A waterfowl seed-dispersal network from the Neotropical region is nested and modular

Seed dispersal by vertebrates is fundamental for the persistence of plant species, forming networks of interactions that are often nested and modular. Networks involving angiosperms and frugivorous birds are relatively well-studied in the Neotropical region, but there are no previous studies of networks involving waterbirds. Here, we describe the structure of a Neotropical waterfowl seed-dispersal network and identify the species that have an important role for the network structure. We used information on 40 plant taxa found in fecal samples of five common waterfowl species to calculate the nestedness (NODF), weighted nestedness (WNODF), modularity, and weighted modularity of the network. We found that the network was nested, with yellow-billed teal showing the highest contribution both to nestedness and weighted nestedness. Twenty-four plant species contributed positively to weighted nestedness, with Salzmann's mille graines presenting the highest influence both to nestedness and weighted nestedness. The network was modular, but the weighted modularity was not significant. These results need to be considered with caution due to incomplete interaction sampling for two species. Ringed teal, Brazilian teal, and yellow-billed teal were considered hub modular species. Among plants, beak sedges and water snowflake were considered modular hub species, while Salzmann's mille graines and spikerush were network connectors. The structure of this Neotropical waterbird seed-dispersal network differed from the only previous waterfowl network study, from Europe, which found similar level of nestedness but no significant modularity. We include several possible explanations for this discrepancy and identified priorities for future research into waterbird–plant interaction networks. Abstract in Portuguese is available with online material.     

Interaction network of vascular epiphytes and trees in a subtropical forest

The commensalistic interaction between vascular epiphytes and host trees is a type of biotic interaction that has been recently analysed with a network approach. This approach is useful to describe the network structure with metrics such as nestedness, specialization and interaction evenness, which can be compared with other vascular epiphyte-host tree networks from different forests of the world. However, in several cases these comparisons showed different and inconsistent patterns between these networks, and their possible ecological and evolutionary determinants have been scarcely studied. In this study, the interactions between vascular epiphytes and host trees of a subtropical forest of sierra de San Javier (Tucuman, Argentina) were analysed with a network approach. We calculated metrics to characterize the network and we analysed factors such as the abundance of species, tree size, tree bark texture, and tree wood density in order to predict interaction frequencies and network structure. The interaction network analysed exhibited a nested structure, an even distribution of interactions, and low specialization, properties shared with other obligated vascular epiphyte-host tree networks with a different assemblage structure. Interaction frequencies were predicted by the abundance of species, tree size and tree bark texture. Species abundance and tree size also predicted nestedness. Abundance indicated that abundant species interact more frequently; and tree size was an important predictor, since larger-diameter trees hosted more vascular epiphyte species than small-diameter trees. This is one of the first studies analyzing interactions between vascular epiphytes and host trees using a network approach in a subtropical forest, and taking the whole vascular epiphyte assemblage of the sampled community into account.     

Interaction type and intimacy structure networks between forest-dwelling organisms and their host trees

Species interact in many ways. Potentially, the type of interaction, e.g. mutualistic, commensalistic or antagonistic, determines the structure of interaction networks, but this remains poorly tested. Here we investigate whether epiphytes and wood decomposers, having different types of interaction with their host trees, show different network properties. We also test whether the traits of host trees affect network architecture. We recorded presence/absence of organisms colonizing trees, and traits of host trees, in 102 forest plots. Epiphytic bryophytes (64 species) and lichens (119 species) were recorded on c. 2300 trees. Similarly, wood-inhabiting fungi (193 species) were recorded on c. 900 dead wood items. We studied the patterns of species aggregation on host trees by comparing network metrics of species specialization, nestedness and modularity. Next, we tested whether the prevalence of interactions was influenced by host tree traits. We found non-random interaction patterns between host trees and the three ecological groups (bryophytes, lichens and fungi), with nested and modular structures associated with high host specificity. A higher modularity and number of modules was found for fungi than for epiphytes, which is likely related to their trophic relationship with the host plant, whilst the stronger nestedness for epiphytes is likely reflecting the commensalistic nature of their interactions. For all three groups, the difference in prevalence of interaction across modules was determined by a gradient in interaction intimacy (i.e. host tree specialization), driven by host tree traits. We conclude that the type of interaction with host trees defines the properties of each network: while autotrophic epiphyte networks show similar properties to mutualistic networks, the heterotrophic wood decomposers show similarity with antagonistic networks.     

