Ant-Partners Play a Minor Role on Occurrence of the Myrmecophilous Butterfly Leptotes Cassius in Its Host Plant

Arthropod-Plant Interactions (2018) 12:377–384 https://doi.org/10.1007/s11829-017-9586-5

ORIGINAL PAPER

Ant-partners play a minor role on occurrence of the myrmecophilous butterfly Leptotes cassius in its host plant

Alexandra Bächtold1,2 · Kleber Del‑Claro1

Received: 17 August 2016 / Accepted: 13 November 2017 / Published online: 20 November 2017 © Springer Science+Business Media B.V., part of Springer Nature 2017

Abstract Ant-related oviposition in facultatively myrmecophilous lycaenid butterflies is common, but not universal. In fact, our knowledge of ant-related oviposition in lycaenids is based on some common species (e.g., Rekoa marius, Allosmaitia strophius, Parrhasius polibetes), which limits generalizations about these systems. In this study, we experimentally investigated whether the oviposition pattern of the florivorous lycaenidLeptotes cassius was influenced by the presence ofCamponotus ants and whether larvae were attended, rather than attacked, by ants. This might be evidence of myrmecophily. Both L. cassius and Camponotus ants occur on Bionia coriacea, an extrafloral nectaried legume shrub that grows in the Brazilian cerrado. Plants were randomly assigned to ant-present and ant-excluded treatments and were observed twice throughout the short reproductive season. Larvae of L. cassius were tended by ants, whose attendance was characterized by active antennation on the last body segments of the caterpillars. Therefore, Camponotus can be considered a partner of L. cassius. Lycaenid abundance was on average 1.9- and 1.21-fold higher in plants with ants in each sampling period, respectively, indicating a tendency of L. cassius to occur in plants with ants. Nonetheless, results were not statistically significant, suggesting that in this case ants are not a major cue for lycaenid oviposition. In many ant–lycaenid mutualisms, butterfly immatures benefit from reduced parasitism rates. However, no L. cassius immature, regardless of ant presence or absence, was parasitized. Furthermore, larvae may occur inside flower buds that may provide protection from natural enemies; thus, ants may not be required for immature protection.

Keywords Brazilian savanna · Camponotus · Extrafloral nectaries · Myrmecophily · Polyommatinae

Introduction perforated cupola organs—Baylis and Kitching 1988; Fie- dler 1991; Axén et al. 1996). Many larvae of Lycaenidae (Lepidoptera) associate with ants Some lycaenids may exhibit strong associations with ants (i.e., myrmecophily—Pierce et al. 2002). Both facultative so that females use ants as a cue for oviposition (Pierce and and obligate associations are possible because larvae pos- Elgar 1985; Fiedler and Maschwitz 1989), and this occurs sess so-called ant-organs that are responsible for the attrac- even in facultatively myrmecophilous species (Wagner and tion and maintenance of otherwise aggressive ants. This is Kurina 1997; Kaminski et al. 2010). Larvae of facultatively achieved by releasing sugared food resources or appeas- myrmecophilous species may have advantages in using ing volatiles (e.g., dorsal nectar organs, tentacle organs, plants visited by ants compared to non-myrmecophilous larvae, especially in extrafloral nectaried species (Fiedler 2001; Kaminski and Freitas 2010). The intense patrolling activity Handling Editor: Jouni Sorvari. of ants upon and in the vicinity of myrmecophilous larvae creates both an enemy- and a competitor-free space, improv- * Alexandra Bächtold [email protected] ing the survival of caterpillars (Kaminski et al. 2010). In this context, it is expected that host plant selection by facultative 1 Instituto de Biologia, Universidade Federal de Uberlândia, myrmecophilous lycaenids may be related to ant presence, Rua Ceará, s/n. Bloco 2D‑Campus Umuarama, Uberlândia, and indeed, this occurs (Pierce et al. 2002). Nonetheless, Minas Gerais CEP 38400902, Brazil there is a remarkable variation in the interactions between 2 Universidade de São Paulo, Avenida Bandeirantes 3900, facultative lycaenids and ants, and ant-related oviposition is Ribeirão Preto, São Paulo CEP 14040901, Brazil

