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Effect of Pleometrosis and Brood Transplantation on Colony Growth of the Black Garden Ant, Lasius Niger

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Effect of Pleometrosis and Brood Transplantation on Colony Growth of the Black Garden Ant, Lasius Niger ASIAN MYRMECOLOGY Volume 9, e009003, 2017 ISSN 1985-1944 | eISSN: 2462-2362 © Natalia E. L. Madsen and Joachim Offenberg DOI: 10.20362/am.009003 Effect of pleometrosis and brood transplantation on colony growth of the black garden ant, Lasius niger Natalia E. L. Madsen* and Joachim Offenberg Department of Bioscience, Aarhus University, Vejlsøvej 25, DK-8600, Silkeborg, Denmark *Corresponding author: [email protected] ABSTRACT. In response to intra- and interspecific competition, ant colonies have evolved mechanisms to maximize early colony growth. Through the analysis of photographic images, this study describes the individual effect of two different strategies, pleometrosis and pupae transplantation, on the growth of Lasius niger colonies. Results showed that both methods had an overall positive effect on colony production, albeit the magnitude at which pupae transplantation boosted colony growth was much larger than pleometrosis. After six weeks, colonies with two and three queens contained, respectively, 26% and 83% more brood than control colonies with only one queen. Queen association also led to overall decreased queen fecundity and death of all resident queens in over 60% of the colonies due to queen fights. After eight weeks, colonies that received 30 or 60 foreign pupae had produced on average 256% more brood than controls and nanitic workers that were 7% longer than those from colonies that did not receive transplanted pupae. Keywords: pupae adoption, multiple queens, ant reproduction, colony found- ing, social insects, formicidae INTRODUCTION queens have a limited amount of resources to invest in reproduction, to produce more workers Most colonies of Lasius niger are started by a than single queens and to do so in shorter time single claustral queen that uses her stored energy (Waloff 1957, Bartz & Hölldobler 1982, Sommer reserves to raise her first brood. However, the & Hölldobler 1995). That may translate into (i) a high numbers of Lasius queens occurring after higher foraging success, which will improve ear- mating flights, coupled with the fact that queens ly colony growth and survival and (ii) an advan- avoid areas frequented by workers of established tage when it comes to successfully defending the colonies (Sommer & Hölldobler 1995), leads to a nest against brood raiding, usurpation (Bartz & high density of new nest foundations in the field. Hölldobler 1982, Rissing & Pollock 1991, Tsch- In response to crowding and high inter-colony inkel 1992b, Bales & Adams 1997) and predation competition, incipient colonies may use two dif- (Jerome et al. 1998) by neighbouring colonies. ferent strategies to gain competitive advantage. That said, an alternative reason for pleometrosis One strategy is pleometrosis, where queens enter can also be a strong pressure for the newly mated facultative associations with non-related queens queens to leave the soil surface and use any avail- during colony founding (Bernasconi & Stras- able holes, including those excavated by other smann 1999). Such pleometrotic association al- queens (Tschinkel 1998). Despite the potential lows claustral species like L. niger, in which benefits of pleometrosis, there is also an impor- 2 Natalia E. L. Madsen & Joachim Offenberg tant cost at the queen level, as the emergence of viduals necessary for the multiple queen experi- workers and the start of foraging activities gener- ment (pleometrosis) were placed in groups of 1, ally trigger the onset of queen fights that continue 2 or 3 queens per test tube less than 60 minutes until only one of the queens survives (Sommer after collection to minimize a build-up of territo- & Hölldobler 1995, Bernasconi & Keller 1998, rial behaviour over time. The day after collection, Aron et al. 2009). queens were cooled to 16.5°C for 60 minutes for A second strategy to improve competi- easier manipulation before being transferred into tion by increasing early colony growth is the se- artificial nests. questration of foreign pupae and larvae through The artificial nests were created in Petri the raiding of brood from neighbouring colo- dishes (5.5cm Ø). The lids of the dishes were per- nies. This strategy provides ant colonies in the forated (single hole, 4mm Ø) to allow the move- field with a competitive advantage derived from ment of workers in and out of the nest; the bases a rapid increase in the number of individuals in were partly filled (approximately 2/3 of their vol- the nest, which leads to earlier colony matura- ume) with a substrate consisting of a combina- tion (Pollock & Rissing 1989, Rissing & Pollock tion of Plaster of Paris and charcoal (8:1). A small 1991, Tschinkel 1992a, Gadau et al. 2003). piece of tape on the substrate allowed the provi- In addition to building up higher num- sion of a drop of water