Fruit traits and temporal abundance shape plant-frugivore interaction networks in a seasonal tropical forest.

Interactions between fleshy fruited plants and frugivores are crucial for the structuring and functioning of biotic communities, particularly in tropical forests where both groups are diverse and play different roles in network organization. However, it remains poorly understood how different groups of frugivore species and fruit traits contribute to network structure. We recorded interactions among 28 plant species and three groups of frugivores (birds, bats, and non-flying mammals) in a seasonal forest in Mexico to determine which species contribute more to network structure and evaluate the importance of each species. We also determined whether fruit abundance, water content, morphology traits, and fruiting phenology are related to network parameters: the number of interactions, species contribution to nestedness, and species strength. We found that plants did not depend on a single group of frugivores, but rather on one species of each group: the bird Pitangus sulphuratus, the bat Sturnira parvidens, and the non-flying mammal Procyon lotor. The abundance, size, and water content of the fruits were significantly related to the contribution to nestedness, number of interactions, and species strength index of plant species. Tree species and birds contributed mainly to the nested structure of the network. We show that the structure of plant-frugivore networks in this seasonal forest is non-random and that fruit traits (i.e., abundance, phenology, size, and water content) are important factors shaping plant-frugivore networks. Identification of the key species and their traits that maintain the complex structure of species interactions is therefore fundamental for the integral conservation of tropical forests.

Influence of plant resistance traits in selectiveness and species strength in a tropical plant-herbivore network.

PREMISE OF THE STUDY: Plant-herbivore networks are highly specialized in their interactions, yet they are highly variable with regard to the relative importance of specific host species for herbivores. How host species traits determine specialization and species strength in this antagonistic network is still an unanswered question that we addressed in this study. METHODS: We assessed plant cover and antiherbivore resistance traits to assess the extent to which they accounted for the variation in specialization and strength of interactions among species in a plant-herbivore network. We studied a tropical antagonistic network including a diverse herbivore-host plant assemblages in different habitat types and climatic seasons, including host plants with different life histories. KEY RESULTS: Particular combinations of leaf toughness, trichome density, and phenolic compounds influenced herbivore specialization and host species strength, but with a significant spatiotemporal variation among plant life histories. Conversely, plant-herbivore network parameters were not influenced by plant cover. CONCLUSIONS: Our study highlights the importance of species-specific resistance traits of plants to understand the ecological and evolutionary consequences of plant-herbivore interaction networks. The novelty of our research lies in the use of a trait-based approach to understand the variation observed in diverse plant-herbivore networks.

Vascular epiphytes and host trees of ant-gardens in an anthropic landscape in southeastern Mexico.

Ant-gardens (AGs) are considered one of the most complex mutualist systems between ants and plants, since interactions involving dispersal, protection, and nutrition occur simultaneously in them; however, little is known about the effects of the transformation of ecosystems on their diversity and interactions. In five environments with different land use within an anthropic landscape in southeastern Mexico, we investigated the diversity and composition of epiphytes and host trees of AGs built by Azteca gnava. A total of 10,871 individuals of 26 epiphytic species, associating with 859 AGs located in 161 host trees, were recorded. The diversity and composition of epiphytes tended to be different between environments; however, Aechmea tillandsioides and Codonanthe uleana were the most important species and considered true AG epiphytes, because they were the most frequent, abundant, and occurred exclusively in AGs. Other important species were the orchids Epidendrum flexuosum, Coryanthes picturata, and Epidendrum pachyrachis, and should also be considered true AG epiphytes, because they occurred almost exclusively in the AGs. The AG abundance in agroforestry plantations was similar or even greater than in riparian vegetation (natural habitat). The AGs were registered in 37 host species but were more frequent in Mangifera indica and Citrus sinensis. We conclude that true epiphytes of A. gnava AGs persist in different environments and host trees, and even these AGs could proliferate in agroforestry plantations of anthropic landscapes.

Nectar secretion on fern fronds associated with lower levels of herbivore damage: field experiments with a widespread epiphyte of Mexican cloud forest remnants.

