichment (6.9%‰) was found. The factors potentially relevant to these deviations are discussed. Possible errors in estimating the dietary isotopic composition and minor modifications of dietary habits revealed by food consumption surveys could explain most of the discrepancies. However, the great enrichment of ¹⁵N found in one of the villages remains unexplained. © 1996 Wiley-Liss, Inc.     

Stable isotopic composition of carbon and nitrogen and nitrogen content in vascular epiphytes along an altitudinal transect*

The foliar content of nitrogen and the relative abundances of ¹³C and ¹⁵N were analysed in vascular epiphytes collected from six sites along an altitudinal gradient from tropical dry forests to humid montane forests in eastern Mexico. The proportion of epiphyte species showing crassulacean acid metabolism (CAM) (atmospheric bromeliads, thick-leaved orchids, Cactaceae, and Crassulaceae) decreased with increasing elevation and precipitation from 58 to 6%. Atmospheric bromeliads, almost all of which had δ ¹³C values indicating CAM, were more depleted in ¹⁵N (x = ? 10·9‰ ± 2·11) than the C₃ bromeliads which form water-storing tanks ( ? 6·05‰ ± 2·26). As there was no difference in δ ¹⁵N values between C₃ and CAM orchids, the difference in bromeliads was not related to photosynthetic pathways but to different nitrogen sources. While epiphytes with strong ¹⁵N depletion appear to obtain their nitrogen mainly from direct atmospheric deposition, others have access to nitrogen in intercepted water and from organic matter decomposing on branches and in their phytotelmata. Bromeliads and succulent orchids had a lower foliar nitrogen content than thin-leaved orchids, ferns and Piperaceae. Ground-rooted hemi-epiphytes exhibited the highest nitrogen contents and δ ¹⁵N values.     

Climate change impacts on body size and food web structure on mountain ecosystems

The current distribution of climatic conditions will be rearranged on the globe. To survive, species will have to keep pace with climates as they move. Mountains are among the most affected regions owing to both climate and land-use change. Here, we explore the effects of climate change in the vertebrate food web of the Pyrenees. We investigate elevation range expansions between two time-periods illustrative of warming conditions, to assess: (i) the taxonomic composition of range expanders; (ii) changes in food web properties such as the distribution of links per species and community size-structure; and (iii) what are the specific traits of range expanders that set them apart from the other species in the community—in particular, body mass, diet generalism, vulnerability and trophic position within the food web. We found an upward expansion of species at all elevations, which was not even for all taxonomic groups and trophic positions. At low and intermediate elevations, predator : prey mass ratios were significantly reduced. Expanders were larger, had fewer predators and were, in general, more specialists. Our study shows that elevation range expansions as climate warms have important and predictable impacts on the structure and size distribution of food webs across space.     

Influence of decomposer food web structure and nitrogen availability on plant growth

We studied the sensitivity of soil microbial communities and ecosystem processes to variation in the vertical and horizontal structure of decomposer food web under nitrogen poor and N-enriched conditions. Microcosms with humus and litter layer of boreal forest floor, birch seedlings infected with mycorrhizal fungi, and decomposer food webs with differing trophic group and species composition of soil fauna were constructed. During the second growing period for the birch, we irrigated half of the microcosms with urea solution, and the other half with de-ionised water to create two levels of N concentration in the substrate. During the experiment night time respirations of the microcosms were measured, and the water leached through the microcosms was analysed for concentration of mineral N, and nematode numbers. The microcosms were destructively sampled after 37 weeks for plant biomass and N uptake, structure of soil animal and microbial community (indicated by PLFA profiles), and physical and chemical properties of the humus and litter materials. Predatory mites and nematodes had a negative influence on the biomass of their microbivorous and microbi-detritivorous prey, and microbi-detritivores affected the biomass and community structure of microbes (indicated by PLFA-analysis). Moreover, predatory mites and nematodes increased microbial biomass and changed the microbial community structure. The decomposer food web structure affected also N uptake and growth of plants. Microbi-detritivorous fauna had a positive effect, whereas predators of microbial and detritus feeding fauna exerted a negative influence on plant N uptake and biomass production. The impact of a trophic group on the microbes and plant was also strongly dependent on species composition within the group. Nitrogen addition magnified the influence of food web structure on microbial biomass and plant N uptake. We suggest that addition of urea-N to the soil modified the animal-microbe interaction by increasing microbial growth and altering community structure of microbes. The presence of microbi-detritivores and predators reduced loss of carbon from the microcosms, and the food web structure influenced also water holding capacity of the materials. The changes in plant growth, nutrient cycling, size of N and C pools, and in the physical properties of the soil emphasize the importance and diversity of indirect consequences of decomposer food web structure. This revised version was published online in June 2006 with corrections to the Cover Date.     

