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Initiation of Absconding-Swarm Emigration in the Social Wasp Polybia Occidentalis Author(S): Peter J

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Initiation of Absconding-Swarm Emigration in the Social Wasp Polybia Occidentalis Author(S): Peter J Initiation of Absconding-Swarm Emigration in the Social Wasp Polybia occidentalis Author(s): Peter J. Sonnentag and Robert L. Jeanne Source: Journal of Insect Science, 9(11):1-11. 2009. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1673/031.009.1101 URL: http://www.bioone.org/doi/full/10.1673/031.009.1101 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. .SYVREPSJ-RWIGX7GMIRGI:SP`%VXMGPI 7SRRIRXEKIXEP Initiation of absconding-swarm emigration in the social wasp Polybia occidentalis 4IXIV.7SRRIRXEK ERH6SFIVX0.IERRI (ITEVXQIRXSJ)RXSQSPSK]9RMZIVWMX]SJ;MWGSRWMR1EHMWSR;- Abstract When a colony of the swarm-founding social wasp &#" loses its nest to severe weather or predation, the adult population evacuates and temporarily clusters on nearby foliage. Most of the adults remain inactive in the cluster, while foragers bring in nectar and scout wasps search the surrounding area for a new nesting site. After several hours, the scouts stimulate the rest of the swarm to leave the cluster and follow their pheromone trail to the chosen site. How scouts communicate to their swarm-mates that a site has been chosen and how they induce the swarm to depart are unknown. Video records of six Costa Rican swarms were used to quantitatively document changes in the frequencies of social behaviors leading to swarm departure. This was accomplished by going backward through the video record and following the behavior of individuals prior to their departure. Analysis of the behavior of scouts and inactive wasps indicated an increase in the frequency with which scouts bump into inactive wasps prior to swarm departure, as well as a shift in the behavior of inactive wasps from primarily receiving bumps to bumping others before departure. Thus, bumping is propagated by recently activated individuals before they take off. These observations suggest that not only is bumping an activation stimulus that causes swarm members to depart for the new nest site, but it is contagious, leading to its amplification throughout the swarm. Keywords: &ILEZMSV'SWXE6MGEXVSTMGEP[EWTHMWTIVWEPLSYWILYRXMRKEQTPMJMGEXMSRGSRXEKMSR Correspondence: TNWSRRIRXEK$Y[EPYQRMGSQ NIERRI$IRXSQSPSK][MWGIHY Received: .YP]` Accepted: *IFVYEV]` Published: %TVMP Copyright: 8LMWMWERSTIREGGIWWTETIV;IYWIXLI'VIEXMZI'SQQSRW%XXVMFYXMSRPMGIRWIXLEXTIVQMXWYRVIWXVMGXIHYWITVSZMHIHXLEX XLITETIVMWTVSTIVP]EXXVMFYXIH ISSN: `:SP2YQFIV Cite this paper as: 7SRRIRXEK4..IERRI60-RMXMEXMSRSJEFWGSRHMRKW[EVQIQMKVEXMSRMRXLIWSGMEP[EWT Polybia occidentalisTT Journal of Insect Science EZEMPEFPISRPMRIMRWIGXWGMIRGISVK .SYVREPSJ-RWIGX7GMIRGI`[[[MRWIGXWGMIRGISVK .SYVREPSJ-RWIGX7GMIRGI:SP`%VXMGPI 7SRRIRXEKIXEP Introduction Still unknown is how the scouts communicate to the cluster that a site has been chosen and how they stimu- In social insects that initiate colonies by means of swarm- late inactives to leave for the new site. Here, for & ing, three organizational challenges must be faced when #" we describe the behavior of wasps on the swarms disperse. The first is that a subset of the popula- swarm cluster during the period leading up to and in- tion, the scouts, must find and agree upon a suitable nest cluding liftoff. The mean frequency with which scouts site. Second, the scouts must induce the rest of the colony bump others in the swarm was found to increase as the to leave for the new site. Finally, scouts must guide the departure approaches and that wasps in the cluster shift migrating swarm of adults to the new site. The process their behavior from inactivity and receiving bumps to has been best studied in the honey bee, " ! moving over the cluster and bumping others before tak- Scout bees leave the temporary cluster to search for po- ing off. This is the first detailed, quantitative study of pre- tential nesting sites. After several hours or days, a single emigration swarm behavior for a social wasp and the first site is agreed upon based on quorum sensing by scouts; systematic study done on multiple swarms of the same that is, scouts favor a site at which they observe a suffi- species. cient number of other scouts visiting (Seeley and Visscher 2004). Once a site is selected, scouts return to the swarm and prepare it for liftoff by rapidly running over the Methods clustered bees and piping (producing short, rising sound Location pulses) (Seeley and Tautz 2001). The actual liftoff of the The study was conducted at Las Pumas, a private estate swarm, however, is