for the queens to drink bers of workers, it may also be advantageous for until the first workers emerged, at which point young colonies to produce larger individuals, as workers provided the queen with water. The nests size influences the range of tasks a worker can were then placed in a bigger Petri dish (14cm Ø) perform (Wilson 1978, Mirenda & Vinson 1981, that contained a 10cm test tube filled with water Arnan et al. 2011). For example, mature colo- and also served as an arena where the feedings nies in the field have not only a higher number would take place. Ad libitum sources of carbo- of workers, but also larger individuals compared hydrate (20% sucrose solution in water) and pro- to incipient colonies (Porter & Tschinkel 1985, tein were added when the first workers emerged. Hasegawa & Imai 2011), and worker polymor- Protein consisted of house flies, pieces of shrimp phism and task repertoire have been found to and a 1% solution of peptone from bovine meat correlate with colony size (Anderson & McShea (Sigma-Aldrich, product number 70175), alter- 2001). Whether increased colony growth via nated weekly. All arenas and nests were kept in- pleometrosis or sequestration of foreign pupae side opaque paper boxes in a dark 25°C chamber leads to production of larger workers, and in this for the duration of the experiment. Feedings and way further improves the competitive abilities population assessments were performed outside of incipient colonies of Lasius niger, remains to the chamber, in light conditions and at room tem- be tested. perature (August-September, 2014, mean daily In order to elucidate the potential com- temperature approximately 18°C). petitive advantage colonies can achieve through pleometrosis and brood raiding, the present study Colony assessments focused on analysing the effects of multiple During the experiment, non-destructive assess- founding queens and pupae adoption on early ments of colony sizes were based on photographic colony growth and worker size of Lasius niger. images taken every two weeks (camera: Canon EOS 1000D, lens: EF-S 18-55mm f/3.5-5.6 IS). The software ImageJ (Open Source: http://imagej. METHODS nih.gov/ij/) was used to analyse the images in or- der to perform counts of brood and workers and to Experimental colonies measure colony areas and lengths of nanitic work- Queens of Lasius niger were collected immedi- ers. After the last assessment on week eight after ately after their mating flight on 17th July, 2014, in the nuptial flight, the ant colonies were exposed Aarhus, Denmark (56°11’04.9”N 10°07’01.1”E), to gradually decreasing temperatures (5 days at and were kept individually in plastic test tubes 20°C followed by 5 more days at 15°C). Lower containing water behind a cotton plug. The indi- temperatures reduce the activity level of the colo- Boosting colony growth in the garden ant Lasius niger 3 nies, making it possible to perform a more exact Poisson distribution and log-link function, Chi- manual count of the number of pupae and work- square = 526.34, p < 0.0001) than that in the 30 ers that were used to correct the estimations ob- pupae treatment (mean ± SE = 19.93 ± 1.31). The tained through photographic analysis. The number exact number of transplanted pupae hatching into of eggs and larvae, however, could not be counted imago workers was monitored and used for cal- manually in this way due to the delicate nature of culations of survival in relation to the number of these stages and our need for keeping the individu- pupae initially transplanted into the colony; i.e. als alive for future experiments. discarded pupae were included as non-surviving. The number of surviving pupae in each colony Multiple queen experiment was subtracted from the total number individuals The multiple queen experiment started with to obtain intrinsic queen production. 16 single queen colonies, 16 colonies with two queens and 13 colonies with three queens. Dur- Nanitic size ing the experiment, a mite infection reduced the In the analysis of nanitic size, photos were used sample size to a total of 29 usable colonies: 11 to measure length longitudinally from mandibles control colonies with one queen, 8 colonies with to the distal end of gaster using ImageJ software. two queens and 10 colonies with three queens. In Five to eight nanitic workers (much smaller and addition, queen fights resulted in 12 colonies be- lighter in colour than adopted ones) were selected coming orphan, as all the queens died from their per colony, depending on availability, and their injuries, which led to a stop in brood production. lengths averaged to provide a single value per ex- As a result, the set of data (images) analysed in perimental colony. this experiment belonged to week six of the ex- periment, where all queens were still present in Statistical analysis all colonies and queen fights had not yet started. Seven response variables were analysed in the two experiments. Four of them were readily ob- Pupae transplantation experiment tained through analysis of photographic images.
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