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Download PDF File Myrmecological News 14 99-106 Vienna, January 2011 Aggressive behaviour of the two European Argentine ant supercolonies (Hymenoptera: Formicidae) towards displaced native ant species of the northeastern Iberian Peninsula Sílvia ABRIL & Crisanto GÓMEZ Abstract The present study attempts to analyse the aggressive behaviour of the two European supercolonies of the Argentine ant, Linepithema humile, in confrontations between them and with several native ant species, to assess if there are differences in their aggressiveness, and consequently, in their competitive ability. Workers from the Main supercolony were more aggressive than workers from the Catalan supercolony. They showed a higher significant aggressiveness index in confrontations with workers from the Catalan supercolony and with two of the three native ant species studied: in Tapinoma nigerrimum (NYLANDER, 1856) and Lasius cinereus SEIFERT, 1992, but not in Pheidole pallidula (NYLANDER, 1849). They also mainly initiated the aggressive encounters and responded ag- gressively to attacks during confrontations both with the Catalan supercolony and with the native ant species. As com- pared to the Main supercolony, workers from the Catalan one showed a reduced aggressive behaviour in confrontations with the native ant species studied. Moreover, the native ant species attacked workers from the Catalan supercolony more frequently than workers from the Main one. These results may support the hypothesis of a weaker competitive ability of the Catalan supercolony, and therefore, a minor invasiveness power, and could also partially explain why that supercolony is clearly less widely distributed throughout Europe than the Main supercolony. Key words: Aggressiveness, competitive ability, Lasius cinereus, Linepithema humile, Pheidole pallidula, Tapinoma nigerrimum. Myrmecol. News 14: 99-106 (online 1 October 2010) ISSN 1994-4136 (print), ISSN 1997-3500 (online) Received 22 May 2010; revision received 1 August 2010; accepted 3 August 2010 Dr. Sílvia Abril (contact author) & Dr. Crisanto Gómez, Department of Environmental Sciences, University of Girona, Montilivi Campus s/n, 17071 Girona, Spain. E-mail: [email protected]; [email protected] Introduction The Argentine ant (Linepithema humile MAYR, 1868), well extreme case of polydomy and polygyny in ants. Unicolo- known as an invasive ant species (MCGLYNN 1999), has nial ants have huge colonies formed by a complex network spread worldwide (HÖLLDOBLER & WILSON 1990, SUAREZ of hundreds or thousands of nests, each with multiple queens & al. 2001) as a result of human commercial activities in (HELANTERÄ & al. 2009). This behaviour gives to the Ar- habitats usually associated with human modification (HOL- gentine ant the ability to create enormous aggregations of WAY 1998a, SUAREZ & al. 2001). However, its ability to nests that are non-aggressive among each other. These large occupy non-altered habitats has also been reported (COLE aggregations are called "supercolonies" and can range over & al. 1992, HOLWAY 1998a, GÓMEZ & al. 2003). Its dis- thousands of kilometers (SUAREZ & al. 1999, TSUTSUI & tribution range includes areas with Mediterranean-type cli- al. 2000, TSUTSUI & CASE 2001, GIRAUD & al. 2002, SU- mates throughout the world (PASSERA 1994, SUAREZ & al. NAMURA & al. 2009, VAN WILGENBURG & al. 2010). 1998) and in the specific case of the Iberian Peninsula it is This unusual social structure allows them to obtain high found over the entire coastal band (WETTERER & al. 2009). densities of workers and numerical dominance over na- In its introduced range, the Argentine ant has impacted tive ant species. It is widely recognised that the European native ant faunas leading to changes in arthropod commu- population of the Argentine ant consists of two super- nities (HUMAN & GORDON 1996, 1997, HOLWAY 1998b, colonies: the Main supercolony, which follows the entire SUAREZ & al. 1998), ant-vertebrate interactions (SUAREZ & coastal band from Italy to the Iberian Peninsula, and the al. 2000) and ant-plant relationships (BOND & SLINGSBY Catalan supercolony, which is smaller than the Main su- 1984, VISSER & al. 1996, GÓMEZ & OLIVERAS 2003, BLAN- percolony and apparently restricted to the eastern part of CAFORT & GÓMEZ 2005). Its presence has also had negative the Iberian Peninsula but mainly present in Catalonia (GI- effects on crops and plantations due to its mutualistic inter- RAUD & al. 2002). There is a total absence of intraspecific actions with hemipterans, which reduce plant growth and aggression between workers from the same supercolony, production (BUCKLEY 1987, NESS & BRONSTEIN 2004). despite the long distance that can separate them (as many As with most invasive species, there are several charac- as 6000 kilometers