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Resumo Expandido Feeding Behavior of Atta Sexdens

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Resumo Expandido Feeding Behavior of Atta Sexdens RESUMOXVIII Simpósio EXPANDIDO de Mirmecologia 031 379A FEEDING BEHAVIOR OF ATTA SEXDENS (HYMENOPTERA: FORMICIDAE) QUEENS DURING ESTABLISHMENT OF NEW COLONIES IN LABORATORY Comportamento alimentar de rainhas de Atta sexdens (Hymenoptera: Formicidae) durante o estabelecimento de novas colônias em condições de laboratório J.O. Augustin* & J.F.S. Lopes Universidade Federal de Juiz de Fora, Instituto de Ciências Biológicas, Campus Universitário Martelos, CEP 36036-330, Juiz de Fora, MG, Brazil. E-mail: [email protected] In ants, colony founding is the most vulnerable the fungus in the mother colony, but also the stage in their life cycle (Hymenoptera, Formicidae) establishment, growth and care of the initial garden (HÖLLDOBLER & WILSON, 1990). In response, a number (HÖLLDOBLER & WILSON, 1990; FERNÁNDEZ-MARÍN et al., of strategies that allowed colonial success were 2003, 2004). selected (PEETERS & ITO, 2001; BROWN & BONHOEFFER, Considering that the establishment of a new colony 2003). Ants were favored with two main reproductive is a key feature to understand the evolutionary strategies. The dependent strategy, in which a adaptations by which many social insects went founding queen depends on their nestmates to start a through (HÖLLDOBLER & WILSON, 1990; PEETERS & ITO, new colony, and the independent strategy, in which 2001; BROWN & BONHOEFFER, 2003; FERNÁNDEZ-MARÍN et the founding queen does not require the help of al., 2003, 2004; JOHNSON, 2006), this study focuses on nestmates (HÖLLDOBLER & WILSON, 1990). Independent the feeding behavior of Atta sexdens queens during founding strategies can be of two types. First, in the both colony founding and early ergonomic stages in semiclaustral founding, queens leave the nest to forage laboratory. for food for themselves and their brood. Second, in the Just after their nuptial flight in early november, 211 fully claustral founding, queens seal themselves in reproductive females of A. sexdens were collected in the nest after mating and never leave the nest to forage. pasture fields within the City of Juiz de Fora (21°46‘S– It is generally assumed that fully claustral founding 43°21‘W, 678m average altitude), southeastern Brazil. queens rely solely on their body reserves to rear the Queens were collected with the aid of tweezers and first generation of workers (KELLER & PASSERA, 1989; individually placed in transparent plastic containers WHEELER, 1994; LIU et al., 2001; HAHN et al., 2004). The (12 cm of diameter and 10,5 cm of height) with a 1 cm definition of claustral colony founding, however, is layer of dampened plaster at their base to keep humidity more precise than that. It has been proposed by inside the chambers. With the aid of needle tip, small HÖLLDOBLER & WILSON (1990) and is given as “the holes on the chamber covers were made in order to procedure during which queens (or royal pairs in the allow the entrance of oxygen in the colonies. Labelled case of termites) seal themselves off in cells and rear containers permitted the identification of the colonies the first generations of workers on nutrients obtained to be observed every week. Laboratory conditions mostly or entirely from their own storage tissues, were kept in 25 ± 5° C, 70 to 80% of relative humidity including fat bodies and histolysed wing muscles”. and photoperiod of 12 hours. The claustral founding in ants is performed by the The Focal Animal Sampling method (ALTMANN, sub-families Dolichoderinae, Formicinae and 1974) was conducted in 30 colonies, starting from the Myrmicinae (HÖLLDOBLER & WILSON, 1990; PEETERS & following day since the collection of the queens. It took ITO, 2001; BROWN & BONHOEFFER, 2003), which are the colonies 11 weeks before the first workers emerged, considered morphologically more complex ants. so that observations were extended for more 11 weeks Particularly more elaborated is the founding behavior after workers emerged in each colony. Six colonies of the fungus-growing ants (Formicidae: Attini) were observed per day, during 30 uninterrupted (HÖLLDOBLER & W ILSON, 1990; WIRTH et al., 2003). Great minutes each. After 22 weeks, colonies had been part of this merit is due to the fact that Attini founding observed for 11 hours each, during both the founding queens must alone accomplish tasks such as both the and the ergonomic stages. care of the offspring and the fungus garden. The later Queens that eventually perished were replaced by includes not only the collection of an initial portion of other ones from the same sample. *Programa de Pós-Graduação em Comportamento e Biologia Animal. Biológico, São Paulo, v.69, suplemento 2, p.379-383, 2007 380 XVIII Simpósio de Mirmecologia Behaviors were recorded as single occurrences using unidirectional movements of the glossa (from regardless of their duration. Data were found to be not within to the outside of the mouth) they ingested the normally distributed according to