triggered by buzz running, wherein 5 km west of Cañas, Guanacaste, Costa Rica (10°25N, excited bees run in zigzag patterns while pushing their 85°7W). Cañas is located in the tropical dry forest life way through the cluster and occasionally buzzing the zone and has a wet season extending from May into wings (Lindauer 1955; Esch 1967; Seeley and Tautz November. Much of the native vegetation at the site was 2001). Within a few minutes, the entire swarm takes to replaced by pasture, but tropical dry-forest species were the air and is then guided by scouts, which streak still represented. The colonies used in this study nested in through the flying swarm in the direction of the new nest trees and shrubs. site (Beekman et al. 2005). Swarming and videotaping Much less is known about the emigration process in The study site consisted of a pasture containing a scatter- swarm-founding social wasps. Best studied in this regard ing of low trees, predominantly $'$ $ is the Neotropical species & #".In (Sterculiaceae). Active nests of #" were moved Guanacaste, Costa Rica, reproductive swarms typically into this area for ease of observation. The twig bearing depart from the natal nest at the end of the wet season the nest was cut and carried to the study site, where it (Forsyth 1978; Jeanne 1999). Absconding swarms, was spring-clipped to a low twig in a tree in the pasture. however, may be produced at any time as a result of Each translocation was done after dark to ensure that all brood-raiding by ants (Bouwma et al. 2007) or when a adult wasps remained inside the nest. nest is damaged beyond repair by a vertebrate predator, severe weather, or accident. The entire adult population The day following the move, the colony was induced to abandons the nest, at first scattering widely, but within swarm by dismantling the nest in mid-afternoon. This 30 minutes coalescing into a single dense cluster on a gave the absconding population time to coalesce into a leafy twig near the nest (Bouwma et al. 2000, 2003). single cluster, but not enough time to find and move to a Scouts then begin to explore the surrounding area for a new nest site before dark. The size of the swarm was de- suitable nesting site (Bouwma et al. 2003). Scouts appear termined using a highly accurate predawn census tech- to be drawn from the ranks of foragers (Forsyth 1981; nique described elsewhere (Bouwma et al. 2000). After West-Eberhard 1982). They inspect several sites at first, the swarm had re-clustered following the census, its sup- but within a day or so narrow the choices to one. After porting twig was cut and spring-clipped to a wooden the scouts have reinforced their trail to the chosen nest board (30 cm wide x 22.5 cm high) mounted vertically on site by leaving pheromone spots on leaves, they induce a tripod and placed in the shade of the tree near where the other wasps in the cluster to follow the trail (Jeanne the swarm had re-clustered. We coaxed the swarm from 1975; Naumann 1975). Unlike in honey bees, the its twig and onto the board by carefully removing one clustered individuals of #" leave their resting leaf at a time (Figure 1). With the removal of the last bit site one by one over a period of 10 to 30 minutes and fol- of twig, the swarm formed a cluster no more than two low the trail individually (Jeanne 1981a; PJS, personal wasps deep on the board. At this point we began video- observation). If there are males in the swarm, they re- taping the swarm, using a digital camcorder (Sony DCH- main behind (Bouwma et al. 2000). Upon arrival at the HC1000) zoomed so that the board almost filled the new site, workers immediately begin construction of the LCD screen of the camcorder. Videotaping continued new nest (Jeanne and Bouwma 2002). until the last wasp departed from the board. Behavioral .SYVREPSJ-RWIGX7GMIRGI`[[[MRWIGXWGMIRGISVK .SYVREPSJ-RWIGX7GMIRGI:SP`%VXMGPI 7SRRIRXEKIXEP Figure 1. 8LIW[EVQSFWIVZEXMSRFSEVH&VERGLFIEVMRKXLIW[EVQLEWFIIRWTVMRKGPMTTIHXSXLIFSEVHERHPIEZIWERHX[MKWEVIFIMRK GEVIJYPP]VIQSZIHSRIF]SRIXSGSE\[EWTWSRXSXLIJPEXWYVJEGIJSVZMHISVIGSVHMRKSJFILEZMSV data were taken by viewing the tapes on a digital video- their designation, inactives did in fact become active cassette recorder (Sony DHR-1000). One swarm was shortly before departing, as we describe below. studied in 1998 and five in 2005. An additional swarm on a twig was videotaped and the video reviewed to de- The total number of wasps present on the board was de- termine whether the behavior of unmanipulated swarms termined every ten minutes by pausing the digital tape of differed qualitatively from that of our manipulated the swarm and counting. This record was used as a meas- swarms.
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