Vol.:(0123456789)1 3 378 A. Bächtold, K. Del‑Claro not universal (Schmidt and Rice 2002; Trager et al. 2013). Materials and methods This variation in the interactions occurs because lycaenid females may also use other factors, like food quality, when Study area selecting the most appropriate plants to oviposit, in order to maximize the survival rate of caterpillars (Rodrigues et al. Fieldwork was carried out in a Brazilian savanna (230 ha, 2010). 18°59′ S, 48°18′ W), in a private natural reserve in Uber- In the Neotropics, studies on ant–lycaenid interactions lândia city, southeastern Brazil, from March to April 2013, have been conducted with the most common species of (fac- which corresponded to the peak flowering period of B. ultative myrmecophilous) butterflies such as Rekoa marius coriacea. The vegetation in this area consists of a cerrado (Lucas, 1857), Allosmaitia strophius (Godart, 1824), and sensu stricto characterized by the presence of shrubs and Parrhasius polibetes (Stoll, 1781), whose oviposition behav- trees ranging between 2 and 4 m tall. The climate has two ior is related to ant presence on plants (Kaminski et al. 2010; well-defined periods: a dry winter (May to September) and Bächtold et al. 2014). Considering the richness of neotropi- a rainy summer (October–April). This latter concentrates cal lycaenids (~ 1200 species; Robbins 2004), there is still more than 90% of the annual average rainfall (Laboratory of a huge gap in the knowledge of the less-studied species, Climatology, Federal University of Uberlandia). especially concerning ant associations. Investigations on some overlooked lycaenid species might provide a further Study system advance in the knowledge of ant–plant interactions and the factors responsible for the occurrence of caterpillars in one The legume shrub B. coriacea (previously known as Camp- plant and not in others. tosema coriaceum) is an herbaceous shrub (up to 1.5 m in Neotropical savannas such as the Cerrado (Brazilian height) common in the cerrado vegetation. In the study area, savanna) are good models for the study of interactions B. coriacea occurs mostly on the edges of the reserve, but it between lycaenids and ants, because both insect groups are is also found in areas in the process of secondary succession rich, diverse, and quite easy to sample (Oliveira and Freitas (after fire or mowing). Leaves are dark green, petiolate, and 2004; Silva et al. 2011). In addition, the presence of extraflo- glabrous. Flower buds (~ 1.5 cm long) are pyriform with nar- ral nectaried plants and their associated ants provides further row acuminate apex and are densely covered with trichomes; opportunities to verify the role of ants in the occurrence of both flower buds and flowers are red. Flowers (~ 3 cm long) lycaenids (Kaminski et al. 2012). are tubular and occur in pseudoraceme inflorescences (Quei- The extrafloral nectaried Bionia coriacea (Nees & Mart.) roz 2008) (Fig. 1a). The extrafloral nectaries ofB. coriacea Benth. (Fabaceae) is a host plant for the florivorous lycaenid occur near buds/flowers (in Fig. 1a, b) and are responsible Leptotes cassius (Crammer, 1775) (Polyommatinae), com- for the attraction and visitation of several ant species (e.g., monly tended by Camponotus Mayr, 1861 (Formicinae) Camponotus, Ectatomma, Cephalotes, Pachycondyla, and (pers. obs.). Leptotes cassius is a facultatively myrmeco- Crematogaster), of which Camponotus are by far the most philous species with little information about its behavior and frequent (Bächtold 2014). Ants in this genus (e.g., C. cras- ecological interactions (Downey and Allyn 1979). This spe- sus, C. blandus, C. rufipes) usually establish mutualistic cies is frequently sampled in inventories of adults in Brazil associations with extrafloral nectaried plants (Korndörfer (Motta 2002; Emery et al. 2006; Sackis and Morais 2008). In and Del-Caro 2006; Nascimento and Del-Claro 2010) and contrast, the ecological data based on immatures are recent, are known for their aggressive behavior against herbivores indicating the population shows marked interannual varia- and some predatory arthropods (Oliveira 1997; Lange and tion, with periods of outbreaks intermingled with surprising Del-Claro 2014; Del-Claro et al. 2016). Camponotus ants absence in field samplings (Bächtold2014 ). also engage in mutualistic relationships with myrmecophil- In our study, we investigated whether the occurrence of ous insects (Fagundes et al. 2013; Bächtold and Del-Claro L. cassius was related to ant presence in B. coriacea (e.g., 2013), including lycaenid larvae (Robbins 1991; Kaminski Bächtold et al. 2014). Specifically, we examined (i) whether et al. 2012; Trager et al. 2013). Larvae of L. cassius are L. cassius larvae establish associations with Camponotus common in B. coriacea and feed on flower buds and flowers ants and (ii) whether adult butterflies use ant presence as (pers. obs. 2011–2014). oviposition cues. Because partnerships between Campono- tus ants and lycaenid species are common (Fiedler 2006; Ant‑related oviposition of lycaenids Alves-Silva et al. 2013), we expected that (ia) L. cassius larvae would be attended rather than attacked by ants; (iia) plants occupied by ants would possess a significantly higher In order to evaluate whether the oviposition pattern of L. cassius number of L. cassius eggs compared to plants without ants. females was based on the presence of ants, we selected and tagged 62 B. coriacea individuals spread over

