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Ant–Treehopper Associations Deter Pollinators and Reduce Reproduction in a Tropical Shrub

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Ant–Treehopper Associations Deter Pollinators and Reduce Reproduction in a Tropical Shrub Indirect effects of mutualism: ant– treehopper associations deter pollinators and reduce reproduction in a tropical shrub Javier Ibarra-Isassi & Paulo S. Oliveira Oecologia ISSN 0029-8549 Volume 186 Number 3 Oecologia (2018) 186:691-701 DOI 10.1007/s00442-017-4045-7 1 23 Oecologia (2018) 186:691–701 https://doi.org/10.1007/s00442-017-4045-7 PLANT-MICROBE-ANIMAL INTERACTIONS - ORIGINAL RESEARCH Indirect efects of mutualism: ant–treehopper associations deter pollinators and reduce reproduction in a tropical shrub Javier Ibarra‑Isassi1,2 · Paulo S. Oliveira1 Received: 22 September 2017 / Accepted: 10 December 2017 / Published online: 15 December 2017 © Springer-Verlag GmbH Germany, part of Springer Nature 2017 Abstract Animal-pollinated plants can be susceptible to changes in pollinator availability. Honeydew-producing treehoppers fre- quently occur on inforescences, potentially enhancing ant-mediated negative efects on pollination services. However, the efect of ant-attended, honeydew-producing insects on plant reproduction remains uncertain. We recorded the abundance of treehoppers and ants on Byrsonima intermedia (Malpighiaceae), and monitored foral visitors in a Brazilian cerrado savanna. We manipulated the presence of ants and ant–treehopper associations on inforescences to assess their efect on pollination and fruit formation. We used dried ants pinned to inforescences to evaluate the efect of ant presence and ant identity on potential pollinators. Results show that the presence of treehoppers increases ant abundance on fowers and disrupts pollination by oil-collecting bees, decreasing the frequency and duration of foral visits and reducing fruit and seed set. Treehopper herbivory has no direct efect on fruit or seed production, which are independent of treehopper density. Pinned ants promote avoidance by foral visitors, reducing the number of visits. Ant identity mediates visitation decisions, with Ectatomma brunneum causing greater avoidance by foral visitors than Camponotus rufpes. Field videos show that pollinating bees are harassed by ants near fowers, prompting avoidance behavior by the bees. This is the frst demonstration of indirect efects by honeydew-gathering ants, via disrupted pollination, on plant reproduction in tropical cerrado savanna. Our results highlight the importance of studying other interactions near fowers, in addition to just observing pollinators, for a proper understanding of plant reproduction. Keywords Ant–plant–herbivore interaction · Byrsonima · Cerrado savanna · Oil-collecting bees · Flower avoidance behavior Introduction may be one of the most ecologically important types of ani- mal–plant interactions, and about 87% of fowering plant Arthropods and fowering plants are among the most abun- species are animal pollinated (Ollerton et al. 2011). In addi- dant organisms in terrestrial landscapes and their life histo- tion to anthropogenic disturbance that can jeopardize pol- ries are bound to cross in many environments (Price et al. lination services, the presence of other organisms near fow- 2011). The relationships between plants and pollinators ers and their interactions with potential pollinators can also afect the fnal outcome of fower visitation (Romero et al. 2011). Thus, the study of ecological factors afecting animal Communicated by Susan Whitehead. pollination is becoming increasingly important. Electronic supplementary material The online version of this Plant species have diferent degrees of fexibility regard- article (https://doi.org/10.1007/s00442-017-4045-7) contains ing pollinators, being able to be pollinated by a single or supplementary material, which is available to authorized users. by several species (Waser et al. 1996; Alarcón et al. 2008). * Paulo S. Oliveira In highly diverse communities, however, negative interac- [email protected] tions between foral visitors may disrupt pollination ser- vices (Alarcón et al. 2008). The relationship between plants 1 Departamento de Biologia Animal, Universidade Estadual and low-efciency pollinators can vary from commensal- de Campinas, Campinas, SP 13083‑862, Brazil ism to antagonism (Lau and Galloway 2004). In addition, 2 Present Address: Department of Biology, Concordia these negative interactions involving foral visitors include University, Montreal, QC H4B‑1R6, Canada Vol.:(0123456789)1 3 692 Oecologia (2018) 186:691–701 predation by arthropods such as wasps and spiders, which (Solenopsis invicta) and cotton aphids (Aphis gossypii) may can afect pollination services (and in turn fruit formation) enhance cotton plant (Gossypium hirsutum) reproduction by by harassing pollinators and prompting fower avoidance