Denise Lange

Influence of extrafloral nectary phenology on ant–plant mutualistic networks in a neotropical savanna

Temporal variation has been one remarkable feature of ecological interactions. In ant–plant mutualism, this variation is widely known, although little is understood about the mechanisms that shape these variations. This study tested whether or not the phenology of extrafloral nectaries (EFNs) influences the temporal variation of two properties of an ant–plant interaction network. The network under investigation exhibited a nested pattern and low specialisation over most months. Monthly nestedness and specialisation in the network were negatively correlated, both being influenced by temporal variations in extrafloral nectar production of the plant community. The months of highest activity in the nectaries (August–November) were those when the level of generalisation in the network was at its highest. Although there were temporal variations in the properties of the network, the generalist core of the species remained the same over time. The stable core enhances the coevolutionary importance of ant–plant interactions for the community. Thus, it can be concluded that the phenology of EFNs is one effective mechanism shaping the temporal variation in ant–plant interaction.

Ant-plant interaction in a tropical savanna: may the network structure vary over time and influence on the outcomes of associations?

Plant-animal interactions occur in a community context of dynamic and complex ecological interactive networks. The understanding of who interacts with whom is a basic information, but the outcomes of interactions among associates are fundamental to draw valid conclusions about the functional structure of the network. Ecological networks studies in general gave little importance to know the true outcomes of interactions and how they may change over time. We evaluate the dynamic of an interaction network between ants and plants with extrafloral nectaries, by verifying the temporal variation in structure and outcomes of mutualism for the plant community (leaf herbivory). To reach this goal, we used two tools: bipartite network analysis and experimental manipulation. The networks exhibited the same general pattern as other mutualistic networks: nestedness, asymmetry and low specialization and this pattern was maintained over time, but with internal changes (species degree, connectance and ant abundance). These changes influenced the protection effectiveness of plants by ants, which varied over time. Our study shows that interaction networks between ants and plants are dynamic over time, and that these alterations affect the outcomes of mutualisms. In addition, our study proposes that the set of single systems that shape ecological networks can be manipulated for a greater understanding of the entire system.

Loss and gains in ant–plant interactions mediated by extrafloral nectar: fidelity, cheats, and lies

All mutualistic plant–animal interactions are mediated by costs and benefits in relationships where resources (from plants) are exchanged by services (from animals). The most common trading coin that plants offer to pay for animal services is nectar; the main servers are hymenopterans. Extrafloral nectar (EFN) is produced in almost all aboveground plant parts not directly related with pollination, and their true function has long been an issue of discussion among naturalists and will be our main subject. The protective function of extrafloral nectaries (EFNs) is reviewed and considered with an alternative hypothesis, presenting not only ants, but also spiders and wasps as potential and effective agents in these protective interactions. Despite their likely relevance, the phenological variation (mainly sequential flowering and resprouting) of host plants mediating these interactions have been generally ignored. We discuss how the outcomes of each ant–EFN bearing plant interaction vary depending on physical and biotic changes in interacting organisms (internal factors such as phenology and species identity) as well as in their environments (external factors such as climatic variation), all of which may modify the character of each interaction. We propose that ant–EFN bearing plant interactions serve an excellent and unique model to test the “Geographic Mosaic Theory” of coevolution providing us a more clear view of how evolution has structured these plant–animal ecological networks.

Importance of interaction frequency in analysis of ant-plant networks in tropical environments

Several studies have shown that qualitative (binary) ant-plant networks are highly nested in tropical environments, in which specialist species (with fewer interactions) are connected with generalists (with the most interactions) in cohesive subgroups. Interactions occur in both qualitative and quantitative networks, however, how their frequency may structure the nestedness in ecological networks involving these organisms is, we believe, unknown. Based on this perspective, we used nestedness analysis to address the effect of interaction frequency on antplant networks (n = 14 networks). Unlike binary networks, quantitative networks are often significantly non-nested. In addition, species with a higher interaction frequency have a higher number of links, indicating that these species are possibly more abundant and/or competitive. Moreover, different biological parameters can change the nature of ant-plant interactions, as a plant can be a good resource for one ant and a ‘bad’ resource for another. Thus, this suggests a new perspective for the study of interaction networks in the tropics, since species with lower interaction frequency are not necessarily subsets of species with higher frequency, and consequently generate the non-nested pattern in quantitative networks.

Flower-Visiting Social Wasps and Plants Interaction: Network Pattern and Environmental Complexity

Network analysis as a tool for ecological interactions studies has been widely used since last decade. However, there are few studies on the factors that shape network patterns in communities. In this sense, we compared the topological properties of the interaction network between flower-visiting social wasps and plants in two distinct phytophysiognomies in a Brazilian savanna (Riparian Forest and Rocky Grassland). Results showed that the landscapes differed in species richness and composition, and also the interaction networks between wasps and plants had different patterns. The network was more complex in the Riparian Forest, with a larger number of species and individuals and a greater amount of connections between them. The network specialization degree was more generalist in the Riparian Forest than in the Rocky Grassland. This result was corroborated by means of the nestedness index. In both networks was found asymmetry, with a large number of wasps per plant species. In general aspects, most wasps had low niche amplitude, visiting from one to three plant species. Our results suggest that differences in structural complexity of the environment directly influence the structure of the interaction network between flower-visiting social wasps and plants.

