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Fire and Leaf Fungus Infection Reduce Benefits to Plants with Extrafloral Nectarines Author(S): L Variation in the Outcomes of an Ant-Plant System: Fire and Leaf Fungus Infection Reduce Benefits to Plants with Extrafloral Nectarines Author(s): L. P. Pires and K. Del-Claro Source: Journal of Insect Science, 14(84):1-10. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1673/031.014.84 URL: http://www.bioone.org/doi/full/10.1673/031.014.84 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Journal of Insect Science: Vol. 14 | Article 84 Pires and Del-Claro Variation in the outcomes of an ant-plant system: Fire and leaf fungus infection reduce benefits to plants with extrafloral nectaries L. P. Piresa and K. Del-Clarob* Universidade Federal de Uberlândia, Laboratório de Ecologia Comportamental e de Interação, Instituto de Biologia, MG, Brazil Abstract Interactions between species are evolutionary malleable and may suffer changes in small timescales. Environmental disturbances, such as fire, can deeply affect species interactions, but how they influence the outcome of a mutualistic interaction has yet to be studied. In order to test the hypothesis that an environmental disturbance, in this case fire, may produce differences in the outcome of the association of ants with the extrafloral-nectaries-bearing plant Qualea multiflora Mart. (Myrtales: Vochysiaceae), a previous study was replicated, but this time after fire incidence, at the same study site and with the same plant species. Eight ant species visited Q. multiflora, and the most abundant genera were Crematogaster, Cephalotes, and Camponotus. Herbivores were found in branches with and without ants with no statistical difference, but foliar herbivory was always higher in branchs where ants were absent. Leaves were infested by fungi, and fungi spots were higher in branches where ants were present. Compared to the previous study, it was clearly observed that ant benefits to Q. multiflora varied over time. The most common ant species still protected leaves against chewing herbivores, but a new kind of leaf damage appeared, namely fungi spots. Data also support that ants may be acting as vectors of fungi spores on plants, as ant visited branches had higher fungus incidence than non-visited branches. Fire is a major source of disturbance in tropical savannas, and we suggest that it can cause strong variation in the outcomes of interactions between ants and plants with extrafloral nectaries in the Brazilian tropical savanna. Keywords: arthropods, cerrado, fungus, mutualism, Qualea multiflora Correspondence: a [email protected], b [email protected], *Corresponding author Editor: Nadir Erbilgin was editor of this paper. Received: 30 August 2012 Accepted: 16 November 2012 Published: 7 July 2014 Copyright: This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 14, Number 84 Cite this paper as: Pires LP, Del-Claro K. 2014. Variation in the outcomes of an ant-plant system: Fire and leaf fungus infection reduce benefits to plants with extrafloral nectaries. Journal of Insect Science 14(84). Available online: http://www.insectscience.org/14.84 Journal of Insect Science | http://www.insectscience.org 1 Journal of Insect Science: Vol. 14 | Article 84 Pires and Del-Claro Introduction Catchpole 1983; Del-Claro and Oliveira 2000). In the latter case, although habitat dis- In ant-plant interactions, plants offer resources turbance, for example fire (Friese et al. 1997), such as shelter (domatia, dead stems) and/or is a cause of reduction in biodiversity (Bond food (exudates, food bodies) to ants, who in et al. 2005), environmental disturbances influ- return may protect the plants against herbi- encing the outcome of plant-ant interactions vores, which they may prey upon or drive have yet to be studied. away (Rico-Gray and Oliveira 2007). Extra- floral nectar (EFN) is the most common Of the world’s biomes, ant-plant interactions nutritional reward that plants offer to ants are most frequent in tropical habitats, occur- (Koptur 1992), increasing survivorship, ring in the full range of habitats where trees growth and reproduction of associated ant are found, from savannah to dense forest. colonies (Byk and Del-Claro 2011). EFN- These interactions are particularly pervasive producing glands have been present in Angio- in cerrado (Brazilian tropical savannah) due to sperms since at least the Oligocene the high incidence of insect- and plant-derived (Pemberton 1992), attracting ants that eat, mo- exudates on foliage, which promote intense lest, or drive away