BACKGROUND AND AIMS: The oldest group of plants in which nectar secretions have been observed are the Polypodiopsida (ferns sensu lato). Nectaries have been reported in a dozen extant genera. The function of these nectaries has been investigated in several fern species, and in some circumstances has been demonstrated to have an antiherbivore role, attracting and maintaining biotic defence (ants and/or other predatory arthropods). This study documents foliar nectaries in Pleopeltis crassinervata, a widespread Central American epiphyte growing on a variety of trees in cloud forest areas of Veracruz, Mexico. This is a new record for this genus and species. METHODS: As previous experimental work on epiphytic species of Polypodium has demonstrated a protective role of ants for developing fronds, we conducted similar experiments (using nylon nail polish to cover nectaries rather than excluding ants with bands of sticky resin as in earlier work). The fronds of Pl. crassinervata developed over 6 weeks, at which time damage was assessed. The experiment was simultaneously conducted on a sympatric species lacking nectaries, Polypodium furfuraceum. Herbivore placement experiments were conducted with large and small caterpillars on both of these ferns. KEY RESULTS: Fronds with nectaries covered suffered greater damage from herbivores over the course of their development, compared with fronds that had uncovered nectaries functioning normally. The parallel experiment on Po. furfuraceum showed no difference between manipulated and control fronds. Six species of ants (Brachymyrmex minutus, Crematogaster formosa, Paratrechina longicornis, Solenopsis geminata, S. picea and Wasmannia auropunctata) were observed visiting nectaries of Pl. crassinervata; most were effective in removing herbivore larvae placed on the fronds. CONCLUSIONS: The long evolutionary history of ferns may explain why some previous studies of fern nectaries have shown little or no benefit to ferns from nectary visitors, as any coevolved herbivores are those resistant to ant defence. The results suggest that ants protect Pl. crassinervata fronds against herbivory. The presence of nectaries, and the relationship with ants, may contribute to this fern's widespread occurrence and persistence in the face of disturbance, though many other factors also play a role. Ant defence may be more likely to benefit a widespread species of disturbed habitats that encounters a wide range of non-adapted herbivores.

Long-term temporal variation in the organization of an ant-plant network.

BACKGROUND AND AIMS: Functional groups of species interact and coevolve in space and time, forming complex networks of interacting species. A long-term study of temporal variation of an ant-plant network is presented with the aims of: (1) depicting its structural changes over a 20-year period; (2) detailing temporal variation in network topology, as revealed by nestedness and modularity analysis and other parameters (i.e. connectance, niche overlap); and (3) identifying long-term turnover in taxonomic structure (i.e. switches in ant resource use or plant visitor assemblages according to taxa). METHODS: Fieldwork was carried out at La Mancha, Mexico, and ant-plant interactions were observed between 1989 and 1991, between 1998 and 2000, and between May 2010 and 2011. Occurrences of ants on extrafloral nectaries (EFNs) were recorded. The resulting ant-plant networks were constructed from qualitative presence-absence data determined by a species-species matrix defined by the frequency of occurrence of each pairwise ant-plant interaction. KEY RESULTS: Network variation across time was stable and a persistent nested structure may have contributed to the maintenance of resilient and species-rich communities. Modularity was lower than expected, especially in the most recent networks, indicating that the community exhibited high overlap among interacting species (e.g. few species were hubs in the more recent network, being partly responsible for the nested pattern). Structurally, the connections created among modules by super-generalists gave cohesion to subsets of species that otherwise would remain unconnected. This may have allowed an increasing cascade-effect of evolutionary events among modules. Mutualistic ant-plant interactions were structured 20 years ago mainly by the subdominant nectarivorous ant species Camponotus planatus and Crematogaster brevispinosa, which monopolized the best extrafloral nectar resources and out-competed other species with broader feeding habits. Through time, these ants, which are still present, lost their position as network hubs and diminished in their importance in structuring the network; simultaneously, plants gained in importance. CONCLUSIONS: The long-term network analysis reveals a decrease in attended plant species richness, a notable increase in plant species participation from 1990 to 2010 (sustained by less plant taxonomic similarity in the older 1990 network), an increase in the number of ant species and a diminishing dominance of super-generalist ants. The structure of the community has remained highly nested and connected with low modularity, suggesting overall a more participative, homogeneous, cohesive interaction network. Although previous studies have suggested that interactions between ants and EFN-bearing plants are susceptible to seasonality, abiotic factors and perturbation, this cohesive structure appears to be the key for biodiversity and community maintenance.

