Species Co-occurrence Networks Research Articles

The impact of gut microbiome enterotypes on ulcerative colitis: identifying key bacterial species and revealing species co-occurrence networks using machine learning

ABSTRACT Ulcerative colitis (UC) is a chronic inflammatory intestinal disease affecting the colon and rectum, with its pathogenesis attributed to genetic background, environmental factors, and gut microbes. This study aimed to investigate the role of enterotypes in UC by conducting a hierarchical analysis, determining differential bacteria using machine learning, and performing Species Co-occurrence Network (SCN) analysis. Fecal bacterial data were collected from UC patients, and a 16S rRNA metagenomic analysis was performed using the QIIME2 bioinformatics pipeline. Enterotype clustering was conducted at the family level, and deep neural network (DNN) classification models were trained for UC and healthy controls (HC) in each enterotype. Results from eleven 16S rRNA gut microbiome datasets revealed three enterotypes: Bacteroidaceae (ET-B), Lachnospiraceae (ET-L), and Clostridiaceae (ET-C). Ruminococcus (R. gnavus) abundance was significantly higher in UC subjects with ET-B and ET-C than in those with ET-L. R. gnavus also showed a positive correlation with Clostridia in UC SCN for ET-B and ET-C subjects, with a higher correlation in ET-C subjects. Conversely, Odoribacter (O.) splanchnicus and Bacteroides (B.) uniformis exhibited a positive correlation with tryptophan metabolism and AMP-activated protein kinase (AMPK) signaling pathways, while R. gnavus showed a negative correlation. In vitro co-culture experiments with Clostridium (C.) difficile demonstrated that fecal microbiota from ET-B subjects had a higher abundance of C. difficile than ET-L subjects. In conclusion, the ET-B enterotype predisposes individuals to UC, with R. gnavus as a potential risk factor and O. splanchnicus and B. uniformis as protective bacteria, and those with UC may have ultimately become ET-C.

Soil rare microorganisms mediated the plant cadmium uptake: The central role of protists

Plants and microorganisms symbiotically mediate and/or catalyse the turnover of elements in rhizosphere soils, thus directly influencing the effectiveness of phytoremediation in addressing heavy metal contamination. Soil rare microbial communities are diverse but not well understood in terms of their importance for phytoremediation. In this study, we simulated the loss of rare microorganisms through dilution-to-extinction approach, and investigated the effects on integrated rhizosphere microbiome with soil microcosm experiments, including bacteria, fungi, protists, and microfauna. Additionally, we explored the implications for ryegrass (Lolium multiflorum Lam.) growth and its uptake of Cd (cadmium). Compared with the undiluted group, ryegrass exhibited a significant decrease in Cd uptake ranging from 52.34 % to 73.71 % in the rare species-loss soils, indicating a lack of functional redundancy in rhizosphere soil microbial community following rare species loss. Interestingly, these soils displayed a remarkable 1.79-fold increase in plant biomass and a 41.02 % increase in plant height. By sequencing the 16S, 18S, and ITS rRNA gene amplicons of rhizosphere microbes, we found that soil rare species loss decreased the rhizosphere microbial α-diversity, changed the community structures, and shifted the functional potential. Protists were particularly affected. Through the analysis of species co-occurrence networks, along with the partial least squares path modeling, we found that the diversity of protists and bacteria and the co-occurring network connectivity of protists and fungi contributed most to plant Cd uptake and growth. These results highlighted the potential significance of rare microorganisms, particularly protists, in phytoextraction of Cd-contaminated soils, owing to their central role in the microbial food web.

Organic management increases beneficial microorganisms and promotes the stability of microecological networks in tea plantation soil.