reducing non-aphid herbivore damage (Styrsky and Eubanks behavior (Romero et al. 2011). The interaction between 2010). In contrast, LeVan and Holway (2015) demonstrated mutualisms occurring near inforescences (such as ants–tree- that ant–aphid interactions negatively afect cotton plant hoppers and pollinators–fowers) and their efect on plant reproduction, because the ants harass other foral visitors. reproduction remain understudied (Lach 2007; Styrsky and Here, we investigate if associations between ants and Eubanks 2010; LeVan and Holway 2015). honeydew-producing treehoppers (Membracidae) on info- Although ants are often observed visiting fowers in a rescences of Byrsonima intermedia (Malpighiaceae) afect variety of environments, the outcomes of these interactions bee pollination (Fig. 1) and plant reproduction in a reserve are frequently reported as negative for the fowers, rarely of cerrado savanna in southeast Brazil. We manipulated the resulting in efective pollination (Beattie 2007; Rico-Gray presence/absence of ants and treehoppers to determine their and Oliveira 2007). Furthermore, foral visitation by ants can efect on the behavior of bee pollinators, as well as on the result in reduced pollen viability when pollen grains come reproductive output of the plant. Specifcally, our observa- into contact with antibiotic substances on ant integuments tions under natural conditions and experimental manipula- (Beattie et al. 1984; Dutton and Frederickson 2012). Ants tions assessed (1) how the presence of treehoppers afects may also cause damage to reproductive structures (Palmer ant abundance near fowers of B. intermedia; (2) how ant et al. 2010), harassment of foral visitors (Tsuji et al. 2004; presence near the fowers and ant species identity afect visi- Ness 2006), avoidance or alteration of floral visitation tation behavior of potential pollinators (evaluated as duration behavior in response to olfactory (Li et al. 2014) and visual of visits and number of approaches to fowers); and (3) how cues (Cembrowski et al. 2014), and depletion or modifca- ant–treehopper associations afect fruit and seed set. tion of foral resources (Lach 2008; de Vega and Herrera 2013). However, foral visitation by ants may also result in positive outcomes for the plant if the ants can deter less Materials and methods efective pollinators (Gonzálvez et al. 2013), foral herbi- vores (Oliveira 1997; Bleil et al. 2011), or even pollinate the Study area and system fowers (see review in de Vega and Gómez 2014). Although foral visitation by ants is pervasive in almost every terres- Fieldwork was carried out at the cerrado reserve of the trial environment, few studies have assessed how ant–fower Estação Ecológica de Itirapina (22º14′43″S, 47º53′23″W), interactions can ultimately afect plant reproduction (Hanna in the state of São Paulo, southeast Brazil. The vegetation et al. 2015; LeVan and Holway 2015). Moreover, the impor- physiognomy corresponds to a cerrado sensu stricto, charac- tance of the identity of the ant species as a factor afecting terized by a dense scrub of shrubs and trees in an herbaceous plant reproduction remains uncertain (see Ness 2006). matrix (Oliveira-Filho and Ratter 2002). The climate of the Given that the presence of honeydew-producing insects (i.e., trophobionts) is known to increase ant visitation to plants (Del-Claro and Oliveira 1996), foral visitation by ants can be enhanced by honeydew-producing insects that feed near fowers. Yet, the ecological consequences for the plant remain unclear (Styrsky and Eubanks 2007). Ants tending honeydew-producing insects often drive of other insects (predators, parasitoids, other herbivores, etc.), causing a pos- itive efect on the trophobionts as well as an indirect positive efect on the plant by reducing herbivore damage (Oliveira and Del-Claro 2005; Pringle et al. 2011). Even though ants tending honeydew-producing insects may be distracted from foraging any foral resource (e.g., nectar), ants are com- monly found on inforescences (Lach 2007). Moreover, and despite the ubiquity of ant-mediated interspecifc interac- tions near fowers in many terrestrial ecosystems, few studies Fig. 1 a Oil-collecting bee, Centris varia, approaches an infores- have experimentally assessed the efects of ant–trophobiont cence of Byrsonima intermedia occupied by workers of Camponotus rufpes, which attend nymphs of honeydew-producing Amastris associations on pollinators and their consequences for plant b undulata treehoppers. c Bees pollinate the fowers of B. interme- reproduction. Previous work focusing on ant–aphid associa- dia while collecting oil at the calyx glands (arrow). Photo credits: J. tions demonstrated that the interaction between red fre ants Ibarra-Isassi (a, b); H. Soares Jr (c) 1 3 Oecologia (2018) 186:691–701 693 region is characterized by a rainy/warm season from Novem- April 2015, during
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