Interaction Networks Help to Infer the Vulnerability of the Saproxylic Beetle Communities That Inhabit Tree Hollows in Mediterranean Forests.

Insect communities are facing contrasting responses due to global change. However, knowledge on impacts of communities' reorganizations is scarce. Network approaches could help to envision community changes in different environmental scenarios. Saproxylic beetles were selected to examine long-term variations in insect interaction/diversity patterns and their vulnerability to global change. We evaluated interannual differences in network patterns in the tree hollow-saproxylic beetle interaction using absolute samplings over an 11-year interval in three Mediterranean woodland types. We explored saproxylic communities' vulnerability to microhabitat loss via simulated extinctions and by recreating threat scenarios based on decreasing microhabitat suitability. Although temporal diversity patterns varied between woodland types, network descriptors showed an interaction decline. The temporal beta-diversity of interactions depended more on interaction than on species turnover. Interaction and diversity temporal shifts promoted less specialized and more vulnerable networks, which is particularly worrisome in the riparian woodland. Network procedures evidenced that saproxylic communities are more vulnerable today than 11 years ago irrespective of whether species richness increased or decreased, and the situation could worsen in the future depending on tree hollow suitability. Network approaches were useful for predicting saproxylic communities' vulnerability across temporal scenarios and, thus, for providing valuable information for management and conservation programs.

Changes in the core species of the ant-plant network of oak forest converted to grassland: replacement of its ant functional groups.

Land-use change in terrestrial environments is one of the main threats to biodiversity. The study of ant-plant networks has increased our knowledge of the diversity of interactions and structure of these communities; however, little is known about how land-use change affects ant-plant networks. Here we determine whether the change in land use, from native oak forest to induced grassland, affected the network properties of ant-plant networks in a temperate forest in Mexico. We hypothesize that the disturbed vegetation will be more nested and generalized due to the addition of generalist species to the network. The oak forest network comprises 47 plant species and 11 ant species, while the induced grassland network has 35 and 13, respectively. Floral nectar was the resource used most intensely by the ants in both vegetation types. The ant-plant network of the induced grassland was significantly more nested and generalist than that of the oak forest; however, none of the networks were nested when considering the frequency of interaction. In both vegetation types, the ants were more specialized than the plants, and niche overlap was low. This could be related to the dominant species present in each type of vegetation: Prenolepis imparis in the oak forest and Camponotus rubrithorax in the grassland. The central core of cold climate ant species in the oak forest was replaced by a central core of subordinate Camponotini and tropical specialists in the induced grassland. These results suggest that the increase in nestedness and generalization in the grassland may be related to the loss of the cold climate specialists from the core of the oak forest network. Our findings provide evidence that land-use change increases the level of generalization in the ant-plant interaction networks of temperate forests.

Short-Term Temporal Patterns in Herbivore Beetle Assemblages in Polyculture Neotropical Forest Plantations.

