1979] Seeley, Morse & Visscher — Honey Bee Swarms 
107 
the nestbox surface, punctuated by bursts of wing buzzing. They 
would butt against individual nestmates and, buzzing steadily, 
attempt to bulldoze through small clusters of scouts. Esch’s (1967) 
analysis of these movements reveals that when running about a buzz 
runner produces short pulses of 180-250 Hz wing vibrations, and 
then shifts to steady blasts of 500 Hz buzzing when contacting other 
bees. 
The importance of the scouts’ return to the swarm was revealed 
when we measured what fraction of the swarm had visited the nest 
site before the entire swarm flew there. This was done while awaiting 
the island swarm’s first flight by recording over the last 60 min 
before lift-off the fraction of bees visible at the nestbox that were 
marked with blue. We marked 143 bees; the percentage of blue- 
marked bees among the bees observed at the nestbox, based on 130 
counts, was 28.9% ± 16.3% (x ± 95% confidence margins). Thus 
approximately 495 bees from the swarm (95% confidence limits: 
316-877 bees), or only about 5% of the swarm, had visited the nest 
site before lift-off. 3 Evidently only a very small minority of a 
swarm’s bees knows the precise location of the new nest site, and it is 
their task to guide the large majority to the new home. Because the 
ratio of ignorant to informed (scout) bees is so large, it appears 
important that as many scouts as possible be back at the swarm 
when lift-off occurs to guide the swarm to the new nest site. 
2. Lift-off of Swarm 
Upon returning to the swarm, some of the blue-marked scouts sat 
quietly on the swarm cluster, others resumed their dances advertis¬ 
ing the chosen nest site, and still others began performing buzzing 
runs across the swarm’s surface. The first vigorous, distinct buzz 
runner was sighted 43 min before lift-off, the second at 33 min, the 
third at 26 min, and by 18 min before lift-off 4 vigorous buzz 
runners were seen simultaneously scrambling over the swarm. Con¬ 
current with the rise in buzzing run frequency came an increase in 
the intensity of a much higher-pitched piping sound. We could 
neither identify which bees were producing this sound, nor had we 
the means of characterizing it or of quantifying its intensification. 
3 This 5% estimate is actually an overestimate since only 46 of the 143 blue-marked 
bees had been painted by the time the proportion counts began at the nest box. 
Therefore early estimates of the proportion of marked bees were smaller than if all 
143 bees had been marked and so, in turn, the net average of 28.9% of the nestbox 
bees being marked is an underestimate of the fraction of scouts which were painted. 
