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      Variation in Extrafloral Nectary Productivity Influences the Ant Foraging

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      Author(s): 1 , 2 , 3 , *
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      Publication date (Electronic):
      Journal: PLoS ONE
      Publisher: Public Library of Science

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          Abstract

          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.

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          Ants on plants: a meta-analysis of the role of ants as plant biotic defenses.

          Felix Rosumek, Fernando Silveira, Frederico de S Neves … (2009)
          We reviewed the evidence on the role of ants as plant biotic defenses, by conducting meta-analyses for the effects of experimental removal of ants on plant herbivory and fitness with data pooled from 81 studies. Effects reviewed were plant herbivory, herbivore abundance, hemipteran abundance, predator abundance, plant biomass and reproduction in studies where ants were experimentally removed (n = 273 independent comparisons). Ant removal exhibited strong effects on herbivory rates, as plants without ants suffered almost twice as much damage and exhibited 50% more herbivores than plants with ants. Ants also influenced several parameters of plant fitness, as plants without ants suffered a reduction in biomass (-23.7%), leaf production (-51.8%), and reproduction (-24.3%). Effects were much stronger in tropical regions compared to temperate ones. Tropical plants suffered almost threefold higher herbivore damage than plants from temperate regions and exhibited three times more herbivores. Ant removal in tropical plants resulted in a decrease in plant fitness of about 59%, whereas in temperate plants this reduction was not statistically significant. Ant removal effects were also more important in obligate ant-plants (=myrmecophytes) compared to plants exhibiting facultative relationships with hemiptera or those plants with extrafloral nectaries and food bodies. When only tropical plants were considered and the strength of the association between ants and plants taken into account, plants with obligate association with ants exhibited almost four times higher herbivory compared to plants with facultative associations with ants, but similar reductions in plant reproduction. The removal of a single ant species increased plant herbivory by almost three times compared to the removal of several ant species. Altogether, these results suggest that ants do act as plant biotic defenses, but the effects of their presence are more pronounced in tropical systems, especially in myrmecophytic plants.
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            How plants shape the ant community in the Amazonian rainforest canopy: the key role of extrafloral nectaries and homopteran honeydew.

            W Barthlott, M. Verhaagh, W Morawetz … (2000)
            Ant-plant interactions in the canopy of a lowland Amazonian rainforest of the upper Orinoco, Venezuela, were studied using a modified commercial crane on rails (Surumoni project). Our observations show a strong correlation between plant sap exudates and both abundance of ants and co-occurrence of ant species in tree canopies. Two types of plant sap sources were compared: extrafloral nectaries (EFNs) and honeydew secretions by homopterans. EFNs were a frequent food source for ants on epiphytes (Philodendron spp., Araceae) and lianas (Dioclea, Fabaceae), but rare on canopy trees in the study area, whereas the majority of trees were host to aggregations of homopterans tended by honeydew-seeking ants (on 62% of the trees examined). These aggregations rarely occurred on epiphytes. Baited ant traps were installed on plants with EFNs and in the crowns of trees from three common genera, including trees with and without ant-tended homopterans: Goupia glabra (Celastraceae), Vochysia spp. (Vochysiaceae), and Xylopia spp. (Annonaceae). The number of ant workers per trap was significantly higher on plants offering one of the two plant sap sources than on trees without such resources. Extrafloral nectaries were used by a much broader spectrum of ant species and genera than honeydew, and co-occurrence of ant species (in traps) was significantly higher on plants bearing EFNs than on trees. Homopteran honeydew (Coccidae and Membracidae), on the other hand, was mostly monopolised by a single ant colony per tree. Homopteran-tending ants were generally among the most dominant ants in the canopy. The most prominent genera were Azteca, Dolichoderus (both Dolichoderinae), Cephalotes, Pheidole, Crematogaster (all Myrmicinae), and Ectatomma (Ponerinae). Potential preferences were recorded between ant and homopteran species, and also between ant-homopteran associations and tree genera. We hypothesize that the high availability of homopteran honeydew provides a key resource for ant mosaics, where dominant ant colonies and species maintain mutually exclusive territories on trees. In turn, we propose that for nourishment of numerous ants of lower competitive capacity, Philodendron and other sources of EFNs might be particularly important.
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              Extrafloral nectar at the plant-insect interface: a spotlight on chemical ecology, phenotypic plasticity, and food webs.