Patterns of interactions of a large fish-parasite network in a tropical floodplain

1.?Describing and explaining the structure of species interaction networks is of paramount importance for community ecology. Yet much has to be learned about the mechanisms responsible for major patterns, such as nestedness and modularity in different kinds of systems, of which large and diverse networks are a still underrepresented and scarcely studied fraction. 2.?We assembled information on fishes and their parasites living in a large floodplain of key ecological importance for freshwater ecosystems in the Paraná River basin in South America. The resulting fish-parasite network containing 72 and 324 species of fishes and parasites, respectively, was analysed to investigate the patterns of nestedness and modularity as related to fish and parasite features. 3.?Nestedness was found in the entire network and among endoparasites, multiple-host life cycle parasites and native hosts, but not in networks of ectoparasites, single-host life cycle parasites and non-native fishes. All networks were significantly modular. Taxonomy was the major host's attribute influencing both nestedness and modularity: more closely related host species tended to be associated with more nested parasite compositions and had greater chance of belonging to the same network module. Nevertheless, host abundance had a positive relationship with nestedness when only native host species pairs of the same network module were considered for analysis. 4.?These results highlight the importance of evolutionary history of hosts in linking patterns of nestedness and formation of modules in the network. They also show that functional attributes of parasites (i.e. parasitism mode and life cycle) and origin of host populations (i.e. natives versus non-natives) are crucial to define the relative contribution of these two network properties and their dependence on other ecological factors (e.g. host abundance), with potential implications for community dynamics and stability.     

The network structure of plant-arbuscular mycorrhizal fungi

Ecological network theory predicts that in mutualistic systems specialists tend to interact with a subset of species with which generalists interact (i.e. nestedness). Approaching plant-arbuscular mycorrhizal fungi (AMF) association using network analyses will allow the generality of this pattern to be expanded to the ubiquitous plant-AMF mutualism. Based on certain plant-AMF specificity recently suggested, networks are expected to be nested as a result of their mutualistic nature, and modular, with certain species interacting more tightly than others. Network analyses were used to test for nestedness and modularity and to compare the different contribution of plant and AMF to the overall nestedness. Plant-AMF networks share general network properties with other mutualisms. Plant species with few AMFs in their roots tend to associate with those AMFs recorded in most plant species. AMFs present in a few plant species occur in plant species sheltering most AMF (i.e. nestedness). This plant-AMF network presents weakly interlinked subsets of species, strongly connected internally (i.e. modularity). Both plants and AMF show a nested structure, although AMFs have lower nestedness than plants. The plant-AMF interaction pattern is interpreted in the context of how plant-AMF associations can be underlying mechanisms shaping plant community assemblages.     

Temporal variation in bat-fruit interactions: Foraging strategies influence network structure over time

Mutualistic interactions, such as seed dispersal, are important for the maintenance of structure and stability of tropical communities. However, there is a lack of information about spatial and temporal variation in plant-animal interaction networks. Thus, our goal was to assess the effect of bat's foraging strategies on temporal variation in the structure and robustness of bat-fruit networks in both a dry and a rain tropical forest. We evaluated monthly variation in bat-fruit networks by using seven structure metrics: network size, average path length, nestedness, modularity, complementary specialization, normalized degree and betweenness centrality. Seed dispersal networks showed variations in size, species composition and modularity; did not present nested structures and their complementary specialization was high compared to other studies. Both networks presented short path lengths, and a constantly high robustness, despite their monthly variations. Sedentary bat species were recorded during all the study periods and occupied more central positions than nomadic species. We conclude that foraging strategies are important structuring factors that affect the dynamic of networks by determining the functional roles of frugivorous bats over time; thus sedentary bats are more important than nomadic species for the maintenance of the network structure, and their conservation is a must.     