Although insect herbivorous communities in tropical forests are known to exhibit strong seasonality, few studies have systematically assessed temporal patterns of variation in community structure and plant-herbivore interactions in early successional arboreal communities. We assessed seasonal and interannual variation of the diversity and composition of herbivorous beetles and the tree-herbivore network in a recently established polyculture forest plantation, during the dry and the rainy seasons of 2012 and of 2013. Species richness was similar between years, while the ecological diversity was higher in 2012. Comparing seasons, no differences were found in 2012, whereas in 2013, the species richness and ecological diversity were higher during the dry season. The species composition differed radically across years and seasons. Moreover, a quantitative nested pattern was consistently found across both temporal scales, more influenced by species densities. We found temporal changes in the species strength, whereas connectance and interaction evenness remained stable. Rapid temporal changes in the structural complexity of recently established polyculture plantations and the availability and quality of the trophic resources they offer may act as drivers of beetle diversity patterns, promoting rapid variation in herbivore composition and some interacting attributes. Nonetheless, network structure, connectance, and interaction evenness remained similar, suggesting that reorganizations in the distribution of species may determine the maintenance of the patterns of interaction. Further work assessing long-term temporal dynamics of herbivore beetle assemblages are needed to more robustly relate diversity and interaction patterns to biotic and abiotic factors and their implications in management programs.

Lowering the density: ants associated with the myrmecophyte Tillandsia caput-medusae diminish the establishment of epiphytes.

Ants benefit myrmecophytic plants by two main activities defending them from herbivores and offering nutrients. Ants' territorial defence behaviour also benefits their myrmecophytic plants; in the case of trees, this behaviour includes eliminating structural parasites (epiphytes and lianas). These benefits could also occur with myrmecophytic epiphytes by decreasing the abundance of competing epiphytes. In two subunits of a tropical dry forest in the centre of Mexico, we (i) recorded the diversity of ants associated with the myrmecophyte Tillandsia caput-medusae, and experimentally tested: (ii) the effect of the ants associated with the myrmecophyte in the removal of its seeds and the seeds of other sympatric non-myrmecophyte species of Tillandsia; and (iii) if seed remotion by ants corresponds with epiphyte load in the preferred (Bursera copallifera) and limiting phorophyte species (B. fagaroides, Ipomoea pauciflora and Sapium macrocarpum). In five trees per species, we tied seed batches of T. caput-medusae, T. hubertiana, T. schiedeana and T. recurvata. One seed batch was close, and the other far away from a T. caput-medusae with active ants. Between forest subunits, ant richness was similar, but diversity and evenness differed. Ants diminish seed establishment of all the Tillandsia species; this effect is stronger in the forest subunit with a large ant diversity, maybe because of ant competition. Seed remotion by ants is independent of phorophyte species identity. Although ants can provide benefits to T. caput-medusae, they also could be lowering their abundance.

Changes of a mutualistic network over time: reanalysis over a 10-year period.

We analyzed the structure of a multispecific network of interacting ants and plants bearing extrafloral nectaries recorded in 1990 and again in 2000 in La Mancha, Veracruz, Mexico. We assessed the replicability of the number of interactions found among species and also whether there had been changes in the network structure associated with appearance of new ant and plant species during that 10-year period. Our results show that the nested topology of the network was similar between sampling dates, group dissimilarity increased, mean number of interactions for ant species increased, the frequency distribution of standardized degrees reached higher values for plant species, more ant species and fewer plant species constituted the core of the more recent network, and the presence of new ant and plant species increased while their contribution to nestedness remained the same. Generalist species (i.e., those with the most links or interactions) appeared to maintain the stability of the network because the new species incorporated into the communities were linked to this core of generalists. Camponotus planatus was the most extreme generalist ant species (the one with the most links) in both networks, followed by four other ant species; but other species changed either their position along the continuum of generalists relative to specialists or their presence or absence within the network. Even though new species moved into the area during the decade between the surveys, the overall network structure remained unmodified.

Relative contribution of ecological and biological attributes in the fine-grain structure of ant-plant networks.