Organic agriculture is highly regarded by people for its commitment to health, ecology, care, and fairness. The soil microbial community responds quickly to environmental changes and is a good indicator for evaluating soil microecology. Therefore, from the perspective of soil microbial communities, elucidating the impact of organic management on soil microecology in tea plantations has great significance for improving local tea plantation systems. The study collected bulk soil from organic management (OM) and conventional management (CM) tea plantations in Pu'er City, a major tea-producing area in China, and analyzed their species diversity, structural composition, and co-occurrence networks using metagenomics technology. Compared with CM, the diversity index (Shannon) and evenness index (Heip) of soil fungi increased by 7.38% and 84.2% in OM tea plantations, respectively. The relative abundance of microorganisms related to the nitrogen cycle increased. Specifically, there was a significant increase in Rhodobiales, a 2-fold increase in Nitrospirae, and approximately 1.95 and 2.03 times increases in unclassified genera within Betaproteobacteria and Deltaproteobacteria, respectively. The relative abundance of plant residue degradation species, Gemmatimonadetes, Ascomycota, and Basidiomycota, increased by 2.8, 1, and 1.4 times, respectively. The OM was conducive to the establishment of collaborative relationships among bacterial species and increased the diversity and complexity of species relationships in fungal communities. The network stability of soil ecosystems was promoted. The organic tea plantations' keystone taxa contained mycorrhizal fungi (Pezoloma_ericae, Rhizophagus_irregularis, Rhizophagus_clarus), as well as species involved in soil nitrogen metabolism (Acidobacteria_bacterium, Acidobacteriia_bacterium_AA117, Sphingomonas_sp._URHD0007, Enhydrobacter_aerosaccus), pathogen (Erysiphe_pulchra), and parasites (Paramycosporidium saccamoeba). The partial least squares method (PLS-SEM) indicated that OM affected N-NH negatively, increasing the abundance of fungi, and thereby positively affecting the Shannon index. In brief, reasonable organic management can improve the diversity of soil microorganisms, increase the relative abundance of beneficial bacteria in tea plantation soil, and promote the stability of the soil microbial ecological network.

Effect of Experiment Warming on Soil Fungi Community of Medicago sativa, Elymus nutans and Hordeum vulgare in Tibet.

The uncertainty response of soil fungi community to climate warming in alpine agroecosystems will limit our ability to fully exploit and utilize soil fungi resources, especially in alpine regions. In this study, a warming experiment was conducted in one perennial leguminous agroecosystem [i.e., alfalfa (Medicago sativa)], perennial gramineous agroecosystem (i.e., Elymus nutans) and annual gramineous agroecosystem [i.e., highland barley (Hordeum vulgare L)] in Tibet since 2016 to investigate the response of soil fungi community to climate warming. Soils at two layers (i.e., 0-10 cm and 10-20 cm) were collected in August 2017 to estimate soil fungi community based on the ITS method. The α-diversity, community composition and functional group abundance of soil fungi in the leguminous agroecosystem were more sensitive to climate warming. The α-diversity of soil fungi in the perennial gramineous agroecosystem were more sensitive to climate warming, but topology parameters of soil fungi species cooccurrence network in the annual gramineous agroecosystem were more sensitive to climate warming. Compared with 0-10 cm, soil fungal α-diversity, community composition and functional group abundance at 10-20 cm were more sensitive to climate warming. The topological parameters of soil fungi species cooccurrence network at 0-10 cm in the gramineous agroecosystem were more sensitive to climate warming, but those at 10-20 cm in the leguminous agroecosystem were more sensitive to climate warming. Warming increased the differences of soil fungi α-diversity and functional composition. For the Medicago sativa agroecosystem, warming increased the abundance of soil pathogenic fungi but decreased the abundance of soil symbiotic and saprophytic fungi at 10-20 cm. Therefore, responses of the soil fungi community to climate warming varied with agroecosystem types and soil depth. Climate warming can alter the differences of the soil fungi community among agroecosystems. Changes in soil fungi community caused by climate warming may be detrimental to the growth of alpine crops, at least for perennial Medicago sativa in Tibet.

