Pitcher Plant Sarracenia Purpurea Research Articles

Dispersal Limitation Governs Bacterial Community Assembly in the Northern Pitcher Plant (Sarracenia purpurea) at the Continental Scale

ABSTRACTAimEcological theory suggests that dispersal limitation and selection by climatic factors influence bacterial community assembly at a continental scale, yet the conditions governing the relative importance of each process remains unclear. The carnivorous pitcher plant Sarracenia purpurea provides a model aquatic microecosystem to assess bacterial communities across the host plant's north–south range in North America. This study determined the relative influences of dispersal limitation and environmental selection on the assembly of bacterial communities inhabiting S. purpurea pitchers at the continental scale.LocationEastern United States and Canada.Time Period2016.Major Taxa StudiedBacteria inhabiting S. purpurea pitchers.MethodsPitcher morphology, fluid, inquilines and prey were measured, and pitcher fluid underwent DNA sequencing for bacterial community analysis. Null modelling of β‐diversity provided estimates for the contributions of selection and dispersal limitation to community assembly, complemented by an examination of spatial clustering of individuals. Phylogenetic and ecological associations of co‐occurrence network module bacteria was determined by assessing the phylogenetic diversity and habitat preferences of member taxa.ResultsDispersal limitation was evident from between‐site variation and spatial aggregation of individual bacterial taxa in the S. purpurea pitcher system. Selection pressure was weak across the geographic range, yet network module analysis indicated environmental selection within subgroups. A group of aquatic bacteria held traits under selection in warmer, wetter climates, and midge abundance was associated with selection for traits held by a group of saprotrophs. Processes that increased pitcher fluid volume weakened selection in one module, possibly by supporting greater bacterial dispersal.ConclusionDispersal limitation governed bacterial community assembly in S. purpurea pitchers at a continental scale (74% of between‐site comparisons) and was significantly greater than selection across the range. Network modules showed evidence for selection, demonstrating that multiple processes acted concurrently in bacterial community assembly at the continental scale.

Abstract 154: Inhibition of Staphylococcus aureus SsbA by the stem extract of the pitcher plant Sarracenia purpurea

Abstract 154: Inhibition of Staphylococcus aureus SsbA by the stem extract of the pitcher plant Sarracenia purpurea

Prey capture by the non-native carnivorous pitcher plant Sarracenia purpurea across sites in Britain and Ireland.

The carnivorous pitcher plant Sarracenia purpurea is native to North America, but has been introduced into Europe, where it is now widespread. Understanding of how this species functions in its non-native range is limited. We measured pitcher morphology and prey capture by S. purpurea in its non-native range in Britain and Ireland. Pitchers were removed from different plants at each of six bogs covering the species range in Britain and Ireland (n=10 pitchers per site). For each pitcher we counted and identified every prey item and took measurements of morphology. We also compiled prey capture data for existing studies in Europe and North America. Prey capture characteristics varied between sites in Britain and Ireland. The amount of prey captured varied 20-fold between sites and was partially explained by differences in pitcher size; larger pitchers caught more prey. The primary prey was Formicidae, Diptera and Coleoptera. At the rank of order, prey composition varied between bogs, some contained mainly Formicidae, some mainly Diptera and some a mix. Prey capture was less evenly distributed at some bogs compared to others, suggesting more specialization. There was no overall difference in prey capture (composition or evenness) at the rank of order between plants in Europe compared to those in North America. At the rank of species, prey capture varied between populations even within the same order. This study demonstrates a large amount of variability between sites in prey capture characteristics. This may reflect different site characteristics and/or plant strategies, which will likely impact plant function, and may impact the inquiline community. In terms of prey capture at the rank of order, S. purpurea functions identically in its non-native range. This supports its use as a model system in a natural experiment for understanding food webs.

Effects of Community Richness and Competitive Asymmetry on Protozoa Evolution in Sarracenia purpurea Leaves.