Population sinks resulting from degraded habitats of an obligate life-history pathway

In networks of plant–animal mutualisms, different animal groups interact preferentially with different plants, thus forming distinct modules responsible for different parts of the service. However, what we currently know about seed dispersal networks is based only on birds. Therefore, we wished to fill this gap by studying bat–fruit networks and testing how they differ from bird–fruit networks. As dietary overlap of Neotropical bats and birds is low, they should form distinct mutualistic modules within local networks. Furthermore, since frugivory evolved only once among Neotropical bats, but several times independently among Neotropical birds, greater dietary overlap is expected among bats, and thus connectance and nestedness should be higher in bat–fruit networks. If bat–fruit networks have higher nestedness and connectance, they should be more robust to extinctions. We analyzed 1 mixed network of both bats and birds and 20 networks that consisted exclusively of either bats (11) or birds (9). As expected, the structure of the mixed network was both modular (M = 0.45) and nested (NODF = 0.31); one module contained only birds and two only bats. In 20 datasets with only one disperser group, bat–fruit networks (NODF = 0.53 ± 0.09, C = 0.30 ± 0.11) were more nested and had a higher connectance than bird–fruit networks (NODF = 0.42 ± 0.07, C = 0.22 ± 0.09). Unexpectedly, robustness to extinction of animal species was higher in bird–fruit networks (R = 0.60 ± 0.13) than in bat–fruit networks (R = 0.54 ± 0.09), and differences were explained mainly by species richness. These findings suggest that a modular structure also occurs in seed dispersal networks, similar to pollination networks. The higher nestedness and connectance observed in bat–fruit networks compared with bird–fruit networks may be explained by the monophyletic evolution of frugivory in Neotropical bats, among which the diets of specialists seem to have evolved from the pool of fruits consumed by generalists.     

Availability of food resources and habitat structure shape the individual‐resource network of a Neotropical marsupial

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Agricultural intensification drives changes in hybrid network robustness by modifying network structure

Within ecological communities, species engage in myriad interaction types, yet empirical examples of hybrid species interaction networks composed of multiple types of interactions are still scarce. A key knowledge gap is understanding how the structure and stability of such hybrid networks are affected by anthropogenic disturbance. Using 15,169 interaction observations, we constructed 16 hybrid herbivore‐plant‐pollinator networks along an agricultural intensification gradient to explore changes in network structure and robustness to local extinctions. We found that agricultural intensification led to declines in modularity but increases in nestedness and connectance. Notably, network connectance, a structural feature typically thought to increase robustness, caused declines in hybrid network robustness, but the directionality of changes in robustness along the gradient depended on the order of local species extinctions. Our results not only demonstrate the impacts of anthropogenic disturbance on hybrid network structure, but they also provide unexpected insights into the structure‐stability relationship of hybrid networks.     

Pollination networks of oil-flowers: a tiny world within the smallest of all worlds

1.   In the Neotropics, most plants depend on animals for pollination. Solitary bees are the most important vectors, and among them members of the tribe Centridini depend on oil from flowers (mainly Malpighiaceae) to feed their larvae. This specialized relationship within 'the smallest of all worlds' (a whole pollination network) could result in a 'tiny world' different from the whole system. This 'tiny world' would have higher nestedness, shorter path lengths, lower modularity and higher resilience if compared with the whole pollination network.
2.   In the present study, we contrasted a network of oil-flowers and their visitors from a Brazilian steppe ('caatinga') to whole pollination networks from all over the world.
3.   A network approach was used to measure network structure and, finally, to test fragility. The oil-flower network studied was more nested ( NODF  = 0·84, N  =   0·96) than all of the whole pollination networks studied. Average path lengths in the two-mode network were shorter (one node, both for bee and plant one-mode network projections) and modularity was lower ( M  =   0·22 and four modules) than in all of the whole pollination networks. Extinctions had no or small effects on the network structure, with an average change in nestedness smaller than 2% in most of the cases studied; and only two species caused coextinctions. The higher the degree of the removed species, the stronger the effect and the higher the probability of a decrease in nestedness.
4.   We conclude that the oil-flower subweb is more cohesive and resilient than whole pollination networks. Therefore, the Malpighiaceae have a robust pollination service in the Neotropics. Our findings reinforce the hypothesis that each ecological service is in fact a mosaic of different subservices with a hierarchical structure ('webs within webs').     