Arthropod co-occurrence networks indicate environmental differences between islands and signal introduced species in Azorean native forest remnants

Island biotas are in imminent threat from anthropogenic impacts. Of these impacts, the negative effects of exotic species on the taxonomic and functional diversity of the local fauna are of major concern. Exotics may also have a detrimental effect on interspecific interactions which, in turn, can destabilize ecological networks. Species co-occurrence networks can detect species-to-species associations and are used to predict ecological interaction networks and utilized as tools to assess environmental impacts on community structure. Here, we aim to investigate whether or not topological differences of the arthropod co-occurrence networks among native forest fragments from seven Azorean islands can reveal the influence of the abiotic environment and exotic species on these networks. Co-occurrence networks were sensitive to environmental and community dissimilarities, showing a clear separation between islands and pinpointed differences between indigenous and exotic networks. Most exotics were little connected and exotic networks had a large proportion of unconnected species. The resulting decreased connectance and the increased modularity with the increase of the proportions of exotics in the networks suggest that most exotics have too low prevalence to show associations with other species, and only a few dominants drive co-occurrences. The proportion of negative links, as indicators of competition, did not increase with the increase of exotics in the habitats, suggesting that exotics provided new functional roles when they colonized native forest remnants. However, when the theoretical networks consisting of only indigenous species were investigated, connectance decreased and closeness increased with the increase of exotics, suggesting processes of network degradation. Since our study provides ample evidence for the usefulness of co-occurrence network analysis in studying island ecosystems, we recommend the use of this tool for ecosystem assessments, early warning systems and decision-making in island biodiversity conservation.

Different living environments drive deterministic microbial community assemblages in the gut of Alpine musk deer (Moschus chrysogaster).

Substantial variation in the environment directly causes remodeling of the colonized gut microbiota, controlling community diversity, and functions in the host to tune-up their adaptive states. However, the mechanisms of microbial community assembly in response to environmental changes remain unclear, especially in endangered ruminants. In this study, we analyzed the microbial communities of 37 fecal samples collected from captive and wild Alpine musk deer (Moschus chrysogaster) to characterize the complexity and assembly processes using 16S rRNA gene sequencing. We found significantly different diversities and compositions of gut microbiota among both groups associated with different living environments. Heterogeneous selection was the predominant factor regulating the gut microbiota community under similar climatic conditions, indicating that microbial community assembly was largely driven by deterministic mechanisms. The species co-occurrence network showed complex and tight connections with a higher positive correlation in the wild environment. Moreover, the captive group exhibited significant differences in chemoheterotrophy and fermentation compared with the wild group, but the opposite was observed in animal parasites or symbionts, which might be closely related to diet, energy supply, and healthcare of animals. This study provides a framework basis and new insights into understanding gut microbiota in different environments.

Structure and diversity of mycorrhizal fungi communities of different part of Bulbophyllum tianguii in three terrestrial environments.

Mycorrhizal fungi plays important roles in the seed germination and subsequent growth of orchids. The research of fungi in orchid roots, especially dominant mycorrhizal fungi is critical for orchids protection. In this study, the fungal community and composition of mycorrhizal fungi in roots, rhizomes and rhizosphere soil of Bulbophyllum tianguii grown in three terrestrial environments were analyzed by the second generation sequencing technology. The results of OTU clustering and α and β diversity analysis showed that there were significant differences in fungal communities in roots, rhizomes and rhizosphere soil of B. tianguii. The total number of OTUs in rhizomes was much less than that in roots and rhizosphere soil. The number of OTUs in rhizosphere soil and the diversity of mycorrhizal fungi were the highest. Meanwhile, the species and abundance of mycorrhizal fungi in roots and rhizomes of B. tianguii were different from those in rhizosphere soil. For different elevations, compared with B. tianguii that grow in middle of Tiankeng and top of Tiankeng, the OTUs number of B. tianguii in orchid garden is richest, and the diversity of mycorrhizal fungi in orchid garden was significantly higher than other locations. Among the three different habitats of B. tianguii, the number of OTUs in humus soil and stone habitats was notably higher than tree habitats, and the diversity of mycorrhizal fungi in humus soil was the highest. The analysis of mycorrhizal fungi in different habitats and altitudes of B. tianguii showed that Sebacina and Exophiala were the dominant mycorrhizal fungi in B. tianguii. The results of species annotation, phylogenetic tree and co-occurrence network analysis showed the dominant mycorrhizal fungi of B. tianguii mainly included Sebacina, Cladosporium, Exophiala, Fusarium. This study reveals the symbiotic relationship between Sebacina, Exophiala, Cladosporium and the B. Tianguii. It will provide a theoretical basis for the protection and biological function study of B. Tianguii.