AbstractPredicting evolution in natural systems will require understanding how selection operates in multispecies communities. We predicted that the amount that traits evolve in multispecies mixtures would be less than the amount that would be predicted from the additive contributions of the pairwise interactions and that subordinate species will be more likely to evolve in competitive systems than dominant species. We conducted an experimental test of these predictions using a guild of protozoans found in the water-filled leaves of the pitcher plant Sarracenia purpurea. The response to selection did not significantly change as we increased richness from monocultures to two- and four-species mixtures. In accordance with our second prediction, subordinate species demonstrated greater growth in competition after selection than before, while dominant species generally showed no response to selection. Monod-type experiments to determine minimum resource levels found that the dominant species had much higher resource requirements than the subordinate species and that the minimum resource requirements evolved to be higher in the subordinate species. Importantly, these results suggest that subordinate species evolve to become more similar to dominant species, which may involve resource use convergence. Our findings and other recent works suggest that community diversity can affect evolution in surprising ways that warrant further investigation.

Anticancer and Antioxidant Activities of the Root Extract of the Carnivorous Pitcher Plant Sarracenia purpurea.

The carnivorous pitcher plant Sarracenia purpurea exhibits many ethnobotanical uses, including the treatments of type 2 diabetes and tuberculosis-like symptoms. In this study, we prepared different extracts from the leaves (pitchers), stems, and roots of S. purpurea and investigated their antioxidant and anticancer properties. To evaluate the extraction efficiency, we individually used different solvents, namely methanol, ethanol, acetone, and distilled water, for S. purpurea extract preparations. The root extract of S. purpurea, obtained by 100% acetone (S. purpurea-root-acetone), had the highest anticancer activities, antioxidation capacity (the DPPH activity with IC50 of 89.3 ± 2.2 μg/mL), antibacterial activities, total phenolic content (33.4 ± 0.7 mg GAE/g), and total flavonoid content (107.9 ± 2.2 mg QUE/g). The most abundant compounds in S. purpurea-root-acetone were identified using gas chromatography–mass spectrometry; 7,8-Dihydro-α-ionone was the major compound present in S. purpurea-root-acetone. In addition, the co-cytotoxicity of S. purpurea-root-acetone (combined with the clinical anticancer drug 5-fluorouracil (5-FU) on the survival, apoptosis, proliferation, and migration of the 4T1 mammary carcinoma) was examined. The combination of 5-FU with S. purpurea-root-acetone could be highly efficient for anti-4T1 cells. We also found that S. purpurea-root-acetone could inhibit the enzymatic activity of human dihydroorotase (huDHOase), an attractive target for potential anticancer chemotherapy. The sic most abundant compounds in S. purpurea-root-acetone were tested using an in silico analysis via MOE-Dock software for their binding affinities. The top-ranked docking conformations were observed for 7,8-dihydro-α-ionone and stigmast-5-en-3-ol, suggesting the inhibition potential against huDHOase. Overall, the collective data in this study may indicate the pharmacological potentials of S. purpurea-root-acetone for possible medical applications.

Bacterial Recruitment to Carnivorous Pitcher Plant Communities: Identifying Sources Influencing Plant Microbiome Composition and Function.

Processes influencing recruitment of diverse bacteria to plant microbiomes remain poorly understood. In the carnivorous pitcher plant Sarracenia purpurea model system, individual pitchers open to collect rainwater, invertebrates and a diverse microbial community, and this detrital food web is sustained by captured insect prey. This study examined how potential sources of bacteria affect the development of the bacterial community within pitchers, how the host plant tissue affects community development and how established vs. assembling communities differ. In a controlled greenhouse experiment, seven replicate pitchers were allocated to five treatments to exclude specific bacterial sources or host tissue: milliQ water only, milliQ + insect prey, rainwater + prey, established communities + prey, artificial pitchers with milliQ + prey. Community composition and functions were examined over 8–40 weeks using bacterial gene sequencing and functional predictions, measurements of cell abundance, hydrolytic enzyme activity and nutrient transformations. Distinct community composition and functional differences between artificial and real pitchers confirm an important influence of host plant tissue on community development, but also suggest this could be partially related to host nutrient uptake. Significant recruitment of bacteria to pitchers from air was evident from many taxa common to all treatments, overlap in composition between milliQ, milliQ + prey, and rainwater + prey treatments, and few taxa unique to milliQ only pitchers. Community functions measured as hydrolytic enzyme (chitinase, protease) activity suggested a strong influence of insect prey additions and were linked to rapid transformation of insect nutrients into dissolved and inorganic sources. Bacterial taxa found in 6 of 7 replicate pitchers within treatments, the “core microbiome” showed tighter successional trajectories over 8 weeks than all taxa. Established pitcher community composition was more stable over 8 weeks, suggesting a diversity-stability relationship and effect of microinvertebrates on bacteria. This study broadly demonstrates that bacterial composition in host pitcher plants is related to both stochastic and specific bacterial recruitment and host plants influence microbial selection and support microbiomes through capture of insect prey.