Flow intermittency decreases nestedness and specialisation of diatom communities in Mediterranean rivers

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Influence of taxonomic resolution on mutualistic network properties

1. Ecologists are increasingly interested in plant–pollinator networks that synthesize in a single object the species and the interactions linking them within their ecological context. Numerous indices have been developed to describe the structural properties and resilience of these networks, but currently, these indices are calculated for a network resolved to the species level, thus preventing the full exploitation of numerous datasets with a lower taxonomic resolution. Here, we used datasets from the literature to study whether taxonomic resolution has an impact on the properties of plant–pollinator networks.
2. For a set of 41 plant–pollinator networks from the literature, we calculated nine network index values at three different taxonomic resolutions: species, genus, and family. We used nine common indices assessing the structural properties or resilience of networks: nestedness (estimated using the nestedness index based on overlap and decreasing fill [NODF], weighted NODF, discrepancy [BR], and spectral radius [SR]), connectance, modularity, robustness to species loss, motifs frequencies, and normalized degree.
3. We observed that modifying the taxonomic resolution of these networks significantly changes the absolute values of the indices that describe their properties, except for the spectral radius and robustness. After the standardization of indices measuring nestedness with the Z‐score, three indices—NODF, BR, and SR for binary matrices—are not significantly different at different taxonomic resolutions. Finally, the relative values of all indices are strongly conserved at different taxonomic resolutions.
4. We conclude that it is possible to meaningfully estimate the properties of plant–pollinator interaction networks with a taxonomic resolution lower than the species level. We would advise using either the SR or robustness on untransformed data, or the NODF, discrepancy, or SR (for weighted networks only) on Z‐scores. Additionally, connectance and modularity can be compared between low taxonomic resolution networks using the rank instead of the absolute values.

     

Network analyses reveal the role of large snakes in connecting feeding guilds in a species‐rich Amazonian snake community

In ecological communities, interactions between consumers and resources lead to the emergence of ecological networks and a fundamental problem to solve is to understand which factors shape network structure. Empirical and theoretical studies on ecological networks suggest predator body size is a key factor structuring patterns of interaction. Because larger predators consume a wider resource range, including the prey consumed by smaller predators, we hypothesized that variation in body size favors the rise of nestedness. In contrast, if resource consumption requires specific adaptations, predators are expected to consume distinct sets of resources, thus favoring modularity. We investigate these predictions by characterizing the trophic network of a species‐rich Amazonian snake community (62 species). Our results revealed an intricate network pattern resulting from larger species feeding on higher diversity of prey and therefore promoting nestedness, whereas snakes with specific lifestyles and feeding on distinct resources, promoting modularity. Species removal simulations indicated that the nested structure is favored mainly by the presence of five species of the family Boidae, which because of their body size and generalist lifestyles connect modules in the network. Our study highlights the particular ways traits affect the structure of interactions among consumers and resources at the community level.     