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Fig. 1 a Inflorescence of Bionia coriacea; the location of extrafloral nectaries is indicated by arrows. b Hatched egg of Leptotes cassius (indicated by the arrow below) near an active extrafloral nectary (indicated by the arrow above). c Unhatched egg of Leptotes cassius. Scales: 20 mm (a); 0.5 mm (b); 0.5 mm (c)

an area of approximately 3 ha. Prior to the experiments, all of ant exclusion. We returned to field 5 days later after ant shrubs had similar heights (< 1.5 m), phenological status, exclusion for the first census of lycaenids. The oviposition and were located on the edges of a trail (~ 2 m wide) that pattern of lycaenids (i.e., the occurrence and abundance of crosses the Cerrado reserve (details in Bächtold et al. 2016). immatures per shrub) was evaluated on two occasions (April All plants were shaded by the canopy of tall trees and none 2, 16) and it was restrained by the short plant reproductive received sunlight all day long. This corresponds to the natu- phenology. The reproductive structures of B. coriacea were ral characteristics of B. coriacea in the area. Tagged plants carefully inspected in the search for eggs and young larvae. were consistently visited by Camponotus crassus Mayr, Eggs and larvae were removed from plants during each sam- 1862 and C. blandus (Smith, F., 1858). Both ant species ple period. Eggs were cautiously placed in small (50 ml) present several similar features, such as foraging behavior, transparent plastic pots, which carried an identification num- occurrence in extrafloral nectary plants, aggressive actions ber of the individual plant and status (control or treated). against herbivores, and attendance to myrmecophilous cater- The same procedure was adopted for larvae, which were pillars (Oliveira et al. 1987; Fiedler 1991; Pereira and Trigo maintained in larger plastic pots (250 ml). Both eggs and 2013). Therefore, we did not differentiate between these two larvae were taken to the laboratory where they were reared ant species in B. coriacea, but rather considered the “Cam- to the adult stage (following Bächtold et al. 2013). During ponotus” group to facilitate subsequent analyses. this process, fresh flower buds of B. coriacea, collected from The individuals of B. coriacea (n = 62) were randomly non-studied individual plants, were offered ad libitum to lar- and equally assigned to two groups (control and treated) vae. Pot cleaning was conducted whenever necessary. Adults which correspond to the presence and absence of ants, obtained from these immatures reared in the laboratory were respectively. In order to exclude ants from plants, we applied used to identify and confirm the lycaenid species and then a layer of non-toxic resin (Tanglefoot ®—Grand Rapids, released in the field, at the study area. MI, USA) at the base of treated shrubs. This procedure is commonly used to prevent the access of ants to plants Ant attendance to caterpillars structures (Nahas et al. 2012). To control for the effect of resin, it was also applied to one side of the stems of con- Ant–lycaenid relationships were experimentally examined in trol shrubs, allowing free access for ants to the plant parts. the field. We were specifically interested in whether larvae Ants were excluded before the survey of lycaenids, so all could establish positive associations with Camponotus, that immatures collected were after the experimental design is, whether larvae would be attended, rather than attacked