Interactions between strains govern the eco-evolutionary dynamics of microbial communities.

Genomic data has revealed that genotypic variants of the same species, that is, strains, coexist and are abundant in natural microbial communities. However, it is not clear if strains are ecologically equivalent, and at what characteristic genetic distance they might exhibit distinct interactions and dynamics. Here, we address this problem by tracking 10 taxonomically diverse microbial communities from the pitcher plant Sarracenia purpurea in the laboratory for more than 300 generations. Using metagenomic sequencing, we reconstruct their dynamics over time and across scales, from distant phyla to closely related genotypes. We find that most strains are not ecologically equivalent and exhibit distinct dynamical patterns, often being significantly more correlated with strains from another species than their own. Although even a single mutation can affect laboratory strains, on average, natural strains typically decouple in their dynamics beyond a genetic distance of 100 base pairs. Using mathematical consumer-resource models, we show that these taxonomic patterns emerge naturally from ecological interactions between community members, but only if the interactions are coarse-grained at the level of strains, not species. Finally, by analyzing genomic differences between strains, we identify major functional hubs such as transporters, regulators, and carbohydrate-catabolizing enzymes, which might be the basis for strain-specific interactions. Our work suggests that fine-scale genetic differences in natural communities could be created and stabilized via the rapid diversification of ecological interactions between strains.

Regulation by the Pitcher Plant Sarracenia purpurea of the Structure of its Inquiline Food Web

Phytotelmata, the water-filled habitats in pitcher plants, bromeliad tanks, and tree-holes, host multitrophic food webs that are model experimental systems for studying food-web structure and dynamics. However, the plant usually is considered simply as an inert container, not as an interacting part of the food web. We used a manipulative field experiment with a response-surface design to determine effects of nutrient enrichment (multiple levels of NH4NO3, PO4, and captured prey), top predators (removed or present), and the plant itself (with or without plastic tubes inserted into the pitchers to isolate the food web from the plant) on the macrobial food web within the modified leaves (“pitchers”) of the carnivorous pitcher plant Sarracenia purpurea. Connection to the plant, addition of NH4NO3, and removal of the top predator significantly increased the food web's saturation, defined as its trophic depth and number of interactions. No effects on food-web saturation resulted from addition of PO4 or supplemental prey. Plants such as S. purpurea that create phytotelmata are more than inert containers and their inhabitants are more than commensal inquilines. Rather, both the plant and the inquilines are partners in a complex network of interactions.

Environment-host-microbial interactions shape the Sarracenia purpurea microbiome at the continental scale.

The importance of climate, habitat structure, and higher trophic levels on microbial diversity is only beginning to be understood. Here, we examined the influence of climate variables, plant morphology, and the abundance of aquatic invertebrates on the microbial biodiversity of the northern pitcher plant Sarracenia purpurea. The plant's cup-shaped leaves fill with rainwater and support a miniature, yet full-fledged, ecosystem with a diverse microbiome that decomposes captured prey and a small network of shredding and filter-feeding aquatic invertebrates that feed on microbes. We characterized pitcher microbiomes of 108 plants sampled at 36 sites from Florida to Quebec. Structural equation models revealed that annual precipitation and temperature, plant size, and midge abundance had direct effects on microbiome taxonomic and phylogenetic diversity. Climate variables also exerted indirect effects through plant size and midge abundance. Further, spatial structure and climate influenced taxonomic composition, but not phylogenetic composition. Our results suggest that direct effects of midge abundance and climate and indirect effects of climate through its effect on plant-associated factors lead to greater richness of microbial phylotypes in warmer, wetter sites.