Abiotic factors shape temporal variation in the structure of an ant–plant network

Despite recognition of key biotic processes in shaping the structure of biological communities, few empirical studies have explored the influences of abiotic factors on the structural properties of mutualistic networks. We tested whether temperature and precipitation contribute to temporal variation in the nestedness of mutualistic ant–plant networks. While maintaining their nested structure, nestedness increased with mean monthly precipitation and, particularly, with monthly temperature. Moreover, some species changed their role in network structure, shifting from peripheral to core species within the nested network. We could summarize that abiotic factors affect plant species in the vegetation (e.g., phenology), meaning presence/absence of food sources, consequently an increase/decrease of associations with ants, and finally, these variations to fluctuations in nestedness. While biotic factors are certainly important, greater attention needs to be given to abiotic factors as underlying determinants of the structures of ecological networks.     

Nestedness of Ectoparasite-Vertebrate Host Networks

Determining the structure of ectoparasite-host networks will enable disease ecologists to better understand and predict the spread of vector-borne diseases. If these networks have consistent properties, then studying the structure of well-understood networks could lead to extrapolation of these properties to others, including those that support emerging pathogens. Borrowing a quantitative measure of network structure from studies of mutualistic relationships between plants and their pollinators, we analyzed 29 ectoparasite-vertebrate host networks—including three derived from molecular bloodmeal analysis of mosquito feeding patterns—using measures of nestedness to identify non-random interactions among species. We found significant nestedness in ectoparasite-vertebrate host lists for habitats ranging from tropical rainforests to polar environments. These networks showed non-random patterns of nesting, and did not differ significantly from published estimates of nestedness from mutualistic networks. Mutualistic and antagonistic networks appear to be organized similarly, with generalized ectoparasites interacting with hosts that attract many ectoparasites and more specialized ectoparasites usually interacting with these same “generalized” hosts. This finding has implications for understanding the network dynamics of vector-born pathogens. We suggest that nestedness (rather than random ectoparasite-host associations) can allow rapid transfer of pathogens throughout a network, and expand upon such concepts as the dilution effect, bridge vectors, and host switching in the context of nested ectoparasite-vertebrate host networks.     

A comparative analysis of nested subset patterns of species composition

We present a broad comparative assessment of nested subsets in species composition among ecological communities. We assembled presence-absence data from a broad range of taxa, geographic regions, and spatial scales; and subjected this collection of datasets to common analyses, including a variety of metrics for measuring nestedness and null hypotheses against which to evaluate them. Here we identify ecological patterns in the prevalence and strength of nested subset structure, and assess differences and biases among the available methodologies. In all, we compiled 279 presence-absence matrices, of which 163 do not overlap in their coverage of species and sites. The survey includes studies on vertebrates, arthropods, mollusks, plants, and other taxa; from north temperate, tropical, and south temperate latitudes. Our results were as follows. Statistically significant nestedness was common. Assemblages from landbridge archipelagos were strongly nested, and immigration experiments were least nested. This adds further empirical support to the hypothesis that extinction plays a major role in producing nested structure. Nestedness was positively correlated with the ratio of the areas of the largest and smallest sites, suggesting that the range in area of sites affects nestedness. Taxonomic differences in nestedness were weak. Higher taxonomic levels showed stronger nesting than their constituent lower taxa. We observed no effect of distance of isolation on nestedness; nor any effects of latitude. With regard to methodology, the metrics Nc and Ut yielded similar results, although Nc proved slightly more flexible in use, and deals differently with tied sites. Similarities also exist in the behavior of N0 (“N”) and Up, and between N1 and Ua. Standardized nestedness metrics were mostly insensitive to matrix size, and were useful in comparative analyses among presence-absence matrices. Most metrics were affected by the proportion of presences in the matrix. All analyses of nestedness, therefore, should test for bias due to matrix fill. We suggest that the factors controlling nested subset structure can be thought of as four filters that species pass to occur at a site: a sampling filter, a distance filter, a habitat filter, and an area filter – and three constraints on community homogeneity: evolutionary history, recent history, and spatial variation in the environment. The scale of examination can also have important effects on the degree of nestedness observed. Received: 13 September 1996 / Accepted: 16 September 1997     

Architecture of an antagonistic tree/fungus network: the asymmetric influence of past evolutionary history

Background

Compartmentalization and nestedness are common patterns in ecological networks. The aim of this study was to elucidate some of the processes shaping these patterns in a well resolved network of host/pathogen interactions.