herbivores, thus reducing ant activity on vegetation (Oliveira and Freitas leaf area loss (Rutter and Rausher 2004) 2004). The cerrado biome of South America and/or increasing fruit (Nascimento and Del- covers about 2 million km2, representing ca. Claro 2010) and seed production (Vesprini et 22% of the Brazilian land surface, and is the al. 2003), with a positive impact on seed via- most diverse tropical savanna in the world bility (Sobrinho et al. 2002). (Oliveira and Marquis 2002). Despite its great importance in area and diversity, few studies However, the effectiveness of ant services to on interspecific associations have been con- plants may vary from positive to negative. For ducted in cerrados (Oliveira and Marquis example, Nahas et al. (2012) showed com- 2002; Del-Claro and Torezan-Silingardi plementary effects of multiple predators 2009). resulting in benefits to a plant possessing EFNs. Some authors pointed out a few ant In 1996, researchers performed an ant- partners as parasites (Ness et al. 2006; Byk exclusion experiment in a preserved cerrado and Del-Claro 2010), and others failed to de- reserve protected from fire occurrence, having tect any benefit of the presence of ants to as a model the tree Qualea multiflora Mart. plants (Rashbrook et al. 1992; Mody and (Myrtales: Vochysiaceae), which has extraflo- Lisenmair 2004). Lack of ant protection can ral nectaries. The results showed that ants be explained by (1) differences in ant-foraging provide benefits to the plants, significantly behavior among habitats (Inouye and Taylor reducing leaf area loss and increasing fruit set 1979), (2) variable susceptibility of distinct production (Del-Claro et al. 1996). By the end herbivore groups to ant predation (Ito and Hi- of the dry season of 1999 (October) up to gashi 1991), (3) differences in the deterring 2003, that cerrado reserve suffered incidental skills among species of visiting ants (Dejean fire that completely burned the area in the dry et al. 2000; Byk and Del-Claro 2010; Nahas et season for three consecutive years (K. Del- al. 2012) or (4) difficulties in assessing sea- Claro, personal observations). Because fire is sonal variation and long-term outcomes in the such an important environmental factor in sa- ant-plant-herbivore interactions (O’Dowd and vannah vegetation (Bond et al. 2005), it may Journal of Insect Science | http://www.insectscience.org 2 Journal of Insect Science: Vol. 14 | Article 84 Pires and Del-Claro influence the nature of tritrophic interactions plant. By the flip of a coin, two of these between ants, herbivores, and plants with branches were designated as control and two EFNs. Thus, our current study repeated the as treatment. In control groups, branches did experiments of Del-Claro et al. (1996 in the not receive any manipulation and ants had free same study site and with the same system af- access to them. In treatment groups, ants were ter measures to stop fires were taken and the all manually removed from the brances, and vegetation clearly recovered. The main aim of the intersection of the branch and the trunk this study was to test the hypothesis that envi- was covered with a 3 cm large adhesive paper ronmental disturbance, in this case fire, may strap and a layer of sticky resin was applied produce significant differences in the out- over it (Tree Tanglefoot®) to prevent ants comes (reduction of leaf area loss and fruit set from climbing. All structures that could be production) of an ant-plant association. used by ants as bridges to get access to these experimental branches were removed, and the Materials and Methods integrity of the sticky barrier was checked weekly. To ensure that the sticky resin did not Field work was conducted from September interfere with the results in control brances, 2008 to March 2009 in the cerrado sensu the branch and trunk intersection was covered stricto (Oliveira-Filho and Ratter 2002) eco- with a paper strap, and the resin was applied logical reserve (400 ha) of Clube de Caça e only on one side of the branch, allowing ants Pesca Itororó de Uberlândia (CCPIU) – MG, to climb freely. Brazil (18°59’S; 48°18’W). Fishing and hunt- ing activities in the area have been prohibited In order to sample EFN-visiting ants and plant since 1984. Experiments took place at the herbivores, branchs of both groups were same area in CCPIU where Del-Claro et al. checked fortnightly regarding number and (1996) established their field experiments. species (or morpho species) of observed indi- The regional climate is markedly seasonal viduals.
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