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.     

Cascading trophic effects in pampean shallow lakes: results of a mesocosm experiment using two coexisting fish species with different feeding strategies

Planktivorous fish, both visual predators and filter feeders, enhance eutrophication processes in lakes. In pampean shallow lakes several planktivorous species may coexist but often two species dominate: silverside (Odontesthes bonariensis), a visual planktivorous fish when young adult, and sabalito (Cyphocharax voga), an omnivorous filter feeder. To assess the relative strength of the cascading trophic effects of the two species, a mesocosm experiment was conducted using different proportions of both species as treatments. Differences were found in water transparency, phytoplankton biomass, crustacean abundance, individual size and biomass. Our results suggest that visual predators intensify eutrophication effects more strongly than filter feeders do.     

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.     

Temperature and prey density jointly influence trophic and non‐trophic interactions in multiple predator communities

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First feeding diet of young brown trout fry in a temperate area: disentangling constraints and food selection

Trophic interactions are important factors structuring animal communities. We assessed the trophic relations of four fish species that live in sympatry in the River Ladra (NW Spain), and cluster analysis differentiated two feeding strategies: (1) species with omnivorous feeding habits, feeding mainly on detritus and plant material but with aquatic macroinvertebrates as an important complement (Achondrostoma arcasii and Pseudochondrostoma duriense) and (2) species feeding mainly (Salmo trutta) or exclusively (Gasterosteus gymnurus) on aquatic macroinvertebrates. Concerning ingested macroinvertebrates, the trophic overlap was quantified using Schoener’s index and the results obtained revealed a high diet overlap among the species (from 81.3 up to 99.2%). In order to get a deeper insight into mechanisms of fish species coexistence, we used ten biological and ecological traits of macroinvertebrate prey to discriminate feeding preferences. As a result, despite the high similarity among the diets, our analyses suggest that differences in diel activity patterns and drift behaviour of preys, as well as differences in the prey size, are important adaptive features that may reduce the inter-specific competition in the fish community and permit the partitioning of food that allows coexistence.     

Community structure and food web based on stable isotopes (δN and δC) analysis of a North Eastern Atlantic maerl bed

Maerl beds are highly biodiverse biogenic substrata that have been receiving increasing attention in the last decade. Although maerl beds represent important nursery areas for commercial fishes and molluscs, little is known on the trophic web of their communities. Community structure parameters of maerl bed of the Bay of Brest (species richness, abundance, biomass and dominating species) were studied in parallel with the carbon and nitrogen isotopic composition of their main benthic species (macrofaunal, and megafaunal organisms) in order to assess the trophic levels and differences in the potential food sources of maerl inhabitants. The major potential sources of energy were identified to originate either from epiphytic macroalgae and microphytobenthos both growing on maerl thalli, together with sedimenting (sedimentary) particulate organic matter (POM) originating from the water column. The majority of the macro- and megafaunal organisms investigated were filter feeders, selective-deposit feeders and predators/scavengers. Filter feeders fall into three different groups representing different trophic pathways (i) sponges feeding directly on POM (water column filter feeders I), (ii) ascidians and holothurians feeding on POM and probably captured pelagic preys (water column filter feeders II), and (iii) filter feeding molluscs and crustaceans were hypothesised to feed on microphytobenthos or on decaying sedimented POM (Interface filter feeders). Selective deposit feeders were also divided into two subgroups. Carnivores were also distinguished between those with scavenging habits and true predators. Coupling of the trophic levels observed with the community biomass structure revealed that most of the benthic biomass derives its food from detritic sedimented POM and/or microphytobenthos, with interface filter feeders (23% of the biomass), selective deposit feeders (12%). Carnivores made up to 14% of the total biomass. Generally stable isotopes ratio mean values overlap and cover a large range within feeding types, indicating a strong overlap in food sources and a high degree of complexity of the food web presumably due to the diversity of the potential food sources.     

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.     

Biomanipulation and its feasibility for water quality management in shallow eutrophic water bodies in The Netherlands

Biomanipulation as a tool for lake restoration is discussed mainly using literature data. It is based on the exploitation of the interactions both within and between the trophic levels in an aquatic ecosystem. Important among the interactions are: competition for light and nutrients between aquatic macrophytes and phytoplankton and among different phytoplankton species; grazing by planktonic and benthic filter feeders; and size-selective predation by fish. In several case studies biomanipulation has proved to be successful in restorating mildly eutrophic small waterbodies. However, for long-term stability of the restored ecosystems supplementary measures like reducing the external nutrient loadings are needed. The feasibility of the different biomanipulation measures to improve the water quality in shallow Dutch lakes is discussed. Preliminary results on biomanipulation experiments in enclosures withOscillatoria agardhii and the benthic filter feederDreissena polymorpha are given.     

