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Secretions of Mandibular Glands Are Not Involved in the Elicitation of Rescue Behaviour in Formica Cinerea Ants

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Secretions of Mandibular Glands Are Not Involved in the Elicitation of Rescue Behaviour in Formica Cinerea Ants Insect. Soc. DOI 10.1007/s00040-017-0547-x Insectes Sociaux SHORT COMMUNICATION Secretions of mandibular glands are not involved in the elicitation of rescue behaviour in Formica cinerea ants K. Miler1 · K. Kuszewska1 Received: 10 November 2016 / Revised: 4 January 2017 / Accepted: 20 January 2017 © The Author(s) 2017. This article is published with open access at Springerlink.com Abstract Certain ants perform rescue behaviour for other Chemical communication among nest mates plays a cru- ants that require help, and the expression of rescue behav- cial role in the functioning of an ant colony. For example, iour is hypothesized to depend on signals (“calls for help”) the group predation of ponerine ants occurs via recruit- sent by the imperilled individuals. We studied whether the ment behaviour based on pheromones secreted by scout- mandibular glands were involved in the elicitation of rescue ing individuals (Maschwitz and Schönegge 1977), and the behaviour in Formica cinerea Mayr ants. In the first experi- queens of Pharaoh ants produce pheromones that enable ment, we determined the occurrence and characteristics of their recognition by workers and special functions in the rescue behaviour directed towards nest mates with impaired colony (Edwards and Chambers 1984). Pheromones in ants mandibular gland communication. We did not observe any are likely involved in every aspect of their lives and ensure difference in rescue behaviour directed towards individu- colony integrity (Jackson and Morgan 1993). One of the als who were untreated, treated with paint over the man- best examples of the complex social behaviour controlled dibles, or sham-treated with paint over the thorax. In the by pheromones is provided by certain sand-dwelling ants, second experiment, we determined whether rescue behav- including the species in this study, Formica cinerea Mayr, iour would occur towards dummy ants coated with the con- showing rescue behaviour towards nest mates that require tents of the mandibular glands. Compared with the control help (Czechowski et al. 2002). Indeed, the elicitation of res- untreated nest mates, we found that rescue behaviour was cue behaviour, specifically in Formica, is hypothesized to not directed towards either the untreated dummy ants or depend primarily on pheromonal signals (“calls for help”) the dummy ants covered with crushed mandibular glands. sent by the imperilled individuals (Czechowski et al. 2002). Our results indicated that the “call for help” signal does not Thus, we studied whether mandibular glands were involved originate from the mandibular glands. Therefore, we pro- in the expression of rescue behaviour in F. cinerea ants. pose that gaster-tip glands represent a plausible alternative These glands are the most likely candidates for the source source of rescue-eliciting pheromone(s) for the F. cinerea of rescue-eliciting pheromone(s) because of their involve- ants examined in this study. ment in the related functions, e.g., coordinating, alerting, and attracting (Attygalle and Morgan 1984; Ali and Mor- Keywords Mandibular gland · Pheromone signalling · gan 1990). To demonstrate the potential importance of Reduced communication · Rescue behaviour secretions from mandibular glands, we designed two exper- iments in which mandible-based pheromone communica- tion was blocked between nest mates or the contents of the Electronic supplementary material The online version of this mandibular glands was used to provoke the expression of article (doi:10.1007/s00040-017-0547-x) contains supplementary material, which is available to authorized users. rescue behaviour. Our methods were similar to those used in previous studies (e.g., Hölldobler et al. 2013; Stuttard * K. Miler et al. 2016). [email protected] In the first experiment, the ants were tested in dyadic 1 Institute of Environmental Sciences, Jagiellonian University, encounters of individuals from the same colonies. In Gronostajowa 7, 30-387 Kraków, Poland each test, one ant required help, as it was entrapped on Vol.:(0123456789)1 3 K. Miler, K. Kuszewska the surface of the sand (namely, the entrapment bioassay, behaviour]. However, the effects of pheromone residu- e.g., Nowbahari et al. 2009; 2012), whereas the nest mate als were unlikely, because in the second experiment, the was free. The entrapped ant was either untreated (control contents of the mandibular glands did not elicit rescue group), had a drop of paint applied over the mandibles behaviour towards the dummy ants. All four attempted (group with blocked pheromone communication via man- rescues of the gland-treated group of dummy ants were