Honey Bee Drone Flyways and Congregation Areas: Radar Observations

Author(s): Gerald M. Loper, Wayne W. Wolf and Orley R. Taylor, Jr.

Source: Journal of the Kansas Entomological Society, Vol. 65, No. 3 (Jul., 1992), pp. 223-230
Published by: Allen Press on behalf of Kansas (Central States) Entomological Society
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 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY
65(3), 1992, pp. 223-230

Honey Bee Drone Flyways and Congregation Areas?

Radar Observations

Gerald M. Loper,1 Wayne W. Wolf,2 and Orley R. Taylor, Jr.3

abstract: We an X-band
used radar unit to document honey bee (Apis mellifera) drone
flyways and drone
congregation areas (DCA's) in a nearly flat desert area. Within an
approximate 5.0 x 2.0 km area adjacent to a commercial apiary, 18 km of flyways and at
least 26 DCA's were located. The studies conducted during March and April in each of 4
years (1987-1990) confirmed that the location of flyways and DCA's were re-established
day after day and year after year. Based on film records of radar images, our definition of
a DCA is that site (? 100 m diam) where drones fly higher and are more numerous than
in adjacent flyways. Flyways formed alongside the most prominent structural or physical

features?e.g., tree lines formed in washes?but these flyways also branched, particularly
when additional tree lines became visible on the near (?200 to 250 m) horizon. Most
DCA's occurred at these
branch points; thus, the accumulation of drones at a DCA may
be a result of rapidturning and altitude changes as the drones re-orient and select their
next flight direction.The maximum height of drones in flyways was 21 m, whereas in
DCA's they flew mainly from 30-50 m above ground. Flyways are more predominant than
DCA's, and contain large numbers of drones. It is suggested that queens flying within or
to the upwind side of flyways should attract sufficient drones to become easily mated.

Honey bee (Apis mellifera L.) queens mate outside the hive (Janscha, 1771;
Koeniger, 1986) during one or more mating flights. Drones are produced by
colonies when the protein (pollen) status of a colony is optimal (Weiss, 1969;
Taber and Poole, 1974). When drones are mature they fly in the afternoon,
generally beginning about one hour before flights by virgin queens (Ruttner and
Ruttner, 1965; Ruttner, 1966). Most apicultural scientists believe that drones fly
to specific aerial locations known as "drone congregation areas" (DCA's) (M?ller,
1950; Zmarlicki and Morse, 1963; Ruttner and Ruttner, 1965). Researchers have
located DCA's by a "droning" sound that drones make as they fly and by the use
of queen sex pheromones (9-oxodecenoic acid, 9-ODA (Gary, 1963) or queens
tethered below a kite, balloon or long pole (Zmarlicki and Morse, 1963; B?ttcher,
1975; Tribe, 1982). Although precise information on the size, shape and dynamics
of drone flight within DCA's is not known, the following generalizations (Ruttner
and Ruttner, 1972; Ruttner, 1974) have emerged: (1) Drones appear to assemble
at the same locations year after year even when no queen is present; (2) Drones
respond with greater intensity to queen pheromones when flying within congre
gation areas; (3) Pursuit by drones declines as tethered queens are moved away
from a DCA; (4) Flight altitude of drones within a DCA is inversely related to
wind velocity; and (5) Drones locate DCA's quickly and appear within DCA's on
their first mating flight. It is generally accepted that drones fly to DCA's and then
virgin queens fly to these DCA's to mate and that most queens mate in DCA's.
Drones frequently fly 2 or 3 km but can fly 6 km or more (Ruttner and Ruttner,
1966; Taylor and Rowell, 1988). The literature provides little information on

1
USDA-ARS, Carl Hayden Bee Research Center, 2000 E. Allen Rd., Tucson, Arizona 85719.
2
USDA-ARS, Crop Insect Pests Mgmt. Res. Unit, Rt. 5, Box 808, College Station, Texas 77845.
3
University of Kansas, Department of Entomology, Lawrence, Kansas 66045.
Accepted for publication 10 January 1992.