Clockwise and counterclockwise hysteresis characterize state changes in the same aquatic ecosystem

Incremental increases in a driver variable, such as nutrients or detritus, can trigger abrupt shifts in aquatic ecosystems that may exhibit hysteretic dynamics and a slow return to the initial state. A model system for understanding these dynamics is the microbial assemblage that inhabits the cup-shaped leaves of the pitcher plant Sarracenia purpurea. With enrichment of organic matter, this system flips within three days from an oxygen-rich state to an oxygen-poor state. In a replicated greenhouse experiment, we enriched pitcher-plant leaves at different rates with bovine serum albumin (BSA), a molecular substitute for detritus. Changes in dissolved oxygen (DO) and undigested BSA concentration were monitored during enrichment and recovery phases. With increasing enrichment rates, the dynamics ranged from clockwise hysteresis (low), to environmental tracking (medium), to novel counter-clockwise hysteresis (high). These experiments demonstrate that detrital enrichment rate can modulate a diversity of hysteretic responses within a single aquatic ecosystem, and suggest different management strategies may be needed to mitigate the effects of high vs. low rates of detrital enrichment.

Diverse microbial communities hosted by the model carnivorous pitcher plant Sarracenia purpurea: analysis of both bacterial and eukaryotic composition across distinct host plant populations.

BackgroundThe pitcher plant Sarracenia purpurea supplements nutrient acquisition through carnivory, capturing insect prey which are digested by a food web community of eukaryotes and bacteria. While the food web invertebrates are well studied, and some recent studies have characterized bacteria, detailed genetic analysis of eukaryotic diversity is lacking. This study aimed to compare eukaryotic and bacterial composition and diversity of pitcher communities within and between populations of host plants in nearby but distinct wetland habitats, and to characterize microbial functions across populations and in comparison with another freshwater community.MethodsPitcher fluid was sampled from the two wetlands, Cedarburg and Sapa Bogs, community DNA was extracted, and 16S and 18S rRNA amplicons were sequenced and data processed for community-level comparisons.Results and ConclusionsBacterial diversity in the small pitcher volume rivaled that of larger aquatic communities. Between pitcher plant populations, several bacterial families (Kiloniellaceae, Acetobacteraceae, Xanthobacteraceae, Sanguibacteraceae, Oligoflexaceae, Nitrosomonadaceae, Chromatiaceae, Saprospiraceae) were significantly higher in one population. However, although predicted pitcher bacterial functions were distinct from other freshwater communities, especially for some amino acid metabolism, functions were similar across all the pitchers in the two populations. This suggests some functional redundancy among bacterial taxa, and that functions converge to achieve similar food web processes. The sequencing identified a previously under-appreciated high diversity of ciliates, Acari mites, fungi and flagellates in pitcher communities; the most abundant sequences from eukaryotic taxa were Oligohymenophorea ciliates, millipedes and Ichthyosporea flagellates. Two thirds of taxa were identified as food web inhabitants and less than one third as prey organisms. Although eukaryotic composition was not significantly different between populations, there were different species of core taxonomic groups present in different pitchers—these differences may be driven by wetland habitats providing different populations to colonize new pitchers. Eukaryotic composition was more variable than bacterial composition, and there was a poor relationship between bacterial and eukaryotic composition within individual pitchers, suggesting that colonization by eukaryotes may be more stochastic than for bacteria, and bacterial recruitment to pitchers may involve factors other than prey capture and colonization by eukaryotic food web inhabitants.

DNA metabarcoding reveals changes in the contents of carnivorous plants along an elevation gradient.