Feeding habits and seasonal trophic guilds structuring fish community in the bay mouth region of a tropical estuarine habitat

Understanding trophic relationships of fish in estuarine ecosystem is an important element for sustainable resource management. This study examined the feeding habits of 29 dominant fish species, characterized the trophic guilds, assessed the impact of season and clarified the role of diets in structuring the fish community in the mouth region of Pattani Bay, Thailand. Samples of 5792 fishes collected monthly by gillnets from March 2019 to February 2020 were used for stomach content analyses. It was found that the number of food types and fullness index differed between fish taxa (P < 0.001). Most fishes were specialist feeders feeding on specific food components and were categorized into five trophic guilds: piscivore, shrimp-fish feeder, polychaete feeder, zooplanktivore and planktivore. Six species were piscivorous, considered as apex predators, that fed almost entirely on fishes. High diet overlaps among some species (>0.6) were recorded. Not much variation in seasonal guilds was observed: four guilds in the dry season, three in the moderate rainy season and four in the rainy season. Some species remained in the same guild the whole year round, but some fishes changed seasonally. Two fish communities from different regions of the bay were segregated based on feeding habits. The inner bay community comprised mainly copepod and plankton feeders, but there were more piscivores in the deeper bay mouth area. Results from this study help us to understand the feeding habits and trophic guilds of dominant fish species at the mouth of this tropical estuarine bay.     

The effects of species properties and community context on establishment success

Understanding whether factors important for species establishment in a local community are predictable or context‐ dependent is key for determining the features that affect community stability and species coexistence. A major challenge for scientists addressing this question is that natural systems are complex. This makes it difficult to test multiple properties of species and features of the resident community simultaneously to determine what factors are most important for establishment success of a species into a novel community. We used the model aquatic system inside the leaves of the pitcher plant Sarracenia purpurea to test whether properties predicted to be important for establishment success of a species (initial density, competitive ability, body size) are generalizable across communities varying in resource availability and the presence of a top predator. For intermediate trophic‐level species, we found that both competitive dominance and initial density were important for establishment success. Although a less competitive species was also able to successfully establish in the communities, high resource availability and high initial density were important for its establishment success. Body size of the introduced species, although correlated with competitive ability, was not an important characteristic for establishment success. The presence of a top predator significantly decreased the densities of the introduced species when resources were low, but did not completely inhibit establishment success. The relative importance of each of these factors, and interactions among them, could not have been discerned through single hypothesis testing. The results from this work show the need for detailed experiments that focus on combinations of factors to understand if mechanisms determining community assembly and species establishment can be generalized across systems.     

Cascading effects of belowground predators on plant communities are density‐dependent

Soil food webs comprise a multitude of trophic interactions that can affect the composition and productivity of plant communities. Belowground predators feeding on microbial grazers like Collembola could decelerate nutrient mineralization by reducing microbial turnover in the soil, which in turn could negatively influence plant growth. However, empirical evidences for the ecological significance of belowground predators on nutrient cycling and plant communities are scarce. Here, we manipulated predator density (Hypoaspis aculeifer: predatory mite) with equal densities of three Collembola species as a prey in four functionally dissimilar plant communities in experimental microcosms: grass monoculture (Poa pratensis), herb monoculture (Rumex acetosa), legume monoculture (Trifolium pratense), and all three species as a mixed plant community. Density manipulation of predators allowed us to test for density‐mediated effects of belowground predators on Collembola and lower trophic groups. We hypothesized that predator density will reduce Collembola population causing a decrease in nutrient mineralization and hence detrimentally affect plant growth. First, we found a density‐dependent population change in predators, that is, an increase in low‐density treatments, but a decrease in high‐density treatments. Second, prey suppression was lower at high predator density, which caused a shift in the soil microbial community by increasing the fungal: bacterial biomass ratio, and an increase of nitrification rates, particularly in legume monocultures. Despite the increase in nutrient mineralization, legume monocultures performed worse at high predator density. Further, individual grass shoot biomass decreased in monocultures, while it increased in mixed plant communities with increasing predator density, which coincided with elevated soil N uptake by grasses. As a consequence, high predator density significantly increased plant complementarity effects indicating a decrease in interspecific plant competition. These results highlight that belowground predators can relax interspecific plant competition by increasing nutrient mineralization through their density‐dependent cascading effects on detritivore and soil microbial communities.     