Variation in Extrafloral Nectary Productivity Influences the Ant Foraging

Extrafloral nectar is the main food source offered by plants to predatory ants in most land environments. Although many studies have demonstrated the importance of extrafloral nectaries (EFNs) to plant defense against herbivores, the influence of EFNs secretory activity pattern on predatory ants remains yet not fully understood. Here, we verified the relation between the extrafloral nectar production of a plant community in Cerrado in different times of the day, and its attractiveness to ants. The extrafloral nectaries (EFNs) of seven plant species showed higher productivity overnight. Ant abundance was higher in times of large extrafloral nectar production, however, there was no positive relation between ant richness on plants and EFNs productivity. There was temporal resource partitioning among ant species, and it indicates strong resource competition. The nectar productivity varied among plant species and time of the day, and it influenced the visitation patterns of ants. Therefore, EFNs are a key ant-plant interaction driver in the studied system.

Influence, or the lack thereof, of host phenology, architecture and climate on the occurrence of Udranomia spitzi (Hesperiidae: Lepidoptera)

The occurrence of herbivores in nature is limited by biotic and abiotic factors which affect their development and survival. Udranomia spitzi is an endemic butterfly in the Brazilian savanna (cerrado) that feeds on young leaves of two sympatric plants, Ouratea hexasperma and O. spectabilis. It is not known which factors affect the occurrence of larvae on their hosts. Therefore, in this study we: (i) evaluated the oviposition preference of U. spitzi; (ii) evaluated the larval performance in both Ouratea species; (iii) investigated plant phenology; (iv) investigated climate (temperature and rainfall); and (v) investigated plant architecture (measured as plant height) on the abundance of skippers. Results showed that U. spitzi immatures (eggs and larvae) were far more abundant (n = 41, 96.7%) on O. spectabilis, whereas on O. hexasperma, the number of larvae was negligible (n = 1). In the laboratory, U. spitzi performed better on O. spectabilis than on O. hexasperma. The occurrence of larvae was not related to host phenology or environmental variations, but rather to plant height, since 92.7% (n = 38) of larvae were found on small O. spectabilis trees. A previous study showed that U. spitzi was not influenced by biotic factors (aggressive ants) and this study showed that plant structure plays a major role in skipper choice. The preference of U. spitzi for O. spectabilis is discussed.

Predation of Fruit Fly Larvae Anastrepha (Diptera: Tephritidae) by Ants in Grove

Based on evidence that ants are population regulatory agents, we examined their efficiency in predation of fruit fly larvae Anastrepha Schiner, 1868 (Diptera: Tephritidae). Hence, we considered the differences among species of fruit trees, the degree of soil compaction, and the content of soil moisture as variables that would explain predation by ants because these variables affect burying time of larvae. We carried out the experiment in an orchard containing various fruit bearing trees, of which the guava (Psidium guajava Linn.), jaboticaba (Myrciaria jaboticaba (Vell.) Berg.), and mango trees (Mangifera indica Linn.) were chosen for observations of Anastrepha. We offered live Anastrepha larvae on soil beneath the tree crowns. We observed for 10 min whether ants removed the larvae or the larvae buried themselves. Eight ant species were responsible for removing 1/4 of the larvae offered. The Pheidole Westwood, 1839 ants were the most efficient genus, removing 93% of the larvae. In compacted and dry soils, the rate of predation by ants was greater. Therefore, this study showed that ants, along with specific soil characteristics, may be important regulators of fruit fly populations and contribute to natural pest control in orchards.

The Complex Ant–Plant Relationship Within Tropical Ecological Networks

The tools involved in the study of ecological networks are relatively new and very useful to improve the knowledge about communities, biodiversity, and their conservation. In many tropical habitats, ants form the major part of the arthropod fauna found on vegetation and, therefore, it is extremely common to observe ants establishing ecological interactions with the host plants, where they find and use nectar, oils, pollen, arils, and seeds as food resources. In this chapter, we show that ant–plant interactions are dynamic, diverse, worldwide spread, and very manipulative which fit perfectly as models in studies dealing with interaction networks. For this, we have conducted global review in the distribution of studies on ant–plant networks and highlighted the most recurrent structural patterns observed in ant–plant networks and the main mechanisms and process behind this structure. Finally, we pointed out the limitations and new directions for the study of ant–plant networks in tropical environments.

Might Heterostyly Underlie Spider Occurrence on Inflorescences? A Case Study of Palicourea rigida (Rubiaceae), a Common Shrub from Brazilian Cerrado

We carried out a research on thePalicourea rigida (Rubiaceae) inflorescences, a distylous shrub of Brazilian Cerrado. Our objective was to compare the inflorescence architectural complexity and its quality in the two floral morphs and search for any relationship with spider occurrence. In order to assess the quality of inflorescence resources, we quantified the nectar volume and its sugar concentration and the number of fruits and flowers (intact and aborted) for both inflorescence morphs with and without spiders. For the architectural heterogeneity, we quantified floral structures and inflorescence levels of branching. Spider occurrence was higher in longistylous inflorescences than in brevistylous ones. The sampled spiders were classified into the guilds ambushers, jumpers, or orb-weavers. Ambushers, jumpers, and total richness were much higher among longistylous inflorescences. We found no difference between morphs neither in volume or nectar concentration nor in amount of fruits and flowers. However, longistylous inflorescences presented greater architectural heterogeneity than brevistylous ones. Therefore, we suggested that architectural heterogeneity is an important factor underlying the occurrence of cursorial spiders onP. rigida inflorescences, which possibly arose from the relationship between refuge availability and inflorescence architecture.