Patterns and drivers of species co-occurrence networks in a tropical stream fish metacommunity

Patterns and drivers of species co-occurrence networks in a tropical stream fish metacommunity

Study on tanshinones regulating root-associated microbiomes of Salvia miltiorrhiza

The plant root-associated microbiomes include root microbiome and rhizosphere microbiome, which are closely related to plant life activities. Nearly 30% of photosynthesis products of plants are used to synthesize root compounds, there is evidence that root compounds regulate and significantly affect the root microbiome Tanshinones are the main hydrophobic components in Salvia miltiorrhiza. In order to study whether these compounds can regulate the root-associated microbiomes of S. miltiorrhiza, our study first identified a white root S. miltiorrhiza(BG) which contains little tanshinones. Retain of the fifth intron of tanshinones synthesis key enzyme gene SmCPS1 leading to the early termination of the SmCPS1 gene, and a stable white root phenotype. Further, wild type(WT) and BG were planted in greenhouse with nutrient soil(Pindstrup, Denmark) and Shandong soil(collected from the S. miltiorrhiza base in Weifang, Shandong), then high-throughput sequencing was used to analyze the root-associated microbiomes. The results showed that the tanshinones significantly affected the root-associated microbiomes of S. miltiorrhiza, and the impact on root microbiomes was more significant. There are significant differences between WT and BG root microbiomes in species richness, dominant strains and co-occurrence network. Tanshinones have a certain repelling effect on Bacilli which belongs to Gram-positive, while specifically attract some Gram-negative bacteria such as Betaproteobacteria and some specific genus of Alphaproteobacteria. This study determined the important role of tanshinones in regulating the structure of root-associated microbiomes from multiple angles, and shed a light for further improving the quality and yield of S. miltiorrhiza through microenvironment regulation.

Microbial Community Structure and Diversity During the Enrichment of Anaerobic Ammonium Oxidation Bacteria

To understand the changes in microbial community characteristics during the enrichment of anaerobic ammonium oxidation (ANAMMOX) bacteria, an ASBR reactor was used to culture the ANAMMOX bacteria. The composition, diversity, and species co-occurrence network of the microbial community were investigated under different cultivation times. The results showed that the ANAMMOX bacteria were enriched by gradually increasing the substrate concentration, with removal efficiencies for NH4+-N, NO2--N, and total nitrogen of 97.6%, 95.4%, and 84.9%, respectively. The high-throughput sequencing found that the dominant phyla (relative abundance>5%) were Proteobacteria, Bacteroidetes, Chloroflexi, Planctomycetes, Armatimonadetes, and Actinobacteria in the whole culture process. Candidatus Brocadia was the main ANAMMOX bacteria in the reactor, with its relative abundance increasing from 1.42% to 24.66%. During the cultivation process, the composition of the dominant microbial community did not change, while the relative abundance showed a significant difference (P<0.05). The alpha diversity of the microbial community significantly increased first and then decreased (P<0.05), and the beta diversity of the microbial community was significantly spatially differentiated (R=0.5672, P<0.01) during the culture process. Species network densities were 0.188, 0.068, 0.059, 0.18, and 0.0735 at different times during the culture process. Although the enrichment culture process resulted in weaker correlations between microorganisms, the related group of microorganisms in the phylum Aspergillus became the main node in the network. The enrichment process weakened the correlation between microorganisms; however, the microbial taxa related to the phylum Planctomycetes became the key node in the network.