Resource variation along abiotic gradients influences subsequent trophic interactions and these effects can be transmitted through entire food webs. Interactions along abiotic gradients can provide clues as to how organisms will face changing environmental conditions, such as future range shifts. However, it is challenging to find replicated systems to study these effects. Phytotelmata, such as those found in carnivorous plants, are isolated aquatic communities and thus form a good model for the study of replicated food webs. Due to the degraded nature of the prey, molecular techniques provide a useful tool to study these communities. We studied the pitcher plant Sarracenia purpurea L. in allochthonous populations along an elevational gradient in the Alps and Jura. We predicted that invertebrate richness in the contents of the pitcher plants would decrease with increasing elevation, reflecting harsher environmental conditions. Using metabarcoding of the COI gene, we sequenced the invertebrate contents of these pitcher plants. We assigned Molecular Operational Taxonomic Units at ordinal level as well as recovering species-level data. We found small but significant changes in community composition with elevation. These recovered sequences could belong to invertebrate prey, rotifer inquilines, pollinators and other animals possibly living inside the pitchers. However, we found no directional trend or site-based differences in MOTU richness with elevational gradient. Use of molecular techniques for dietary or contents analysis is a powerful way to examine numerous degraded samples, although factors such as DNA persistence and the relationship with species presence still have to be completely determined.

Exploring the predation of UK bumblebees (Apidae, Bombus spp.) by the invasive pitcher plant Sarracenia purpurea: examining the effects of annual variation, seasonal variation, plant density and bumblebee gender

Invasive carnivorous plant species can impact the native invertebrate communities on which they prey. This article explores the predation of native UK bumblebees (Bombus spp.) by the invasive pitcher plant species Sarracenia purpurea and discusses the potential effect of S. purpurea on native bumblebees. Specifically, it evaluates whether the extent to which bumblebees are captured varies (i) over successive years, (ii) across June and July, (iii) with density of distribution of pitchers or (iv) with bumblebee gender. Pitcher contents were examined from an established population of Sarracenia purpurea growing in Dorset, UK. Results show that the total extent to which bumblebees were captured differed over the years 2012–2014 inclusive. A 1-year study in 2013 showed that more bumblebees were caught in July than in June and more bumblebees were captured when pitchers grew at high density. Results from 2013 also showed that more pitchers caught more than one bumblebee than would be expected based on a normal probability distribution and that this phenomenon affects female and male bumblebees equally. We discuss possible reasons for these results including that the bumblebees may be using S. purpurea as a resource. Further work is required to establish the exact underpinning mechanisms and the relative roles of plant and bumblebee behaviour within the relationship. Such interaction complexity may have consequences for consideration in invasive carnivorous plant management.

Mycobacterium sarraceniae sp. nov. and Mycobacterium helvum sp. nov., isolated from the pitcher plant Sarracenia purpurea.

Several fast- to intermediate-growing, acid-fast, scotochromogenic bacteria were isolated from Sarracenia purpurea pitcher waters in Minnesota sphagnum peat bogs. Two strains (DL734T and DL739T) were among these isolates. On the basis of 16S rRNA gene sequences, the phylogenetic positions of both strains is in the genus Mycobacterium with no obvious relation to any characterized type strains of mycobacteria. Phenotypic characterization revealed that neither strain was similar to the type strains of known species of the genus Mycobacterium in the collective properties of growth, pigmentation or fatty acid composition. Strain DL734T grew at temperatures between 28 and 32 °C, was positive for 3-day arylsulfatase production, and was negative for Tween 80 hydrolysis, urease and nitrate reduction. Strain DL739T grew at temperatures between 28 and 37 °C, and was positive for Tween 80 hydrolysis, urea, nitrate reduction and 3-day arylsulfatase production. Both strains were catalase-negative while only DL739T grew with 5 % NaCl. Fatty acid methyl ester profiles were unique for each strain. DL739T showed an ability to survive at 8 °C with little to no cellular replication and is thus considered to be psychrotolerant. Therefore, strains DL734T and DL739T represent two novel species of the genus Mycobacterium with the proposed names Mycobacterium sarraceniae sp. nov. and Mycobacterium helvum sp. nov., respectively. The type strains are DL734T (=JCM 30395T=NCCB 100519T) and DL739T (=JCM 30396T=NCCB 100520T), respectively.