in the Main supercolony). However, teristics that contribute to their success in their introduced there is high aggression between the two supercolonies (GI- ranges. One of them is unicoloniality (PASSERA 1994), an RAUD & al. 2002). Aggressive behaviour is an important component of eco- ertheless use the name T. nigerrimum because the male logical dominance, because it allows exclusive access to re- genitalia of the taxon we used differ from T. erraticum (cf. sources such as food or nesting sites (ROWLES & O'DOWD SEIFERT 1984; confirmed by comparison with type mate- 2007, CARPINTERO & REYES-LÓPEZ 2008). In the case of rial deposited at the Museu de Ciències Naturals de Ca- the Argentine ant, the combination of its highly aggressive talunya) and because of distinct chromosome numbers in behaviour in comparison with native ants (HÖLLDOBLER & T. nigerrimum and T. erraticum (PALOMEQUE & al. 1988), WILSON 1990, HUMAN & GORDON 1999) and its numerical but we do not propose a formal taxonomic act here. We dominance caused by unicoloniality (HUMAN & GORDON chose these three native ant species because they are suc- 1996, OLIVERAS & al. 2005) make this species very com- cessfully displaced by the Argentine ant in natural invaded petitive, which is likely the key to its high level of inva- areas (OLIVERAS & al. 2005), and therefore can be of use siveness. For this reason, its nestmate recognition system to compare the potential invasiveness of each supercolony has been studied in depth. Intraspecific aggression in this in terms of their ecological dominance. These native ant species may be influenced by several factors. On the one species were sampled from the non-invaded area in Castell hand, it seems that both genetic and environmental cues can d'Aro (CA) (Fig. 1). affect its nestmate-discrimination ability (LIANG & SILVER- The level of aggression in the Argentine ant can vary in MAN 2000, TSUTSUI & al. 2000, CHEN & NONACS 2002, response to the presence of competitors. Repeated encoun- SUAREZ & al. 2002). On the other, the frequency and in- ters with neighbouring individuals may result in a habitu- tensity of aggression between non-nestmates can also be ation process leading to a reduction in the aggression lev- influenced by behavioural factors. THOMAS & al. (2005, els of workers, the so-called "dear-enemy phenomenon" 2006, 2007) reported that workers close to colony bounda- (CHEN & NONACS 2002). Conversely, the Argentine ant can ries exhibit higher levels of aggression than those living also increase its aggressiveness as a consequence of repeated further away, and concluded that the intensity of the intra- exposure to individuals from foreign colonies (THOMAS specific aggression of these ants may change depending on & al. 2007) leading to changes between the intensity of ag- the history of contact and the nature of those interactions. gression of workers from the colony boundaries and that Moreover, it was also reported that the levels of aggression of workers far away from this contact zone. Due to this of this species may, in some cases, change temporally (CHEN plasticity in its recognition system, we collected all the & NONACS 2002, SUAREZ & al. 2002). These studies mainly Argentine ant workers of this study from the invasion cen- focus on the aggressive interactions among Argentine ant tre, far from the contact zones. Conversely, we also collect- workers belonging to different United States supercolo- ed all the native workers from non-invaded areas; therefore, nies, but up to now, not a single study has examined pos- they had had no contact with the invader prior to this study. sible differences in the aggression level of supercolonies Aggressiveness tests: In order to quantify aggressive- towards native ants. It seems that the presence of native ness between the two Argentine ant supercolonies and be- ants in the Argentine ant's advancing front can offer re- tween these supercolonies and the native ant species, we sistance to the invasion progress (HELLER & al. 2006). adapted the protocol employed by GRANGIER & al. (2007). Therefore, the degree of competitiveness exhibited by the To avoid possible changes in the aggression levels of wor- invader against local ants can play an important role in kers through time (CHEN & NONACS 2002, SUAREZ & al. terms of the invasion success of the species. Against this 2002), we carried out all the aggression tests on the same background, the present study aims to analyse the aggres- day we collected them from the field. In the lab, we chose sive behaviour of the two European Argentine ant super- at random one worker from each supercolony or species colonies in confrontations between each other and with sev- and placed them in a neutral arena (diameter = 2.5 cm, eral native ant species
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