Species co-occurrence networks of ground beetles in managed grasslands

Grassland biodiversity, including traditional rural biotopes maintained by traditional agricultural practices, has become threatened worldwide. Road verges have been suggested to be complementary or compensatory habitats for species inhabiting grasslands. Species co-occurrence patterns linked with species traits can be used to separate between the different mechanisms (stochasticity, environmental filtering, biotic interactions) behind community structure. Here, we study species co-occurrence networks and underlying mechanisms of ground beetle species (Carabidae) in three different managed grassland types (meadows, pastures, road verges, n = 12 in each type) in Central Finland. We aimed to find out whether road verges can be considered as compensatory to traditional rural biotopes (meadows and pastures). We found that stochasticity explained over 90% of the pairwise co-occurrences, and the non-random co-occurrences were best explained by environmental filtering, regardless of the grassland type. However, the identities and traits of the species showing non-random co-occurrences differed among the habitat types. Thus, environmental factors behind environmental filtering differ among the habitat types and are related to the site-specific characteristics and variation therein. This poses challenges to habitat management since the species’ response to management action may depend on the site-specific characteristics. Although road verges are not fully compensatory to meadows and pastures, the high similarity of species richness and the high level of shared species suggest that for carabids road verges may be corridors connecting the sparse network of the remaining traditional rural biotopes.

The human respiratory tract microbial community structures in healthy and cystic fibrosis infants

The metagenome development of the human respiratory tract was investigated by shotgun metagenome metagenomic sequencing of cough swabs from healthy children and children with cystic fibrosis (CF) between 3 weeks and 6 years of age. A healthy microbial community signature was associated with increased absolute abundances in terms of bacterial–human cell ratios of core and rare species across all age groups, with a higher diversity of rare species and a tightly interconnected species co-occurrence network, in which individual members were found in close proximity to each other and negative correlations were absent. Even without typical CF pathogens, the CF infant co-occurrence network was found to be less stable and prone to fragmentation due to fewer connections between species, a higher number of bridging species and the presence of negative species correlations. Detection of low-abundant DNA of the CF hallmark pathogen Pseudomonas aeruginosa was neither disease- nor age-associated in our cohort. Healthy and CF children come into contact with P. aeruginosa on a regular basis and from early on.

Carcinogenesis of Male Oral Submucous Fibrosis Alters Salivary Microbiomes

Most oral squamous cell carcinoma (OSCC) tumors arise from oral premalignant lesions. Oral submucous fibrosis (OSF), usually occurring in male chewers of betel quid, is a premalignant stromal disease characterized by a high malignant transformation rate and high prevalence. Although a relationship between the inhabited microbiome and carcinogenesis has been proposed, no detailed information regarding the oral microbiome of patients with OSF exists; the changes of the salivary microbiome during cancer formation remain unclear. This study compared the salivary microbiomes of male patients with OSCC and a predisposing OSF background (OSCC-OSF group) and those with OSF only (OSF group). The results of high-throughput sequencing of the bacterial 16S rRNA gene indicated that OSF-related carcinogenesis and smoking status significantly contributed to phylogenetic composition variations in the salivary microbiome, leading to considerable reductions in species richness and phylogenetic diversity. The microbiome profile of OSF-related malignancy was associated with increased microbial stochastic fluctuation, which dominated the salivary microbiome assembly and caused species co-occurrence network collapse. Artificial intelligence selection algorithms consistently identified 5 key species in the OSCC-OSF group: Porphyromonas catoniae, Prevotella multisaccharivorax, Prevotella sp. HMT-300, Mitsuokella sp. HMT-131, and Treponema sp. HMT-927. Robust accuracy in predicting oral carcinogenesis was obtained with our exploratory and validation data sets. In functional analysis, the microbiome of the OSCC-OSF group had greater potential for S-adenosyl-l-methionine and norspermidine synthesis but lower potential for l-ornithine and pyrimidine deoxyribonucleotide synthesis and formaldehyde metabolism. These findings indicated that the salivary microbiome plays important roles in modulating microbial metabolites during oral carcinogenesis. In conclusion, our results provided new insights into salivary microbiome alterations during the malignant transformation of OSF.