BACKGROUND: Ecological communities of interacting species analyzed as complex networks have shown that species dependence on their counterparts is more complex than expected at random. As for other potentially mutualistic interactions, ant-plant networks mediated by extrafloral nectar show a nested (asymmetric) structure with a core of generalist species dominating the interaction pattern. Proposed factors structuring ecological networks include encounter probability (e.g., species abundances and habitat heterogeneity), behavior, phylogeny, and body size. While the importance of underlying factors that influence the structure of ant-plant networks have been separately explored, the simultaneous contribution of several biological and ecological attributes inherent to the species, guild or habitat level has not been addressed. METHODS: For a tropical seasonal site we recorded (in 48 censuses) the frequency of pairwise ant-plant interactions mediated by extrafloral nectaries (EFN) on different habitats and studied the resultant network structure. We addressed for the first time the role of mechanistic versus neutral determinants at the 'fine-grain' structure (pairwise interactions) of ant-plant networks. We explore the simultaneous contribution of several attributes of plant and ant species (i.e., EFN abundance and distribution, ant head length, behavioral dominance and invasive status), and habitat attributes (i.e., vegetation structure) in prevailing interactions as well as in overall network topology (community). RESULTS: Our studied network was highly-nested and non-modular, with core species having high species strengths (higher strength values for ants than plants) and low specialization. Plants had higher dependences on ants than vice versa. We found that habitat heterogeneity in vegetation structure (open vs. shaded habitats) was the main factor explaining network and fine-grain structure, with no evidence of neutral (abundance) effects. DISCUSSION: Core ant species are relevant to most plants species at the network showing adaptations to nectar consumption and deterrent behavior. Thus larger ants interact with more plant species which, together with higher dependence of plants on ants, suggests potential biotic defense at a community scale. In our study site, heterogeneity in the ant-plant interactions among habitats is so prevalent that it emerges at community-level structural properties. High frequency of morphologically diverse and temporarily-active EFNs in all habitats suggests the relevance and seasonality of plant biotic defense provided by ants. The robust survey of ecological interactions and their biological/ecological correlates that we addressed provides insight of the interplay between adaptive-value traits and neutral effects in ecological networks.

Neither ant dominance nor abundance explain ant-plant network structure in Mexican temperate forests.

BACKGROUND: Ant-plant mutualistic networks tend to have a nested structure that contributes to their stability, but the ecological factors that give rise to this structure are not fully understood. Here, we evaluate whether ant abundance and dominance hierarchy determine the structure of the ant-plant networks in two types of vegetation: oak and grassland, in two temperate environments of Mexico: Flor del Bosque State Park (FBSP) and La Malinche National Park (MNP). We predicted that dominant and abundant ant species make up the core, and submissives, the periphery of the network. We also expected a higher specialization level in the ant trophic level than in plant trophic level due to competition among the ant species for the plant-derived resources. METHODS: The ant-plant interaction network was obtained from the frequency of ant-plant interactions. We calculated a dominance hierarchy index for the ants using sampling with baits and evaluated their abundance using pitfall traps. RESULTS: In MNP, the Formica spp. species complex formed the core of the network (in both the oak forest and the grassland), while in FBSP, the core species were Prenolepis imparis (oak forest) and Camponotus rubrithorax (grassland). Although these core species were dominant in their respective sites, they were not necessarily the most dominant ant species. Three of the four networks (oak forest and grassland in FBSP, and oak forest in MNP) were nested and had a higher number of plant species than ant species. Although greater specialization was observed in the ant trophic level in the two sites and vegetations, possibly due to competition with the more dominant ant species, this was not statistically significant. In three of these networks (grassland and oak forest of MNP and oak forest of FBSP), we found no correlation between the dominance hierarchy and abundance of the ant species and their position within the network. However, a positive correlation was found between the nestedness contribution value and ant dominance hierarchy in the grassland of the site FBSP, which could be due to the richer ant-plant network and higher dominance index of this community. CONCLUSIONS: Our evidence suggests that ant abundance and dominance hierarchy have little influence on network structure in temperate ecosystems, probably due to the species-poor ant-plant network and a dominance hierarchy formed only by the presence of dominant and submissive species with no intermediate dominant species between them (absence of gradient in hierarchy) in these ecosystems.

The indirect paths to cascading effects of extinctions in mutualistic networks.

