The honeybee (Apis mellifera) queen mates during nuptial £ights, in the so-called drone congregation area where many males from surrounding colonies gather. Using 20 highly polymorphic microsatellite loci, we studied a sample of 142 drones captured in a congregation close to Oberursel (Germany). A parentage test based on lod score showed that this sample contained one group of four brothers, six groups of three brothers, 20 groups of two brothers and 80 singletons. These values are very close to a Poisson distribution. Therefore, colonies were apparently equally represented in the drone congregation, and calculations showed that the congregation comprised males that originated from about 240 di¡erent colonies. This ¢gure is surprisingly high. Considering the density of colonies around the congregation area and the average £ight range of males, it suggests that most colonies within the recruitment perimeter delegated drones to the congregation with an equal probability, resulting in an almost perfect panmixis. Consequently, the relatedness between a queen and her mates, and hence the inbreeding coe¤cient of the progeny, should be minimized. The relatedness among the drones mated to the same queen is also very low, maximizing the genetic diversity among the di¡erent patrilines of a colony.
-In Apis mellifera the estimated average number of drones visiting a drone congregation area (DCA) was 11 750 + 2 145. Drones of the species Apis cerana, A. koschevnikovi, A. dorsata and A. mellifera, which pursued a queen dummy moving in circular course, flew in a comet shaped formation. Median numbers of drones in a comet ranged from 9 drones (A. koschevnikovi) to 31 drones (A. mellifera).In none of the species we observed aggression between drones. Drone density behind the queen and distance to the queen seemed to be adjusted to avoid collisions between drones. The median flight speed ranged from 2.6 m/s (A. koschevnikovi) to 4.1 m/s (A. dorsata). The median duration of a drone's presence in the mating comet did not exceed 2 seconds. Drones of all species had the ability of high acceleration (10 to 20 m/s 2 ). Either by overtaking or leaving/entering the comet drones seem to compete for more promising positions. Only drones flying in a limited space of not more than 2000 cm 3 behind the queen were successful in grasping the dummy.Apis reproduction / mating behavior / drone congregation area / drone competition / drone numbers
At a natural drone congregation area freeflying drones were attracted by a fast-moving queen dummy and the pursuits of drones were stereoscopically recorded (Fig. 1). The reconstruction of 192 flight paths from successfully approaching drones in chronological three dimensional sequences (Fig. 4) lead to the following results: 1. The alignment of the drone's longitudinal body axis coincides fairly well with the line connecting drone and queen (drone-queen-axis), its mean angular deviation from this line being only 14 ~ . Angles between -5 ~ and 5 ~ occur most frequently (Fig. 5B). Thus, drones head straight to the queen. 2. Lateral deviations from the drone-queen-axis most frequently lie between -30 ~ and 30 ~ (Fig. 5A) which corresponds to the drone's binocular visual field. 3. The drone's heading was continuously adjusted to the actual target, mean turning speed being 1890~ 4. The results lead to the conclusion that honeybee drones choose the shortest way to a fast and not predictably moving mate. A comparison with earlier observations suggests that a drone's mating success depends not only on his skills to win a race but also on his persistence within a group.
Summary — Colony defense and predatory behavior of Vespa multimaculata was observed at the entrance of a natural nest site of Apis nuluensis. When V multimaculata was present, guard bees frequently performed body shaking behavior: bees lifted the tip of the abdomen slightly spreading the wings and exposed the Nasonov gland. The oscillation of the abdomen tip had an amplitude of 83° (SD = 50°; n = 84) and its duration was 0.08 s (SD = 0.01; n = 84). The individual behavior resulted in the rapid formation of a group of guard bees and induced body shaking of neighboring bees. The hunting V multimaculata workers hovered about 5-10 cm in front of the nest and blocked the direct flight path of bees returning to the nest. Returning foragers are regularly diverted from the nest entrances and attracted to the body shaking guard bees where they land beyond the reach of hovering V multimaculata. In contrast to Apis cerana, A nuluensis exposes its Nasonov gland during body shaking. During a period of 12 h one V multimaculata worker caught 14 bees. At the end of the observation we brought the hornet (tethered to a wire) into contact with a group of bees at the nest entrance. The A nuluensis guard bees immediately balled and killed the hornet with heat.Apis nuluensis / Vespa / behavior / defense / predation
Summary — The recent discovery of a morphologically distinct Apis type in mountain areas of Sabah raised the question of its taxonomic status. After it became evident that the drone flight period did not coincide with other cavity dwelling Apis species of Borneo, a new species, Apis nuluensis was described (Tingek et al, 1996). Here drone flight observations and results are reported in detail. A total number of 255 drone flights (50-60 drones) were observed at one colony. The drones started to fly at 1044 hours and the flight activity reached a peak shortly after 1200 hours. The last drones flew at 1312 hours. The complete separation of A nuluensis drone flight time from the other two cavity dwelling Apis species of Borneo functions as a premating barrier and results in complete reproductive isolation. This seems to be significant because overlap of habitats occurs in the transition zone of the upper dipterocarp forest and lower mountain forest (between 1 500m to 1 700m).Apis nuluensis / reproductive isolation / drone flight / Borneo
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