Methology/Principal Findings

Based on a long-term (1972–2005) survey of forest health at the regional scale (all French forests; 15 million ha), we uncovered an almost fully connected network of 51 tree taxa and 157 parasitic fungal species. Our analyses revealed that the compartmentalization of the network maps out the ancient evolutionary history of seed plants, but not the ancient evolutionary history of fungal species. The very early divergence of the major fungal phyla may account for this asymmetric influence of past evolutionary history. Unlike compartmentalization, nestedness did not reflect any consistent phylogenetic signal. Instead, it seemed to reflect the ecological features of the current species, such as the relative abundance of tree species and the life-history strategies of fungal pathogens. We discussed how the evolution of host range in fungal species may account for the observed nested patterns.

Conclusion/Significance

Overall, our analyses emphasized how the current complexity of ecological networks results from the diversification of the species and their interactions over evolutionary times. They confirmed that the current architecture of ecological networks is not only dependant on recent ecological processes.     

Comparing the plant–herbivore network topology of different insect guilds in Neotropical savannas

1. Herbivorous insects can be classified into several trophic guilds with different levels of specialisation on their host plants, which may influence the topological structure of their trophic networks. The present study tested the hypothesis that the structure of plant–herbivore networks differs between guilds of galling, sucking, and chewing insects. 2. Six areas of Neotropical savannas were studied in two localities in the North of the state of Minas Gerais, Brazil. In each area, interactions between plant and insect species were used to build networks for different guilds. 3. In total, 18 plant–herbivore networks were built, comprising 317 insect morphospecies, 50 plant species, and 489 distinct interactions. The networks were characterised using species richness and different network topological measures (connectance, modularity, nestedness, and specialisation). 4. The results obtained showed no difference in species richness, network size, and connectance between distinct insect herbivore guilds. However, it was found that modularity was higher for exophagous than galling insect networks and nestedness was higher for chewers than for other guilds. On the other hand, galling insect networks showed higher specialisation than exophagous insect networks, and sucking insect networks were more specialised than chewing insect networks. 5. The findings of the present study indicate that, although species richness did not differ between insect guilds of herbivores in Neotropical savannas, the topological structure of networks is sensitive to biological and ecological differences between these herbivore groups. The present study stands out as the first to systematically compare the network structure of different herbivore guilds in Neotropical savannas.     

A network analysis of plant–pollinator interactions in temperate rain forests of Chiloé Island, Chile

This study characterizes the structure of a plant–pollinator network in a temperate rain forest of Chiloé Island, southern Chile, where woody species are strongly dependent on biotic pollinators, and analyzes its robustness to the loss of participating species. Degree distribution, nestedness, and expected species persistence were evaluated. In addition, we assessed the roles of predefined subsets of plants (classified by life forms) and pollinators (grouped by taxonomic orders) in the network’s structure and dynamics. For this, we simulated the complete removal of each plant and pollinator subset and analyzed the resultant connectivity patterns, as well as the expected long-term species losses by running a stochastic model. Finally, we evaluated the sensitivity of the network structure to the loss of single species in order to identify potential targets for conservation. Our results show that the plant–pollinator network of this Chilean temperate rain forest exhibits a nested structure of interactions, with a degree distribution best described by a power law model. Model simulations revealed the importance of trees and hymenopterans as pivotal groups that maintain the core structure of the pollination network and guarantee overall species persistence. The hymenopterans Bombus dahlbomii and Diphaglossa gayi, the shrubs Tepualia stipularis and Ugni molinae, the vines Mitraria coccinea and Asteranthera ovata, and the entire set of tree species exerted a disproportionately large influence on the preservation of network structure and should be considered as focal species for conservation programs given current threats from selective logging and habitat loss.