Biodiversity loss is a hallmark of our times, but predicting its consequences is challenging. Ecological interactions form complex networks with multiple direct and indirect paths through which the impacts of an extinction may propagate. Here we show that accounting for these multiple paths connecting species is necessary to predict how extinctions affect the integrity of ecological networks. Using an approach initially developed for the study of information flow, we estimate indirect effects in plant-pollinator networks and find that even those species with several direct interactions may have much of their influence over others through long indirect paths. Next, we perform extinction simulations in those networks and show that although traditional connectivity metrics fail in the prediction of coextinction patterns, accounting for indirect interaction paths allows predicting species' vulnerability to the cascading effects of an extinction event. Embracing the structural complexity of ecological systems contributes towards a more predictive ecology, which is of paramount importance amid the current biodiversity crisis.

Flower-mediated plant-butterfly interactions in an heterogeneous tropical coastal ecosystem.

BACKGROUND: Interspecific interactions play an important role in determining species richness and persistence in a given locality. However at some sites, the studies, especially for interaction networks on adult butterflies are scarce. The present study aimed the following objectives: (1) determine butterfly species richness and diversity that visit flowering plants, (2) compare species richness and diversity in butterfly-plant interactions among six different vegetation types and (3) analyze the structure of butterfly-flowering plant interaction networks mediated by flowers. METHODS: The study was developed in six vegetation types within the natural reserve of La Mancha, located in Veracruz, Mexico. In each vegetation type, we recorded the frequency of flower visits by butterflies monthly in round plots (of radius 5 m) for 12 months. We calculated Shannon diversity for butterfly species and diversity of interactions per vegetation type. We determined the classic Jaccard similarity index among vegetation types and estimated parameters at network and species-level. RESULTS: We found 123 species of butterflies belonging to 11 families and 87 genera. The highest number of species belonged to Hesperiidae (46 species), followed by Nymphalidae (28) and Pieridae (14). The highest butterfly diversity and interaction diversity was observed in pioneer dune vegetation (PDV), coastal dune scrub (CDS) and tropical deciduous flooding forest and wetland (TDF-W). The same order of vegetation types was found for interaction diversity. Highest species similarity was found between PDV-CDS and PDV-TDF. The butterfly-plant interaction network showed a nested structure with one module. The species Ascia monuste, Euptoieta hegesia and Leptotes cassius were the most generalist in the network, while Horama oedippus, E. hegesia, and L. cassius were the species with highest dependencies per plant species. DISCUSSION: Our study is important because it constitutes a pioneer study of butterfly-plant interactions in this protected area, at least for adult butterflies; it shows the diversity of interactions among flowering plants and butterflies. Our research constitutes the first approach (at a community level) to explore the functional role of pollination services that butterflies provide to plant communities. We highlighted that open areas show a higher diversity and these areas shared a higher number of species that shaded sites. In the interaction networks parameters, our results highlighted the higher dependence of butterflies by the flowers on which they feed than vice versa. In conclusion, the plant species (as a feeding resource) seem to limit the presence of butterfly species. Thus, this protected area is highly relevant for Lepidoptera diversity and the interaction between these insects and flowering plants. We suggest that studying plant and butterfly diversity in tropical habitats will provide insight into their interspecific interactions and community structure.

Effect of seed removal by ants on the host-epiphyte associations in a tropical dry forest of central Mexico.

Seed depredation is recognized as a determining factor in plant community structure and composition. Ants are primary consumers of seeds influencing abundance of epiphytes on trees. This study was conducted in two subunits of a tropical dry forest established on different soil substrates in San Andrés de la Cal, Tepoztlán, in Morelos, Mexico, and experimentally tested whether seed removal activity is higher in tree species with smaller epiphyte loads compared to those with greater epiphyte loads. Five trees were selected at random from six species of trees with high (preferred hosts) or low (limiting hosts) epiphyte loads. Seed removal differed among hosts and different soil substrates in the forest. On relating seed removal to the abundance of arboreal ants, the most consistent pattern was that lower seed removal was related to lower ant abundance, while high seed removal was associated with intermediate to high ant abundance. Epiphyte seed removal by ants influences epiphyte abundance and can contribute considerably to a failure to establish, since it diminishes the quantity of seeds available for germination and establishment.

