Allsopp, M.H.

Beekman, M.

Jordan, L.A.

Oldroyd, B.P.

Wossler, T.C.

Stellenbosch University SUNScholar Repository

Apidologie 38 (2007) 436–437 Available online at:c© INRA/DIB-AGIB/ EDP Sciences, 2007 www.apidologie.orgDOI: 10.1051/apido:2007028Scientific noteA scientific note on the drone flight time of Apis mellifera capensisand A. m. scutellata*Lyndon A. Jordana, Michael H. Allsoppb, Benjamin P. Oldroyda, Theresa C. Wosslerc,Madeleine Beekmanaa Behaviour and Genetics of Social Insects Lab, School of Biological Sciences A12, University of Sydney, NSW 2006,Australiab Honeybee Research Section, ARC-Plant Protection Research Institute, Private Bag, X5017, Stellenbosch, 7599,South Africac DST-NRF Centre of Excellence for Invasion Biology, Department of Botany & Zoology, University of Stellenbosch,Private Bag X1, Matieland, 7602, South AfricaReceived 27 February 2007 – Revised 14 May 2007 – Accepted 15 May 2007Apis mellifera capensis / Apis mellifera scutellata / drone flight time / hybrid zone / South AfricaHoneybee drones go on mating flights during aspecies-specific period of the day. During this pe-riod, drones leave the colony and fly to drone con-gregation areas where they may mate with visitingqueens (Koeniger et al., 2005). Honeybee queensmate multiply, are all of a similar size and se-crete the same sex attractant pheromone, 9-ODA(Koeniger and Koeniger, 2000). Thus, in areaswhere different honeybee species occur in sympa-try, the potential for interspecific mating is high.Where more than one species (Koeniger et al.,1994; Hadisoesilo and Otis, 1996) or subspecies(Koeniger et al., 1989) inhabits the same area, mat-ing times tend to be offset such that there is nooverlap of mating flights of the different species.The drone flight time may vary across the rangeof a species in response to the presence of het-erospecifics. For example, the drone flight time ofApis cerana in Japan (where it is the only nativeApis species present) extends from 1300 h to 1700h (Yoshida et al., 1994). In Sri Lanka however,where multiple species co-occur, the drone flighttime of A. cerana is truncated, with flights only oc-curring between 1530 and 1730 h (Koeniger andWijayagunesekera, 1976). This shift in flight timemay be an adaptive response to the prevention ofinterspecific matings. Divergent drone flight timeswithin a species may also be evidence of speciation,occurring before reproductive isolation is complete(Otis et al., 2000; Oldroyd et al., 2006).Corresponding author: Lyndon A. Jordan,lyndonjordan@gmail.com* Manuscript editor: Stan SchneiderThe honeybees of South Africa, the Cape hon-eybee Apis mellifera capensis (hereafter capensis)and the African honeybee Apis mellifera scutellata(hereafter scutellata), are separated by a zone ofoverlap situated mainly within the Great Karoo eco-tone (Hepburn and Crewe, 1991). The exact natureof this zone of overlap is unclear, but neither sub-species appears to be able to increase its range,despite capensis’s ability to become a social para-site of other honeybee subspecies (Allsopp, 1992)and scutellata’s potential for invasiveness (Winston,1992). This status quo (Hepburn and Radloff, 1998)may be due to an asynchrony of mating flight times.Here we test the hypothesis that scutellata andcapensis have offset mating flight times by compar-ing the flight times of drones of the two subspeciesat a single locality in Stellenbosch, Western Cape,South Africa. We determined the drone flight timesof four colonies of scutellata and four colonies ofcapensis by observing the time at which drones leftthe colonies over a period of 27 days during Novem-ber and December 2006.Scutellata colonies were obtained from a re-gion north of the hybrid zone in Douglas, NorthernCape, two months prior to observations. Capensiscolonies were obtained within the capensis nativerange. Colonies were located at separate apiariesbut within 2 km of each other. The two subspeciescannot be kept in the same apiary because of prob-lems with interspecific parasitism.To determine each subspecies’ typical droneflight times, we made preliminary observationsfrom sunrise to sundown. We then observed theexperimental colonies 30 minutes prior to andArticle published by EDP Sciences and available at http://www.apidologie.org or http://dx.doi.org/10.1051/apido:2007028A scientific note on the drone flight time of Apis mellifera capensis and A. m. scutellata 437051015202530351334 1404 1434 1504 1534 1604 1634Solar TimeMean Percentage of Drone FlightFigure 1. Mean (± s.e.) percentage of total num-ber of drones leaving A. m. capensis (shaded bars)and A. m. scutellata (open bars) colonies per fifteen-minute interval.following the first and last drone flights recordedin the pilot studies. One observer per subspeciescounted all drones leaving each of four coloniesfor one minute before moving to the next colony.Hence, at every colony, drone exits were countedfor one minute every five minutes. Observationswere made simultaneously by the second observerat the other subspecies’ apiary. We calculated thenumber of drones leaving per fifteen-minute periodacross all days for each colony and then averagedacross colonies. Drones were given a score accord-ing to their time of departure equal to the number ofminutes after 1400 h (solar time) that they left thecolony. Solar time was calculated by adding or sub-tracting 4 minutes for each degree of latitude westor east of the centre of the time zone respectively(Otis et al., 2000). As Stellenbosch is 6.5 degreeseast of the time zone centre (Cape Town) solar timewas calculated by subtracting 26 minutes from thelocal time. The frequency distributions of departingdrones across time intervals were compared using aKolmogorov-Smirnov test (Fig. 1).The drone flight times of capensis and scutel-lata were not significantly different (P = 0.541,D = 0.286, N = 14), indicating that drones ofboth subspecies perform mating flights simultane-ously in the zone of natural overlap and that inter-subspecies matings are likely to occur unless dronecongregation areas are spatially isolated (Koenigerand Koeniger, 2000). Hence, the causes that main-tain the status quo must be sought in reasons otherthan offset mating times.ACKNOWLEDGEMENTSWe thank Christian Fransman, Lee-Ann Noach-Pienaar, Willem Jan Boot and Johan Calis for theirhelp in the field, and Theuns Engelbrecht for the giftof scutellata colonies. BPO and MB are supportedby the ARC. MB is further supported via a SydneyUniversity Research Fellowship.Note scientifique sur la période de vol des mâlesd’Apis mellifera capensis et d’A. m. scutellata.Eine wissenschaftliche Notiz über die Drohnen-flugzeit von Apis mellifera capensis und A. m.scutellataREFERENCESAllsopp M. (1992) The Capensis Calamity, S. Afr. Bee J. 64,52–57.Hadisoesilo S., Otis G. (1996) Drone flight times confirm thespecies status of Apis nigrocincta Smith, 1861 to be aspecies distinct from Apis cerana F, 1793, in Sulawesi,Indonesia, Apidologie 27, 361–369.Hepburn H.R., Crewe R.M. (1991) Portrait of the CapeHoneybee, Apis mellifera capensis, Apidologie 22, 567–580.Hepburn H.R., Radloff S.E. (1998) Honeybees of Africa,Berlin, Springer-Verlag.Koeniger G., Koeniger, N., Pechhacker H., Ruttner F. (1989)Assortative mating in a mixed population of Europeanhoneybees, Apis mellifera ligustica and Apis melliferacarnica, Insectes Soc. 36, 129–138.Koeniger G., Koeniger N., Tingek S. (1994) Crossfostereddrones of Apis cerana Fabricius, 1793 and Apiskoschevnikovi v. Buttel-Reepen, 1906 fly at their species-specific mating times, Insectes Soc. 41, 73–78.Koeniger N., Wijayagunesekera H.N.P. (1976) Time of droneflight in three Asian honeybee species (Apis cerana, Apisflorea, Apis dorsata), J. Apic. Res. 15, 67–71.Koeniger N., Koeniger G. (2000) Reproductive isolationamong species of the genus Apis, Apidologie 31, 313–339.Koeniger N., Koeniger G., Gries M., Tingek S. (2005) Dronecompetition at drone congregation areas in four Apisspecies, Apidologie 36, 211–221.Oldroyd B.P., Reddy M.S., Chapman N.C., Thompson G.L.,Beekman M. (2006) Evidence for reproductive isolationbetween two colour morphs of cavity nesting honey bees(Apis) in south India, Insectes Soc. 53, 428–434.Otis G., Koeniger N., Rinderer T.E., Hadisoesilo S.,Yoshida T., Tingek S., Wongsiri S., Mardan M. (2000)Comparative Mating Flight Times of Asian Honey Bees,7th Int. Conf. on Tropical Bees, Chiang Mai, Thailand,International Bee Research Association.Winston M.L. (1992) The Biology and Management ofAfricanized Honey-Bees, Annu. Rev. Entomol. 37, 173–193.Yoshida T., Saito J., Najigaya N. (1994) The mating flighttimes of native Apis cerana japonica Radoszkowski andintroduced Apis mellifera L in sympatric conditions,Apidologie 25, 353–360.

A scientific note on the drone flight time of Apis mellifera capensis and A. m. scutellata

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A scientific note on the drone flight time of Apis mellifera capensis and A. m. scutellata

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