Disentangling the mechanisms shaping the surface ocean microbiota

BackgroundThe ocean microbiota modulates global biogeochemical cycles and changes in its configuration may have large-scale consequences. Yet, the underlying ecological mechanisms structuring it are unclear. Here, we investigate how fundamental ecological mechanisms (selection, dispersal and ecological drift) shape the smallest members of the tropical and subtropical surface-ocean microbiota: prokaryotes and minute eukaryotes (picoeukaryotes). Furthermore, we investigate the agents exerting abiotic selection on this assemblage as well as the spatial patterns emerging from the action of ecological mechanisms. To explore this, we analysed the composition of surface-ocean prokaryotic and picoeukaryotic communities using DNA-sequence data (16S- and 18S-rRNA genes) collected during the circumglobal expeditions Malaspina-2010 and TARA-Oceans.ResultsWe found that the two main components of the tropical and subtropical surface-ocean microbiota, prokaryotes and picoeukaryotes, appear to be structured by different ecological mechanisms. Picoeukaryotic communities were predominantly structured by dispersal-limitation, while prokaryotic counterparts appeared to be shaped by the combined action of dispersal-limitation, selection and drift. Temperature-driven selection appeared as a major factor, out of a few selected factors, influencing species co-occurrence networks in prokaryotes but not in picoeukaryotes, indicating that association patterns may contribute to understand ocean microbiota structure and response to selection. Other measured abiotic variables seemed to have limited selective effects on community structure in the tropical and subtropical ocean. Picoeukaryotes displayed a higher spatial differentiation between communities and a higher distance decay when compared to prokaryotes, consistent with a scenario of higher dispersal limitation in the former after considering environmental heterogeneity. Lastly, random dynamics or drift seemed to have a more important role in structuring prokaryotic communities than picoeukaryotic counterparts.ConclusionsThe differential action of ecological mechanisms seems to cause contrasting biogeography, in the tropical and subtropical ocean, among the smallest surface plankton, prokaryotes and picoeukaryotes. This suggests that the idiosyncrasy of the main constituents of the ocean microbiota should be considered in order to understand its current and future configuration, which is especially relevant in a context of global change, where the reaction of surface ocean plankton to temperature increase is still unclear.8ZJi5SwgFNAgJoQiFLy3zUVideo

The Structure and Species Co-Occurrence Networks of Soil Denitrifying Bacterial Communities Differ Between A Coniferous and A Broadleaved Forests.

Acacia mangium (AM) and Pinus massoniana (PM) are widely planted in tropical regions, whereas their effects on soil microbial communities remain unclear. We did a comprehensive investigation of soil denitrifying bacterial communities in AM and PM monoculture plantations in Southern China based on the high throughput sequencing data of their functional genes: nirK, nirS, and nosZ. The average abundance of nosZ (1.3 × 107) was significantly higher than nirS (5.6 × 106) and nirK (4.9 × 105). Shannon estimator revealed a markedly higher α-diversity of nirS and nosZ communities in PM than in AM plantations. The AM and PM plantations were dominated by different nirS and nosZ taxa belonging to proteobacteria, actinobacteria, thermoleophilia, chloroflexia, and acidobacteria, while the dominant nirK taxa were mainly categorized into proteobacteria in both types of plantations. The structure of nirS and nosZ communities shifted substantially from AM to PM plantations with changes in soil moisture, NH4+, and microbial biomass nitrogen content. The species co-occurrence network of nirK community was better organized in a more modular manner compared to nirS and nosZ communities, and the network keystone species mostly occurred in PM plantations. These results indicated a highly species corporation of nirK community in response to environmental changes, especially in PM plantations. AM and PM plantations can form different soil denitrifying microbial communities via altering soil physicochemical properties, which may further affect soil N transformations.