Trophic and functional cascades in tropical versus temperate aquatic microcosms

Inverse trophic cascades are a well explored and common consequence of the local depletion or extinction of top predators in natural ecosystems. Despite a large body of research, the cascading effects of predator removal on ecosystem functions are not as well understood. Developing microcosm experiments, we explored food web changes in trophic structure and ecosystem functioning following biomass removal of top predators in representative temperate and tropical rock pool communities that contained similar assemblages of zooplankton and benthic invertebrates. We observed changes in species abundances following predator removal in both temperate and tropical communities, in line with expected inverse effects of a trophic cascade, where predation release benefits the predator’s preys and competitors and impacts the preys of the latter. We also observed several changes at the community and ecosystem levels including a decrease in total abundance and mean trophic level of the community, and changes in chlorophyll-a and total dissolved particles. Our results also showed an increase in variability of both community and ecosystem processes following the removal of predators. These results illustrate how predator removal can lead to inverse trophic cascades both in structural and functioning properties, and can increase variability of ecosystem processes. Although observed patterns were consistent between tropical and temperate communities following an inverse cascade pattern, changes were more pronounced in the temperate community. Therefore, aquatic food webs may have inherent traits that condition ecosystem responses to changes in top-down trophic control and render some aquatic ecosystems especially sensitive to the removals of top predators.     

Trophic analysis and trophodynamic functioning of mangrove fish fauna of New Caledonia]

Fish communities from geomorphologically different mangrove forests showed distinct trophic structures. A mangrove area located near fringing reefs (Bouraké) was dominated by large invertebrate (> 2 mm) feeders (40.7% of total wet weight of the fish fauna) and herbivores (26.7%) whilst an estuarine mangrove (Ouenghi) was characterized by detritus feeders (28.2%), piscivorous (18.2%) and large invertebrate feeders (17.9%). In spite of these differences in trophic structure, similar food webs occurred in both areas, whereas the intensity of fluxes between trophic compartments was different. Resident species were usually at the base of the trophic structure. This component of the fish fauna used available sources of energy, such as microalgae in Bouraké or detritus and phytoplankton in Ouenghi. In contrast, transient species were high level predators, mainly piscivorous and large invertebrate feeders. These species actively contributed to net exports of energy from mangrove areas to nearby coastal habitats. Food webs and energy fluxes associated with trophic migration of fishes were particularly important in non-estuarine mangrove forests because of hydrologic conditions (salinity and turbidity) which were more suitable to the invasion of numerous marine foraging species (carangids, lutjanids, sphyraenids).     

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.     

Coexistence of two termite-eating specialists (Araneae)

1. Competition among closely-related specialist predators has rarely been studied, and thus the mechanism of their coexistence remains enigmatic. Interspecific competition among specialised co-occurring predators capturing termites should be high.
2. Here we investigated various niche dimensions, namely temporal, spatial and trophic, of a couple of jumping spider species of the genus Stenaelurillus (Stenaelurillus guttiger and S. modestus) from South Africa, to find whether these two species co-exist and along which niche dimension(s) they differentiate.
3. The two species co-occurred in two out of five study sites. Body size was not significantly different between the species. The phenology was shifted so that one species matured earlier. Circadian activity was not different, as both species were diurnal and active at similar times. Both species preyed almost exclusively on termites. The fundamental trophic niche was very similar and rather narrow. The realised trophic niche at the prey order level of both species was similar, but at the genus level it was different. In S. modestus it was narrower, as it captured mainly Odontotermes, while S. guttiger exploited a few termite species. The size of prey captured was also similar between the two species. The frequency of intraguild predation was negligible.
4. We conclude that both Stenaelurillus species are specialised termitophagous predators. The two species can coexist across broad spatial scales due to spatial segregation on the landscape. At the sites where they co-occur, the two species specialise on different termite prey, promoting local coexistence.

     

Trophic Structure of Benthic Macroinvertebrate Communities from Lakes with Various Water-Salinity Levels

The trophic structure of benthic macroinvertebrate communities in lakes varying in salinity levels (from oligohaline to hyperhaline) in the southern Ob–Irtysh interfluve has been investigated. Four trophic groups of macroinvertebrates have been identified in 48 lakes: (1) predators, (2) grinders, (3) scrapers, and (4) collectors–detritophages and facultative filter feeders. It is found that the proportion of different trophic groups in taxonomic composition and biomass of macroinvertebrate communities changes with increasing water salinity in lakes.     