1 3 380 A. Bächtold, K. Del‑Claro by ants. Tending behavior is defined as active and frequent exclusion experiment), were conducted and compared with antennation by ants on any part of the larva’s body (Hinton a specific ANOVA (analysis of variance) test in R statisti- 1951; Kaminski and Rodrigues 2011). Eight L. cassius lar- cal software. In this test, we aimed to verify whether the vae (n = 4 larvae in 3rd instar and n = 4 larvae in 4th instar) initial abundance of lycaenids (first sampling) was related reared in the laboratory were taken to the field and placed to their abundance at the second sampling. The significance individually in different shrubs of B. coriacea occupied by of the overdispersion was calculated with a Chi-square test Camponotus. We used mature larvae because at these stages that takes into account the residual deviance and degrees of of development, they may be facultatively associated with freedom. several ant species (Downey and Allyn 1979). Each observa- A general linear mixed model (GLMM) was used to tion session of ant–lycaenid interactions lasted 10 min and investigate whether the presence/absence of lycaenids started as soon as the larva was discovered by ants. Each (binary data, intercept) was related to the experimental session was recorded with a digital camera (480 p). Videos design (ant presence/absence), time of sampling (1st and 2nd were then carefully examined to see in detail the functioning weeks, both as fixed effects), and individual plants (random of larval ant-organs, such as the dorsal nectary organs and effect) (following Bächtold et al. 2014). For this GLMM, the eversible tentacle organs (see Malicky 1970 and; Fiedler we used binary data regarding lycaenid presence or absence 1991 for details of these structures). (zero and one values) on each individual plant. A model with Lycaenid feeding behavior was observed both in the field interaction effects was also performed. Statistical procedures and in the laboratory. Observations (approximately 10 h) were performed in R statistical software version 3.2.3. were conducted ad libitum and we registered the behavior In both GLM and GLMM tests, there were no statistical of larvae and which structures they fed on. differences between the models with and without interaction effects (P > 0.05 in both cases); we therefore chose to use the model with interaction effects. Statistical analyses

Quantitative data are presented as mean ± standard error. Our main analyses are separated into two major tests, as we Results combined data of (1) abundance and (2) presence/absence of lycaenids in plants to investigate their relation with ants. Lycaenids on plants with and without ants The frequency of control and treated individual plants with lycaenids was compared through Fisher’s exact test. In this A total of 80 immatures (n = 76 eggs and 4 larvae) of L. analysis, we used data from the two sampling periods (April cassius (cumulative abundance = 1.29 ± 0.18 immatures per 2, 16). plant, n = 62 individual plants) were recorded on B. coria- A generalized linear model test (GLM) with negative cea (see a L. cassius egg in Fig. 1c). No eggs or larva was binomial errors (following Anjos et al. 2016) was used to parasitized. Lycaenids were found on 61.2% (19 plants) and investigate whether the final abundance of L. cassius (second 64.5% (20 plants) of the treated (ant-excluded) and control sampling period) was related to the presence/absence of ants (ant-present) plants, respectively, and this difference was not and the initial numbers of lycaenids (first sampling period). significant (Fisher’s exact test, P = 0.7998). In our first and Among the GLM tests with different errors (see Crawley second samplings, L. cassius were on average 1.9- and 1.21- 2007), the negative binomial accounted for no overdisper- fold higher, respectively, on plants with ants, showing that sion, so it was given priority. Two GLM models, one with throughout the study, lycaenids showed a tendency to occur and one without interaction effects (initial lycaenids + ant on plants with ants (Table 1).

Table 1 Descriptive data of the Descriptive data Lycaenid abundance Total - cumulative abundance of Leptotes cassius in Bionia coriacea according 1st sampling 2nd sampling to the presence (control) or absence (treated) of ants Control Treated Control Treated Control Treated Mean 0.61 0.32 0.90 0.74 1.52 1.06 SD 0.88 0.54 1.13 1.06 1.59 1.18 Median 0 0 0 0 1 1 Range 0–3 0–2 0–4 0–3 0–5 0–4 Sum 19 10 28 23 47 33

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Table 2 Relationship among the final abundance of Leptotes cassius Lycaenid feeding behavior (intercept), its initial abundance, and the ant exclusion experiment Variables Estimate Std. error z value P value Among the immatures of L. cassius collected in the field, some larvae (n = 5, all from second instar) were observed Lycaenids final (intercept) − 0.3001 0.3130 − 0.9587 0.3377 to chew a small portion of the external part of flower buds Ant exclusion experiment 0.0239 0.4352 0.0550 0.9561 and make a hole (one hole per bud). Larvae then managed Lycaenids initial 0.2721 0.2609 1.0429 0.2970 to enter the bud through this aperture and remained inside Interaction effect − 0.3433 0.5617 − 0.6111 0.5411 the bud while feeding on the reproductive structures, e.g., The interaction effect between ant exclusion experiment and lycaenid the stamens, pollen, and stigma. This behavior was noted initial abundance was also tested. Residual deviance = 62, degrees of in both control and treated plants. Detailed observations in freedom = 58, dispersion parameter = 1 the laboratory also revealed that besides feeding on flower buds and flowers, larvae also fed on extrafloral nectaries. This occurred whenever fresh inflorescences were offered Table 3 Influence of ant presence/absence and time of sampling (fixed factors) on the occurrence of lycaenids (binary data) in Bionia to larvae. coriacea Besides L. cassius, four other lycaenid species were also recorded during fieldwork in B. coriacea, albeit in low abun- Coefficients Estimate SE Z value P value dance: Calycopis calor (H. H. Druce, 1907) (n = 7 eggs), Lycaenids (intercept) − 0.3854 0.4277 − 0.9012 0.3675 Panthiades hebraeus (Hewitson, 1867) (n = 5 eggs), Strymon Ant exclusion experiment − 0.1551 0.6017 − 0.2577 0.7967 mulucha (Hewitson 1867) (n = 2 eggs), and Tmolus venustus Time of sampling 0.4629 0.5608 0.8253 0.4092 (Druce 1907) (n = 1 egg). Interaction effect − 0.3053 0.7910 − 0.3860 0.6995