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 plantSarracenia 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.

Species Composition and Seasonal Distribution of Mosquito Larvae (Diptera: Culicidae) in Southern New Jersey, Burlington County.

A total of 36,495 larvae consisting of 45 species from 11 genera were collected from 7,189 sites from southern New Jersey, Burlington County between the months of March and October, 2001-2014. Density and seasonal distribution were determined among natural and artificial habitat. The most dominant species collected from natural habitat was Aedes vexans (Meigen) followed by Ochlerotatus canadensis canadensis (Theobald), Culex restuans Theobald, Culex pipiens L., and Culex territans Walker. The most dominant species collected from artificial habitat was Aedes albopictus (Skuse) followed by Ochlerotatus japonicus japonicus (Theobald), Cx. restuans, Cx. pipiens, and Ochlerotatus triseriatus (Say). Cx. restuans and Cx. pipiens were the only species categorized as dominant among both natural and artificial habitat and comprised greater than half the total density. Sympatry was common among dominant species from artificial habitat where a significant percentage of the total collection contained multiple species. The most common types of natural habitats were forested depressions and stream flood plains whereas rimless vehicle tires and various plastic containers were the most common artificial habitats. The pitcher plant Sarracenia purpurea L. was the only habitat exclusive to one species.

Carnivorous pitcher plant species (Sarracenia purpurea) increases root growth in response to nitrogen addition

Nitrogen (N) deposition from anthropogenic sources can facilitate the encroachment of plant species with high-N demands into nutrient-poor ecosystems such as sphagnum bogs. Prior research has demonstrated that altered leaf morphology of the carnivorous pitcher plant Sarracenia purpurea L. can serve as a biological indicator of increased bog nitrification. Our objective was to assess the effect of N addition on the root morphology of S. purpurea. To make this assessment, nine S. purpurea plants were grown in microcosms with their roots positioned on transparent acrylic tubes so that root growth could be monitored. Three replicate microcosms received either a high-N treatment (1.0 mg NH4-N·L−1), low-N treatment (0.1 mg NH4-N·L−1), or no additional N. After 7 weeks, we scanned the roots with WinRhizo Pro software, recorded leaf dimensions, and measured the dry mass of the roots and leaves. The high-N treatment had significantly greater root length, surface area, and dry biomass than the controls. In contrast, we found no difference in leaf dimensions or aboveground biomass among treatments. The results of this study support our hypothesis that S. purpurea increases root growth to uptake nutrients from the soil under conditions of increased N deposition.

Size Matters: Purple Pitcher Plant (Sarracenia purpurea) Pitcher Size Influences the Presence of Aedes albopictus Larvae

A vector of multiple arboviruses, the invasive Asian tiger mosquito Aedes albopictus was recently found to develop inside the pitchers of the purple pitcher plant Sarracenia purpurea in a forested environment. In this study larvae and pupae of Ae. albopictus were found in S. purpurea pitchers in a peridomestic habitat in central North Carolina. Evidence of direct oviposition on the inner sides of the pitchers was obtained by dissecting pitchers and examining the insides for eggs, which were then raised to adults. To determine if pitcher size and origin (same or different plant) affected the presence of immature Ae. albopictus, these variables were measured and compared against the presence/absence of larvae and/or pupae through a logistic regression analysis. Statistical evidence showed pitchers with larger openings were much more likely to contain larvae and/or pupae, and pitchers from the same plant were not significantly more often colonized than pitchers on other plants. These data, in combination with the presence of dead adult females in pitchers, suggest the morphology of some pitchers may prevent successful oviposition by gravid females.

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.

Predicting food‐web structure with metacommunity models

SynthesisMetacommunity 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.