Diet composition,feeding niche partitioning and trophic organisation of large pelagic predatory fishes in the eastern Arabian Sea

Information on the ecology and feeding behaviour of the large oceanic predatory fishes is crucial for the ecosystem approaches to fisheries management models. Co-existing large pelagic predators in the open oceans may avoid competition for the limited forage by resource partitioning on spatial, temporal or trophic levels. To test this, we studied the prey species composition, diet overlap, trophic level, and trophic organisation of 12 large predatory fishes co-existing in the eastern Arabian Sea. Stomach contents of 1,518 specimens caught by exploratory longline operations in the Indian Exclusive Economic Zone during the years 2006–2009 were analysed. Finfishes were dominant prey of all species except blue marlin (Makaira nigricans) and yellowfin tuna (Thunnus albacares), which fed mainly on cephalopods, and long-snouted lancetfish (Alepisaurus ferox) and pelagic stingray (Pteroplatytrygon violacea), which fed mainly on crustaceans. Common dolphinfish (Coryphaena hippurus) and yellowfin tuna fed on a wider variety of prey than the other species, while the diets of lancetfish and black marlin (Istiompax indica) were narrowest. Pelagic stingray and great barracuda (Sphyraena barracuda) fed on species occupying epipelagic waters, whereas the contribution of mesopelagic prey was higher in the diets of swordfish (Xiphias gladius) and pelagic thresher (Alopias pelagicus). Trophic levels of these fishes ranged from 4.13 to 4.37. Diet overlap index revealed that some of the large pelagic predatory fishes share common prey species. Cluster analysis of the diets revealed four distinct trophic guilds namely ‘flyingfish feeders’ (common dolphinfish and great barracuda); ‘mesopelagic predators’ (pelagic thresher and swordfish); ‘crab feeders’ (lancetfish, pelagic stingray and silky shark) and ‘squid feeders’ (yellowfin tuna, Indo-Pacific sailfish (Istiophorus platypterus), skipjack tuna (Katsuwonus pelamis), black marlin and blue marlin). Large predatory fishes of the eastern Arabian Sea target different prey types, and limit their vertical extent and time of feeding to avoid competing for prey.     

Larval fish-induced phenotypic plasticity of coexisting Daphnia: an enclosure experiment

1. The indirect effects of predators on lower trophic levels have been studied without much attention to phenotypically plastic traits of key food web components. Phenotypic plasticity among species creates phenotypic diversity over a changing environmental landscape. 2. We measured the indirect effects of planktivorous larval walleye (Stizostedion vitreum) on phytoplankton biomass through their effects on the dominant herbivore species, Daphnia pulicaria and D. mendotae. 3. Fish had no effect on phytoplankton biomass or overall Daphnia density. We observed a compensatory response to predation by functionally comparable species within a trophic level in the form of shifting dominance and coexistence of Daphnia species. We hypothesized that this phenotypically plastic response to predation decoupled a potential trophic cascade in this freshwater pelagic system. Daphnia pulicaria density decreased over time with fish predation, but D. mendotae density increased over time with fish predation. 4. Phenotypically plastic life history trait shifts and reproductive rates differed between species in fishless and fish enclosures, accounting for population trends. Daphnia pulicaria were also proportionally higher in walleye larvae stomachs than in the enclosures, indicating that walleye preferred to feed on D. pulcaria over D. mendotae. The resultant shift in dominance may partially explain the overall benign effect of fish on grazers and supports the hypothesis that trophic level diversity can decouple a trophic cascade.     

Effects of an invasive plant transcend ecosystem boundaries through a dragonfly-mediated trophic pathway

Trophic interactions can strongly influence the structure and function of terrestrial and aquatic communities through top-down and bottom-up processes. Species with life stages in both terrestrial and aquatic systems may be particularly likely to link the effects of trophic interactions across ecosystem boundaries. Using experimental wetlands planted with purple loosestrife (Lythrum salicaria), we tested the degree to which the bottom-up effects of floral density of this invasive plant could trigger a chain of interactions, changing the behavior of terrestrial flying insect prey and predators and ultimately cascading through top-down interactions to alter lower trophic levels in the aquatic community. The results of our experiment support the linkage of terrestrial and aquatic food webs through this hypothesized pathway, with high loosestrife floral density treatments attracting high levels of visiting insect pollinators and predatory adult dragonflies. High floral densities were also associated with increased adult dragonfly oviposition and subsequently high larval dragonfly abundance in the aquatic community. Finally, high-flower treatments were coupled with changes in zooplankton species richness and shifts in the composition of zooplankton communities. Through changes in animal behavior and trophic interactions in terrestrial and aquatic systems, this work illustrates the broad and potentially cryptic effects of invasive species, and provides additional compelling motivation for ecologists to conduct investigations that cross traditional ecosystem boundaries.