Temporal changes in the structure of a plant-frugivore network are influenced by bird migration and fruit availability.

Background. Ecological communities are dynamic collections whose composition and structure change over time, making up complex interspecific interaction networks. Mutualistic plant-animal networks can be approached through complex network analysis; these networks are characterized by a nested structure consisting of a core of generalist species, which endows the network with stability and robustness against disturbance. Those mutualistic network structures can vary as a consequence of seasonal fluctuations and food availability, as well as the arrival of new species into the system that might disorder the mutualistic network structure (e.g., a decrease in nested pattern). However, there is no assessment on how the arrival of migratory species into seasonal tropical systems can modify such patterns. Emergent and fine structural temporal patterns are adressed here for the first time for plant-frugivorous bird networks in a highly seasonal tropical environment. Methods. In a plant-frugivorous bird community, we analyzed the temporal turnover of bird species comprising the network core and periphery of ten temporal interaction networks resulting from different bird migration periods. Additionally, we evaluated how fruit abundance and richness, as well as the arrival of migratory birds into the system, explained the temporal changes in network parameters such as network size, connectance, nestedness, specialization, interaction strength asymmetry and niche overlap. The analysis included data from 10 quantitative plant-frugivorous bird networks registered from November 2013 to November 2014. Results. We registered a total of 319 interactions between 42 plant species and 44 frugivorous bird species; only ten bird species were part of the network core. We witnessed a noteworthy turnover of the species comprising the network periphery during migration periods, as opposed to the network core, which did not show significant temporal changes in species composition. Our results revealed that migration and fruit richness explain the temporal variations in network size, connectance, nestedness and interaction strength asymmetry. On the other hand, fruit abundance only explained connectance and nestedness. Discussion. By means of a fine-resolution temporal analysis, we evidenced for the first time how temporal changes in the interaction network structure respond to the arrival of migratory species into the system and to fruit availability. Additionally, few migratory bird species are important links for structuring networks, while most of them were peripheral species. We showed the relevance of studying bird-plant interactions at fine temporal scales, considering changing scenarios of species composition with a quantitative network approach.

Association patterns in saproxylic insect networks in three Iberian Mediterranean woodlands and their resistance to microhabitat loss.

The assessment of the relationship between species diversity, species interactions and environmental characteristics is indispensable for understanding network architecture and ecological distribution in complex networks. Saproxylic insect communities inhabiting tree hollow microhabitats within Mediterranean woodlands are highly dependent on woodland configuration and on microhabitat supply they harbor, so can be studied under the network analysis perspective. We assessed the differences in interacting patterns according to woodland site, and analysed the importance of functional species in modelling network architecture. We then evaluated their implications for saproxylic assemblages' persistence, through simulations of three possible scenarios of loss of tree hollow microhabitat. Tree hollow-saproxylic insect networks per woodland site presented a significant nested pattern. Those woodlands with higher complexity of tree individuals and tree hollow microhabitats also housed higher species/interactions diversity and complexity of saproxylic networks, and exhibited a higher degree of nestedness, suggesting that a higher woodland complexity positively influences saproxylic diversity and interaction complexity, thus determining higher degree of nestedness. Moreover, the number of insects acting as key interconnectors (nodes falling into the core region, using core/periphery tests) was similar among woodland sites, but the species identity varied on each. Such differences in insect core composition among woodland sites suggest the functional role they depict at woodland scale. Tree hollows acting as core corresponded with large tree hollows near the ground and simultaneously housing various breeding microsites, whereas core insects were species mediating relevant ecological interactions within saproxylic communities, e.g. predation, competitive or facilitation interactions. Differences in network patterns and tree hollow characteristics among woodland sites clearly defined different sensitivity to microhabitat loss, and higher saproxylic diversity and woodland complexity showed positive relation with robustness. These results highlight that woodland complexity goes hand in hand with biotic and ecological complexity of saproxylic networks, and together exhibited positive effects on network robustness.