The predatory Chironomidae of an iron-rich stream: feeding ecology and food web structure

Abstract. 1. Three species of Tanypodinae (Chironomidae) were found in an acid and iron-rich stream in southern England. Maximum abundance was achieved in summer and they were sparse at other times. Individuals were aggregated on the stream bed and were overrepresented in accumulations of leaf litter.
2. The diets of all three species consisted of a mixture of prey (prominently detritivorous chironomid larvae) and detritus. More detritus and fewer prey were taken in winter than in summer.
3. When comparing large tanypod species with small and, intraspecifically, late instars with early, the proportion of guts containing prey increased with increasing body size.
4. Stonefly larvae were more prominent in the diet of Zavrelimyia barbatipes (Kieffer) in summer than in winter but for the other two species the reverse was true. A bigger proportion of Trissopelopia longimana (Staeger) guts contained prey in early summer than in August whereas more Macropelopia goetghebueri (Kieffer) guts contained prey in August. This was apparently a consequence of seasonal differences in the distribution of body size among the populations of these two species.
5. The stream contains two further common predators, Plectrocnemia conspersa (Curtis) and Sialis fuliginosa Pict. These are important predators of tanypod larvae but might also compete with them since they severely deplete populations of prey taken in common.
6. Analysis of the food-web in Broadstone Stream reveals remarkably high values of connectance (C and C_max) and of species richness times connectance (SC_max). Such characteristics are theoretically associated with fragile and dynamically unstable food webs, and may be found in 'constant' environments. There is also an apparently unusual prevalence of omnivory in the community.     

Food web structure in a tropical stream ecosystem

Abstract This study investigated the structure and properties of a tropical stream food web in a small spatial scale, characterizing its planktonic, epiphytic and benthic compartments. The study was carried out in the Potreirinho Creek, a second‐order stream located in the south‐east of Brazil. Some attributes of the three subwebs and of the conglomerate food web, composed by the trophic links of the three compartments plus the fish species, were determined. Among compartments, the food webs showed considerable variation in structure. The epiphytic food web was consistently more complex than the planktonic and benthic webs. The values of number of species, number of links and maximum food chain length were significantly higher in the epiphytic compartment than in the other two. Otherwise, the connectance was significantly lower in epiphyton. The significant differences of most food web parameters were determined by the increase in the number of trophic species, represented mainly by basal and intermediate species. High species richness, detritus‐based system and high degree of omnivory characterized the stream food web studied. The aquatic macrophytes probably provide a substratum more stable and structurally complex than the sediment. We suggest that the greater species richness and trophic complexity in the epiphytic subweb might be due to the higher degree of habitat complexity supported by macrophyte substrate. Despite differences observed in the structure of the three subwebs, they are highly connected by trophic interactions, mainly by fishes. The high degree of fish omnivory associated with their movements at different spatial scales suggests that these animals have a significant role in the food web dynamic of Potreirinho Creek. This interface between macrophytes and the interconnections resultant from fish foraging, diluted the compartmentalization of the Potreirinho food web.     

Stable isotopes in physiological ecology and food web research

Mention the word 'isotopes' and most people will think of short-lived radioactive isotopes. Yet far more abundant than the radioactive forms and perhaps much more useful for ecological studies are the stable isotopes. Early interest in stable isotope analyses developed in the geological sciences, and their application in environmental biology has developed slowly until recently. Over the past few years, innovative applications of stable isotope analyses to studies of biological processes have been expanding rapidly, and there is every indication that stable isotope approaches will lead to major advances over the next decade in our understanding of physiological processes and fluxes through ecological systems. Studies of plant and animal physiological ecology and food webs will benefit greatly.     

Relating planktonic microbial food web structure in lentic freshwater ecosystems to water quality and land use

Relationships among picoplankton, protozoa, phytoplankton, plantnutrients, lake type, drainage basin morphology and land coverwere studied in 45 water bodies in South Island, New Zealandthat ranged from large, deep, ultra-oligotrophic lakes to shallow,macrophyte-dominated ponds and swamps. The biomasses of mostheterotrophic components of the pelagic microbial food webswere positively related to phytoplankton and features of thedrainage basin that enhanced nutrient input, and imply strongresource-driven structuring of pelagic microbial food webs.Prokaryotic picophytoplankton biomass was negatively relatedto indices of eutrophication, and the picoautotroph contributionto total microbial food web biomass declined with increasingtotal phosphorus concentration from 16.5% in deep lakes to <0.02%in swamps and ponds. Biomass ratios of (picoplankton plus protozoa):phytoplanktonranged from 40:60 in swamps and ponds to >70:30 in deep lakes,and indicate the potential importance of microbial food websin carbon transfer to higher trophic levels in deep, less productivelakes. Strong relationships exist between land use in the catchmentand pelagic microbial food web structure and biomass acrossa wide range in size and trophic state of water bodies in heterogeneouslandscapes.     