dibular glands), or had a drop of paint applied over the tho- weak and could have resulted from other substances rax (sham-treated group). In each test, we noted whether transferred onto them during experimental procedures the free ant performed rescue behaviour, the latency to the (Bagnères et al. 1991). Thus, our results indicated that first episode of rescue, and the total duration of rescue. Dig- the mandibular glands are not involved in the elicitation ging around the entrapped nest mate, pulling at its limbs, of rescue behaviour in F. cinerea ants. transporting sand particles away from it, and biting the The previous reports have indicated that the man- snare entrapping the nest mate were evaluated as the main dibular gland secretions of ants function at a minimum subcategories of rescue behaviour. In the second experi- to attract conspecifics (Cammaerts et al. 1981; Howard ment, the ants were tested in an analogous situation, but the et al. 1982) and release both alarm and digging behav- ‘trapped’ ant was either untreated (first control group) or a iours, which are responses involved in rescue operations dummy ant that was either untreated (second control group) (Wilson 1958; McGurk et al. 1966). However, these stud- or covered in the crushed contents of a mandibular gland ies involved ants from genera that were not used in the (experimental group). The same type of data was collected present study (Formica), including Pogonomyrmex, Was- in the second experiment, and the same subcategories of mannia, and Myrmica. In addition, the mandibular glands rescue behaviour were evaluated. We used a two-tailed in Formica workers contain low quantities of volatile Fisher’s exact test (FET) to detect the between-group dif- materials (Bagnères et al. 1991). Therefore, other glands ferences in the rate of occurrence of rescue behaviour and in Formica are most likely involved in rescue elicita- a Kruskal–Wallis ANOVA to detect the between-group dif- tion, such as Dufour glands, which function in commu- ferences in the latency and the duration of the behaviours nication (Löfqvist 1976; Attygalle and Morgan 1984). An (see the Supplementary Information for detailed descrip- alternative or complementary explanation could be that tions of the materials and methods). the production of CO2 by nest mates that require help In the first experiment, we found that rescue behaviour attracts other ants and releases the basic forms of rescue occurred in 16 of 30 tests with the first untreated control behaviour, alarm, and digging behaviours, as observed group of ants, in 12 of 30 tests with the second mandible- in Solenopsis ants (Hangartner 1969). Notably, stridula- treated group of ants, and in 11 of 30 tests with the third tion may be an alternative mode of communicating for thorax-treated group of ants. Based on these results, the fre- help. Indeed, stridulation is hypothesized to have evolved quency of rescue behaviour occurrence among the groups among the ants to alert nest mates that rescue is required, was not significantly different (FET yielded nonsignificant although this hypothesis has been largely rejected results for each comparison). In addition, differences were (Golden and Hill 2016). Moreover, stridulation as a call not observed among these three groups in either the latency for help is not relevant in Formica, because stridulatory to the first episode of rescue (K–W ANOVA: H = 0.123, organs are absent in this genus (Czechowski et al. 2002). p = 0.940) or the total duration of rescue (K–W ANOVA: Thus, the “call for help” in our study species of Formica H = 0.192, p = 0.908). In the second experiment, we found could not have involved vibroacoustic signals. that rescue behaviour occurred in 31 of 60 tests with the The current studies on the selected ecological and evolu- first untreated control group of ants, in none of the 60 tests tionary aspects of rescue behaviour are strongly dependent with the third control group of dummy ants, and in 4 of 60 on the hypothesis that individuals who require help emit tests with the second gland-treated group of dummy ants. “call for help” signals; however, this behaviour remains These results indicated that only the live ants in the first largely unknown (e.g., Nowbahari et al. 2009; Miler 2016), group elicited rescue behaviour (FET yielded a nonsignifi- and uncovering the mechanism of rescue behaviours in ants cant result for the comparison between the second and the is an essential component of further research. Although third groups). mandibular gland secretions did not elicit rescue behav- The data from the first experiment could be con- iours in this study, they can possess such a function in other founded by certain minute residual pheromone(s) on the sand-dwelling ants which display rescue behaviours (Hollis body surface of trapped ants with blocked pheromone and Nowbahari 2013). In F. cinerea ants as well as
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