Ant exclusion experiment and time of sampling were considered as the interaction effect. SE = standard error Discussion

Our study revealed that L. cassius larvae establish positive The generalized linear model with negative binomial associations with Camponotus, because (i) no larvae were errors revealed that the final number of lycaenids was nei- attacked or preyed upon by ants; (ii) ants frequently walked ther significantly related to the experiment (ant presence/ upon larvae and antennated the last segments of the larva’s absence) nor to the interaction effect between experiment body; and (iii) larvae everted the tentacle organs in response and lycaenid initial abundance (Table 2). The GLM model to the antennation by ants. These instances are listed as evi- with and without interaction effects were not statistically dence for the mutualistic interaction between myrmecophil- significantly different 2(χ = 1.1050, P = 0.5754) so we pro- ous larvae and ants (Axén et al. 1996; Saarinen and Daniels vide the former. The overdispersion was not significant 2006; Kaminski and Rodrigues 2011). The tentacle organs (P = 0.3247). play a marked role in lycaenid–ant associations because The general linear mixed test (Table 3) indicated that they are presumed to release volatiles that influence the lycaenid occurrence/absence was also neither affected by alert behavior of tending ants, which maintains them close plants with ants (fixed factor) nor by time of sampling to myrmecophilous larvae (DeVries 1988; Fiedler 1991). (random factor). This latter was responsible for low vari- So far, several ant species were observed in associa- ations in lycaenids (0.7253). The models with and with- tion with L. cassius (e.g., Pheidole anastasii Emery, 1896; out interaction effects were not statistically different Nylanderia bourbonica (Forel, 1886); Brachymyrmex heeri (χ2 = 0.1489, P = 0.6996). Forel, 1874, and Crematogaster ashmeadi Mayr, 1886), and most of these are tiny (~ 1 mm in length) opportunistic species that make use of different food sources such as Tending behavior of ants aphids and extrafloral nectaries (Downey and Allyn1979 ). Camponotus is also an opportunistic ant species that has a Observation in the field revealed that all larvae of L. cas- broad diet as collecting seeds and fruits and hunting for live sius (n = 8) were tended by Camponotus ants, which fre- preys (Yamamoto and Del-Claro 2008). It also feeds on plant quently antennated the larvae in intervals ranging from 5 extrafloral nectar, insect exudates, and maintains stable asso- to 10 s. Then ants moved over the body of larvae before ciations with lycaenid larvae, especially in the neotropics returning to the tending behavior. In contact with ants, (Moreira and Del-Claro 2005; Kaminski et al. 2012; Bäch- two larvae exhibited the eversible tentacle organs (TOs). told and Del-Claro 2013; Lange et al. 2013; Alves-Silva and Secretions delivered from the dorsal nectary organ (DNO) Del-Claro 2014). In fact, lycaenid oviposition based on the were not observed. presence of Camponotus is common (Kaminski et al. 2010;