Does fire disturbance affect ant community structure? Insights from spatial co-occurrence networks.

The coexistence of several species involves a complex mix of positive and negative interactions that can be represented as networks. As much as other ecological features, patterns of multispecies co-occurrence are susceptible to anthropogenic disturbance. In ant communities, wildfires may enhance competitive interactions by benefiting active, aggressive species, and by increasing encounter probabilities through decreased space availability. We explored ant co-occurrence patterns by analysing the macro and microscopic structure of their interaction networks in burned and unburned habitats. We built co-occurrence networks using significant aggregations and segregations between species pairs as positive and negative interactions, respectively. We described aggregate network properties and microscopic structural changes by comparing species and interactions turnover between burned and unburned sites. We found no differences in the macroscopic structure of co-occurrence networks between different fire regimes. However, we detected changes in the composition of both species and negative interactions. Interaction turnover between networks of different habitats was mostly explained by rewiring of interactions between shared species rather than by species replacement. Our results reflected changes in ant communities in response to fire although there were no changes in global structural patterns. These changes in species and negative interactions suggest modifications in species roles translated into changes in the spatial distribution of ant species. The analysis of species co-occurrence networks is a useful tool to detect and visualize patterns in ant communities and to understand the mechanisms underlying the effects of disturbance on biodiversity.

Introduced elk alter traits of a native plant and its plant-associated arthropod community

Introduced species are known to have large impacts on native ecosystems, by extirpating species, altering species interactions and changing community composition. The mechanisms by which introduced species impact native communities often include both direct and indirect effects. Here, we combined observational field surveys, an exclosure experiment, network analysis and structural equation modeling to examine the direct and indirect effects of an introduced species on native communities. Specifically, we investigated how introduced elk indirectly impact the composition and co-occurrence patterns within a community of plant-associated arthropods by directly altering the phenotype of a native plant, Solidago velutina. Surveying the arthropods associated with the plant's inflorescence, two main patterns emerged. i) Using field observations across 500 km2 and an exclosure experiment, S. velutina growing in the presence of elk had 67–90% fewer flowering ramets, 15–85% lower percentage of flowering ramets, 33–45% fewer florets per inflorescence and 25–45% lower sexual ramet biomass. ii) Using the same exclosures, the arthropod community on S. velutina in the presence of elk, had 45% fewer species, a 70% reduction in abundance and a significant change to the species composition and co-occurrence network structure. The results from the network analysis suggested that introduced species' impacts on communities can be more than changes in richness or abundance, but include changes to species interactions. Structural equation modeling showed that elk caused a decrease in inflorescence size of S. velutina, which affected the arthropod community, suggesting that communities can change without extirpation of their host plant species. These results highlight the importance of changing intraspecific variation as a mechanism by which invasive species alter community composition and species interactions.

Inferring species roles in metacommunity structure from species co-occurrence networks.

A long-standing question in community ecology is what determines the identity of species that coexist across local communities or metacommunity assembly. To shed light upon this question, we used a network approach to analyse the drivers of species co-occurrence patterns. In particular, we focus on the potential roles of body size and trophic status as determinants of metacommunity cohesion because of their link to resource use and dispersal ability. Small-sized individuals at low-trophic levels, and with limited dispersal potential, are expected to form highly linked subgroups, whereas large-size individuals at higher trophic positions, and with good dispersal potential, will foster the spatial coupling of subgroups and the cohesion of the whole metacommunity. By using modularity analysis, we identified six modules of species with similar responses to ecological conditions and high co-occurrence across local communities. Most species either co-occur with species from a single module or are connectors of the whole network. Among the latter are carnivorous species of intermediate body size, which by virtue of their high incidence provide connectivity to otherwise isolated communities playing the role of spatial couplers. Our study also demonstrates that the incorporation of network tools to the analysis of metacommunity ecology can help unveil the mechanisms underlying patterns and processes in metacommunity assembly.