Microcrustacean prey and macroinvertebrate predators in a stream food web

1. The consumption of microcrustacea by two polyphagous predators, larvae of the caddisfly Plectrocnemia conspersa (Curtis) and the alderfly Sialis fuliginosa Pictet, was investigated in an English stream with a well-known macro- and microinvertebrate fauna. Benthic samples were collected in August, November, December and April, and the gut contents of all individuals of both predators were examined. 2. All the microcrustacean groups (Cyclopoida, Harpacticoida, Chydoridae and Ostracoda) were identified in gut contents. Of the ten taxa present in the benthos, all occurred in the diet of P. conspersa; nine were found in S. fuliginosa. 3. Ontogenetic shifts in the diets of both predators were found, and microcrustacea were consumed more frequently by small than large instars. 4. There was little evidence of selective feeding by P. conspersa, whereas ostracods were over-represented in the diet of S. fuliginosa, compared with benthic relative densities. The Chydoridae were under-represented in the diet of both predators. 5. The food web of Broadstone Stream is perhaps the most detailed web available for any running water habitat. Increased taxonomic resolution produced marked changes in values of connectance and predator-prey ratios. Linkage density remained fairly constant at different levels of resolution and were high, indicative of a web of generalist species. Omnivory was pronounced and may be characteristic of donor-controlled systems where organic detritus is the primary energy base.     

Brown trout and food web interactions in a Minnesota stream

1. We examined indirect, community‐level interactions in a stream that contained non‐native brown trout (Salmo trutta Linnaeus), native brook trout (Salvelinus fontinalis Mitchill) and native slimy sculpin (Cottus cognatus Richardson). Our objectives were to examine benthic invertebrate composition and prey selection of fishes (measured by total invertebrate dry mass, dry mass of individual invertebrate taxa and relative proportion of invertebrate taxa in the benthos and diet) among treatments (no fish, juvenile brook trout alone, juvenile brown trout alone, sculpin with brook trout and sculpin with brown trout). 2. We assigned treatments to 1 m² enclosures/exclosures placed in riffles in Valley Creek, Minnesota, and conducted six experimental trials. We used three designs of fish densities (addition of trout to a constant number of sculpin with unequal numbers of trout and sculpin; addition of trout to a constant number of sculpin with equal numbers of trout and sculpin; and replacement of half the sculpin with an equal number of trout) to investigate the relative strength of interspecific versus intraspecific interactions. 3. Presence of fish (all three species, alone or in combined‐species treatments) was not associated with changes in total dry mass of benthic invertebrates or shifts in relative abundance of benthic invertebrate taxa, regardless of fish density design. 4. Brook trout and sculpin diets did not change when each species was alone compared with treatments of both species together. Likewise, we did not find evidence for shifts in brown trout or sculpin diets when each species was alone or together. 5. We suggest that native brook trout and non‐native brown trout fill similar niches in Valley Creek. We did not find evidence that either species had an effect on stream communities, potentially due to high invertebrate productivity in Valley Creek.     

Incorporation of mineral nitrogen into the soil food web as affected by plant community composition

Although nitrogen (N) deposition is increasing globally, N availability still limits many organisms, such as microorganisms and mesofauna. However, little is known to which extent soil organisms rely on mineral‐derived N and whether plant community composition modifies its incorporation into soil food webs. More diverse plant communities more effectively compete with microorganisms for mineral N likely reducing the incorporation of mineral‐derived N into soil food webs. We set up a field experiment in experimental grasslands with different levels of plant species and functional group richness. We labeled soil with ¹⁵NH₄ ¹⁵NO₃ and analyzed the incorporation of mineral‐derived ¹⁵N into soil microorganisms and mesofauna over 3 months. Mineral‐derived N incorporation decreased over time in all investigated organisms. Plant species richness and presence of legumes reduced the uptake of mineral‐derived N into microorganisms. In parallel, the incorporation of mineral‐derived ¹⁵N into mesofauna species declined with time and decreased with increasing plant species richness in the secondary decomposer springtail Ceratophysella sp. Effects of both plant species richness and functional group richness on other mesofauna species varied with time. The presence of grasses increased the ¹⁵N incorporation into Ceratophysella sp., but decreased it in the primary decomposer oribatid mite Tectocepheus velatus sarekensis. The results highlight that mineral N is quickly channeled into soil animal food webs via microorganisms irrespective of plant diversity. The amount of mineral‐derived N incorporated into soil animals, and the plant community properties affecting this incorporation, differed markedly between soil animal taxa, reflecting species‐specific use of food resources. Our results highlight that plant diversity and community composition alter the competition for N in soil and change the transfer of N across trophic levels in soil food webs, potentially leading to changes in soil animal population dynamics and community composition. Sustaining high plant diversity may buffer detrimental effects of elevated N deposition on soil biota.     

Urbanisation alters fatty acids in stream food webs

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