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Alves-Silva et al. 2013; Bächtold et al. 2014). Our study The knowledge of components in the environment that showed that plants with Camponotus ants had a higher abun- shape lycaenid oviposition behavior is only partly known, dance of L. cassius immatures compared to ant-excluded but is mostly based on food quality/availability and/or the plants, but this was not statistically significant. presence of certain species of tending ants (Pierce 1985; In the present study, we found no statistical support for Baylis and Pierce 1991). Different ant species might result ant-related oviposition patterns in L. cassius, but these in distinct relationships with lycaenid larvae, ranging from results must be seen with caution. According to data sam- mutualist to agonistic (Seufert and Fiedler 1996). For pled in the field (i.e., number of plants and abundance of instance, Ectatomma (ants > 1 mm, with large and strong lycaenids), the power of our analysis was of 0.33. To obtain mandibles and a sharp sting) exhibit two divergent behav- a power of 0.8, which depicts a better scenario, we would iors, as they can attack/kill lycaenid caterpillars or tend lar- need to sample 3.6 times more plants (actual sample = 64, vae for long periods (Robbins 1991; Bächtold and Alves- necessary sample = 232). Nonetheless, the study plant is Silva 2013). In the case of Camponotus, to the best of our not frequent at the area and the mortality of these shrubs knowledge, there is no record of lycaenid larvae being in pervasive. This occurs not only at study area, but also in attacked/killed by ants in this genus. other localities (pers. obs.). The Cerrado contains a rich, As stated above, tending ants might provide an enemy- rather than an abundant flora, thus many investigations on free space for caterpillars (Kaminski et al. 2010), strengthen- plant–lycaenids interactions usually account for samples ing the association between parties. Leptotes cassius spends sizes < 50 plants (see Bächtold et al. 2014). With our cur- at least half of its life cycle (first and second instar larvae) rent sample size and error values, a power 0.8 would also inside flower buds. Living in shelters is a potent anti-par- be possible if the differences in the abundance of lycaenid asitism/predation behavior in butterflies (Fiedler 1991). If were twofold higher in plants with ants, compared to plants we consider that L. cassius benefits from living in shelters, with ants excluded. However, the actual difference was persistent and prolonged association with ants are not nec- 30% (average cumulative abundance in both plant groups). essary to protect larvae from parasitoids. Furthermore, the Study conditions and experimental designs are bounded by production of honey-like solutions and volatiles from lycae- the restrictions of plants and animals in nature. In our case, nids’ ant-organs requires more energy from larvae, which we had a rather good plant sample size and lycaenids were can delay their development and affect adult size (Pierce indeed more abundant in ant-present plants, but statistical et al. 1987; Robbins 1991). Thus, living in shelters might tests did not detect a significant different. enhance larvae development, but this hypothesis requires In general, ant-related oviposition is advantageous for further studies which should also take into account whether both obligate and facultative myrmecophilous lycaenids, as other ant species and the quality of plants also influence the tended larvae can exhibit low levels of parasitism (Pierce oviposition pattern of L. cassius. et al. 1987; Wagner and Kurina 1997; Weeks 2003; Kamin- ski et al. 2010, but see; Bächtold et al. 2014). Nonetheless, Acknowledgements The authors thank the staff of Clube de Caça e Pesca Itororó de Uberlândia where the study was carried out, Pietro lycaenid females may be driven by factors other than ant K.M. Mendonça for information about the host plant, Estevão Alves presence (Nowicki et al. 2005; Rodrigues et al. 2010; Mota Silva for comments and suggestions to for manuscript, and Conselho and Oliveira 2016), so the presence of ants may not be the Nacional de Pesquisa e Desenvolvimento Tecnológico (CNPq) and main cue for oviposition in facultative myrmecophilous Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) for their financial support. lycaenids. For instance, females of Cyclargus thomasi bet- hunebakeri (W. Comstock and Huntington, 1943) show no Compliance with ethical standards preference in laying their eggs on plants occupied by tending ants, even though their larvae may benefit from such Conflict of interest The authors declare that they have no conflict of association (Trager et al. 2013). In other cases, trade-offs interest. between ant occurrence and food quality may also play a role in female oviposition choices (Rodrigues et al. 2010). On the one hand, ant presence may be advantageous for larval development and survival (Fraser et al. 2001; Weeks References 2003); on the other, choosing plants with ants can result in undesirable consequences. In some systems, parasitoids Alves-Silva E, Del-Claro K (2014) Fire triggers the activity of extra- may use ants as cues to locate their hosts (Pierce et al. 1987; floral nectaries, but ants fail to protect the plant against herbivores Fiedler et al. 1992), and high parasitism rates are noted in in a neotropical savanna. Arthropod Plant Interact 8:233–240. https://doi.org/10.1007/s11829-014-9301-8 lycaenid eggs associated with ants (Bächtold et al. 2014). In Alves-Silva E, Bächtold A, Barônio GJ, Del-Claro K (2013) Influ- the case of L. cassius, no immature was parasitized, regard- ence of Camponotus blandus (Formicinae) and flower buds on the less of ant presence or absence. occurrence of Parrhasius polibetes (Lepidoptera: Lycaenidae) in

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