              Martin Heil (2015)
              Plants secrete extrafloral nectar (EFN) as an induced defense against herbivores. EFN contains not only carbohydrates and amino acids but also pathogenesis-related proteins and other protective enzymes, making EFN an exclusive reward. EFN secretion is commonly induced after wounding, likely owing to a jasmonic acid-induced cell wall invertase, and is limited by phloem sucrose availability: Both factors control EFN secretion according to the optimal defense hypothesis. Non-ant EFN consumers include parasitoids, wasps, spiders, mites, bugs, and predatory beetles. Little is known about the relevance of EFN to the nutrition of its consumers and, hence, to the structuring of arthropod communities. The mutualism can be established quickly among noncoevolved (e.g., invasive) species, indicating its easy assembly is due to ecological fitting. Therefore, increasing efforts are directed toward using EFN in biocontrol. However, documentation of the importance of EFN for the communities of plants and arthropods in natural, invasive, and agricultural ecosystems is still limited.
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                Author and article information

                Contributors
                Renee M. Borges: Role: Editor
                Journal
                Journal ID (nlm-ta): PLoS One
                Journal ID (iso-abbrev): PLoS ONE
                Journal ID (publisher-id): plos
                Journal ID (pmc): plosone
                Title: PLoS ONE
                Publisher: Public Library of Science (San Francisco, CA USA )
                ISSN (Electronic): 1932-6203
                Publication date (Electronic): 3 January 2017
                Publication date Collection: 2017
                Volume: 12
                Issue: 1
                Electronic Location Identifier: e0169492
                Affiliations
                [1 ]Universidade Tecnológica Federal do Paraná, Campus Santa Helena, Santa Helena, PR, Brazil
                [2 ]Pós-Graduação em Entomologia, Faculdade de Filosofia, Ciências e Letras, Universidade de São Paulo, Ribeirão Preto, SP, Brazil
                [3 ]Laboratório de Ecologia Comportamental e de Interações (LECI), Instituto de Biologia, Universidade Federal de Uberlândia, Uberlândia, MG, Brazil
                Indian Institute of Science, INDIA
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                • Conceptualization: DL ESC KDC.

                • Data curation: DL ESC KDC.

                • Formal analysis: DL.

                • Funding acquisition: KDC.

                • Investigation: DL ESC KDC.

                • Methodology: DL KDC.

                • Project administration: KDC.

                • Supervision: KDC.

                • Visualization: DL ESC KDC.

                • Writing – original draft: DL ESC KDC.

                • Writing – review & editing: DL ESC KDC.

                Author information
                http://orcid.org/0000-0001-8886-9568
                Article
                Publisher ID: PONE-D-16-37581
                DOI: 10.1371/journal.pone.0169492
                PMC ID: 5207748
                PubMed ID: 28046069
                SO-VID: cbf835c4-a07e-46f4-a043-dafb9c9033a6
                Copyright © © 2017 Lange et al
                License:

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                Date received : 19 September 2016
                Date accepted : 16 December 2016
                Page count
                Figures: 6, Tables: 2, Pages: 13
                Funding
                The authors do not have any specific funding that supported this research.
                Categories
                Subject: Research Article
                Subject: Biology and Life Sciences
                Subject: Organisms
                Subject: Animals
                Subject: Invertebrates
                Subject: Arthropoda
                Subject: Insects
                Subject: Hymenoptera
                Subject: Ants
                Subject: Biology and Life Sciences
                Subject: Organisms
                Subject: Plants
                Subject: Biology and Life Sciences
                Subject: Plant Science
                Subject: Plant Anatomy
                Subject: Nectaries
                Subject: Biology and Life Sciences
                Subject: Behavior
                Subject: Animal Behavior
                Subject: Foraging
                Subject: Biology and Life Sciences
                Subject: Zoology
                Subject: Animal Behavior
                Subject: Foraging
                Subject: Biology and Life Sciences
                Subject: Ecology
                Subject: Plant Ecology
                Subject: Plant-Animal Interactions
                Subject: Plant-Herbivore Interactions
                Subject: Ecology and Environmental Sciences
                Subject: Ecology
                Subject: Plant Ecology
                Subject: Plant-Animal Interactions
                Subject: Plant-Herbivore Interactions
                Subject: Biology and Life Sciences
                Subject: Plant Science
                Subject: Plant Ecology
                Subject: Plant-Animal Interactions
                Subject: Plant-Herbivore Interactions
                Subject: Biology and Life Sciences
                Subject: Ecology
                Subject: Plant Ecology
                Subject: Plant-Animal Interactions
                Subject: Herbivory
                Subject: Ecology and Environmental Sciences
                Subject: Ecology
                Subject: Plant Ecology
                Subject: Plant-Animal Interactions
                Subject: Herbivory
                Subject: Biology and Life Sciences
                Subject: Plant Science
                Subject: Plant Ecology
                Subject: Plant-Animal Interactions
                Subject: Herbivory
                Subject: Biology and Life Sciences
                Subject: Ecology
                Subject: Community Ecology
                Subject: Trophic Interactions
                Subject: Herbivory
                Subject: Ecology and Environmental Sciences
                Subject: Ecology
                Subject: Community Ecology
                Subject: Trophic Interactions
                Subject: Herbivory
                Subject: Biology and Life Sciences
                Subject: Plant Science
                Subject: Plant Physiology
                Subject: Plant Defenses
                Subject: Biology and Life Sciences
                Subject: Organisms
                Subject: Plants
                Subject: Vascular Plants
                Custom metadata
                Data Availability All relevant data are within the paper.

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