Evaluating the spatio-temporal factors that structure network parameters of plant-herbivore interactions.

Despite the dynamic nature of ecological interactions, most studies on species networks offer static representations of their structure, constraining our understanding of the ecological mechanisms involved in their spatio-temporal stability. This is the first study to evaluate plant-herbivore interaction networks on a small spatio-temporal scale. Specifically, we simultaneously assessed the effect of host plant availability, habitat complexity and seasonality on the structure of plant-herbivore networks in a coastal tropical ecosystem. Our results revealed that changes in the host plant community resulting from seasonality and habitat structure are reflected not only in the herbivore community, but also in the emergent properties (network parameters) of the plant-herbivore interaction network such as connectance, selectiveness and modularity. Habitat conditions and periods that are most stressful favored the presence of less selective and susceptible herbivore species, resulting in increased connectance within networks. In contrast, the high degree of selectivennes (i.e. interaction specialization) and modularity of the networks under less stressful conditions was promoted by the diversification in resource use by herbivores. By analyzing networks at a small spatio-temporal scale we identified the ecological factors structuring this network such as habitat complexity and seasonality. Our research offers new evidence on the role of abiotic and biotic factors in the variation of the properties of species interaction networks.

Individual-based ant-plant networks: diurnal-nocturnal structure and species-area relationship.

Despite the importance and increasing knowledge of ecological networks, sampling effort and intrapopulation variation has been widely overlooked. Using continuous daily sampling of ants visiting three plant species in the Brazilian Neotropical savanna, we evaluated for the first time the topological structure over 24 h and species-area relationships (based on the number of extrafloral nectaries available) in individual-based ant-plant networks. We observed that diurnal and nocturnal ant-plant networks exhibited the same pattern of interactions: a nested and non-modular pattern and an average level of network specialization. Despite the high similarity in the ants' composition between the two collection periods, ant species found in the central core of highly interacting species totally changed between diurnal and nocturnal sampling for all plant species. In other words, this "night-turnover" suggests that the ecological dynamics of these ant-plant interactions can be temporally partitioned (day and night) at a small spatial scale. Thus, it is possible that in some cases processes shaping mutualistic networks formed by protective ants and plants may be underestimated by diurnal sampling alone. Moreover, we did not observe any effect of the number of extrafloral nectaries on ant richness and their foraging on such plants in any of the studied ant-plant networks. We hypothesize that competitively superior ants could monopolize individual plants and allow the coexistence of only a few other ant species, however, other alternative hypotheses are also discussed. Thus, sampling period and species-area relationship produces basic information that increases our confidence in how individual-based ant-plant networks are structured, and the need to consider nocturnal records in ant-plant network sampling design so as to decrease inappropriate inferences.

Breaking down complex Saproxylic communities: understanding sub-networks structure and implications to network robustness.

Saproxylic insect communities inhabiting tree hollow microhabitats correspond with large food webs which simultaneously are constituted by multiple types of plant-animal and animal-animal interactions, according to the use of trophic resources (wood- and insect-dependent sub-networks), or to trophic habits or interaction types (xylophagous, saprophagous, xylomycetophagous, predators and commensals). We quantitatively assessed which properties of specialised networks were present in a complex networks involving different interacting types such as saproxylic community, and how they can be organised in trophic food webs. The architecture, interacting patterns and food web composition were evaluated along sub-networks, analysing their implications to network robustness from random and directed extinction simulations. A structure of large and cohesive modules with weakly connected nodes was observed throughout saproxylic sub-networks, composing the main food webs constituting this community. Insect-dependent sub-networks were more modular than wood-dependent sub-networks. Wood-dependent sub-networks presented higher species degree, connectance, links, linkage density, interaction strength, and were less specialised and more aggregated than insect-dependent sub-networks. These attributes defined high network robustness in wood-dependent sub-networks. Finally, our results emphasise the relevance of modularity, differences among interacting types and interrelations among them in modelling the structure of saproxylic communities and in determining their stability.