the Kolmogorov- corporal fluids that come out of the body of the larva. Smirnov modified by Lilliefors test. In order to evaluate In the ergonomic stage, the queens were observed to differences in the absolute frequencies of the receive from a worker’s jaws a portion of the body of behavioural acts both in different phases within the a larva, that they ingested by movements of both the founding stage as well as in between the founding glossa (from within to the outside of the mouth) and and ergonomic stages, all data were analyzed by jaws (in open-and-close movements). means of c2 test. Only once were the queens observed to perform Queens were found to perform five behavioral acts pupophagy (Table 1). Similarly to larvophagy, queen related to the feeding behavior: were observed to receive from a worker’s jaws a portion 1. Ingesting trophic egg of the pupal abdomen, which the queens ingested by 2. Ingesting fungi staphylae movements of both the glossa (from within to the 3. Ingesting reproductive egg outside of the mouth) and jaws (in open-and-close 4. Larvophagy movements). 5. Pupophagy Although it is generally believed that fully The ingestion of trophic eggs was the most claustral queens rear their first clutch of workers frequently recorded behavior (Table 1). Queens were solely on internal reserves (KELLER & PASSERA, 1989; observed to bend their bodies in a dorso-ventral WHEELER, 1994; LIU et al., 2001; H AHN et al., 2004), our direction before a newly-trophic egg was laid. While work shows that this may not always be the case, the egg was still on the anal pore, the queens ingested given the fact that A. sexdens queens were found to it with unidirectional movements of the glossa. Queens ingest both trophic and reproductive eggs as well as also ingested trophic eggs that happened to be on the fungi staphylae. Here we show that the main food mouth of the larvae. sources for the queens were both the trophic eggs and The second most frequently recorded behavior the fungi staphylae. Ingesting extra nutritious was ingesting staphylae (Table 1). The queens were sources would strongly mean that A. sexdens queens observed to grab a portion of the simbiont fungus required more energy for founding new colonies in using their jaws and with unidirectional movements laboratory than their internal reserves could offer. of the glossa (from within to the outside of the mouth) This could largely be because founding attine queens they ingested the fungus. Despite being the most face a challenge that other fully claustral queens do frequently recorded behaviors, no significant not: they must accomplish fungiculture, otherwise differences were found between the ingesting of their colonies will not last for long. Indeed, because trophic eggs and ingesting staphylae in any of the of the obligatory ant-fungus mutualism (CURRIE, 2001), colonial development stages (c2 = 3,42; p = 0,0644). Atta colonies have been found not to last for more Comparing both colonial development stages, than 78 days without the fungus garden (AUTUORI, however, ingesting fungi staphylae (c2 = 60,552; p < 1941, WEBER, 1972; FERNÁNDEZ-MARÍN et al., 2005, 0,001) and ingesting trophic egg (c2 = 54,256; p < AUGUSTIN, 2007). Although claustral queens that 0,001) were more frequent in the founding stage than exhibit the independent mode of colony founding in the ergonomic stage. are capable of accumulating energy reserves to Only twice were the queens observed to ingest support mating, dispersal, and either digging nests reproductive eggs (Table 1). Such behavior was only (KELLER & PASSERA, 1989; JOHNSON, 1996; HAHN et al., observed in the founding stage. Queens were observed 2004) the oophagy behavior is not uncommon among to grab a reproductive egg between her jaws, licking Atta queens (AUTUORI, 1940, 1942; HÖLLDOBLER & it repeated times until it was ingested, by means of WILSON, 1990; AUGUSTIN, 2007). In fact, our study unidirectional movements of the glossa from within shows the ingesting of trophic eggs as the most to the outside of the mouth. The frequency of ingesting frequently recorded behavior, which did not trophic eggs was significantly higher than the significantly differ from the ingesting of staphylae. ingesting of reproductive eggs both in the founding This could mean that both behaviors not only share stage (c2 = 118,127; p <0,001) and the ergonomic stage equivalent costs but also equally benefits for colony (c2 = 32; p <0,001). maintenance. Considering that the claustral Larvophagy was once observed in the founding founding is energetically demanding for newly mated stage and once observed in the ergonomic stage (Table queens that found new colonies independently, this 1). This behavior was observed in two different colonies result strongly indicates that A. sexdens queens do and was performed in two different ways. In the not rely solely on nutrients obtained entirely from founding stage, the queens were observed to perforate their own storage tissues when founding new with their jaws the cuticle of the body a larva and, colonies.
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