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224 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

drone flyways, although Weiss (1969) does suggest that drones "seem to have a
reliable, solidly joined system of orientation, which directs them on a certain flight
route." Some studies suggest that the flyways of drones are related to physical
features in the area, i.e., low points on the horizon (Ruttner and Ruttner, 1966);
but there is no general explanation as to why DCA's are formed in specific areas.
From his studies in S. Africa, Tribe (1982) has described three environmental
factors in DCA formation/location: (1) shelter (wind), (2) barrier (vertical relief),
and (3) convection (thermal wind patterns). Patterns of features associated with
DCA's may have eluded investigators because it has not been possible to precisely
define the location and spatial configuration of DCA's due to changing wind
directions and velocity and the inability to define DCA's without the use of queen
pheromones.
Radar has been used recently to document drone flight activity and the location
of DCA's (Loper et al., 1987). Radar was also used in this study with the following
objectives: (1) document flyways and DCA's near an isolated apiary; (2) locate
flyways used by drones flying to and from the DCA's; (3) characterize physical
features or vegetational patterns that influence the location of either flyways or
DCA's; and (4) determine dynamics of drone flight to, within, and among DCA's
and the apiary.
We assessed drone activity by using an X-band marine radar as previously
described (Loper et al., 1987). "Targets" seen on the radar screen (plan-position
indicator, PPI) were recorded using time-lapse photography (16 mm movie cam
era). Individual frames from the 16 mm movie were evaluated to determine: (1)
location, diameter and height of drone activity in DCA's; (2) location, width and
height of drone flight in flyways; (3) the response of drones to queens or queen
pheromone tethered below a balloon and moved within and outside of DCA's
and flyways. Individual pictures were projected over aerial photos of the study
site in order to relate drone activity to physical and vegetational features. The
film was examined to relate the location of drone activity with respect to the radar
location and these locations were plotted on a map.
In this paper, we describe the results of our studies near Tucson, Az. which
were conducted during 2-3 week periods of March and April in each of four years:
1987-1990.

Experimental Situation

This study was conducted near a commercial apiary consisting of 50 to 70

colonies. This apiary was located 700 m to the west of a dry wash which generally
extended from south to north (Fig. 1). The predominant vegetation in and along
the wash was mesquite trees (Prosopis velutina Woot.) ranging from 4 to 5 m tall.
There were other scattered individual or small clumps of mesquite trees in the
area as well as scattered individual creosote (Larrea tridentata (DC) Coville)
bushes usually 1-2 meters tall. Numerous patches of bare ground also occurred
in the area. A single dirt road ("Trico Rd.," Fig. 1) extended north and south past
the apiary. A 3-wire 115 kV electric power line paralleled the west side of the
road and a 2-wire 14.4 kV line went from east-west, perpendicularly crossing the
major wash.
In order to discern both flyway activity and DCA's, the radar was moved to
triangulate the target area. At each new radar location, pictures were taken either

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VOLUME 65, NUMBER 3 225

rendition.
Artiste notexactly
Positions to scale.

Fig. 1. Honeybee drone flight: radar tracking observations. Flyways and drone congregation areas
(DCA's), Avra Valley, AZ.

before or after drones were flying in order to provide background for accurate
comparisons. The film was examined for the presence or absence of patterns of
drone flight activity in localized areas during drone flight time (colonies were
examined to establish that drones were flying during the observation periods).
During some periods of maximum drone flight, drone flyway and DCA activity
could be adequately documented in as little as 10 min and the radar moved to
another site. However, during studies of the influence of pheromones, temperature,
wind and time of day on drone flight patterns the radar was situated at one location
for the entire drone flight period for that afternoon.

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226 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Results

After locating a few DCA's and discovering some flyways in 1987, the radar
was again used in 1988-1990 to extend our observational area and to obtain data
on the dimensions of the DCA's and flyways. Data was also obtained on one
afternoon to record the relative density of insect targets of two DCA-sized areas
being observed simultaneously. The radar was located 244 m southwest of DCA
#9 & 10 (Fig. 1) and the number of insect targets in the DCA's were compared
to the number in a similar-sized area 244 m south of the radar (control area).
Periodically during the afternoon, the number of insect targets were counted in
both areas, first with the radar at 3? above the horizontal and then immediately
afterward at 5? elevation. Thus, insects within the areas flying at 16.4 ? 9 m and
25 ? 9 m above ground were counted (see Loper et al., 1987). At the same time,
two observers counted the number of drones leaving the colonies at the nearby
apiary (1 km SW) (Fig. 1). Each observer counted drones for 2 min/colony, and
each rotated among 7 colonies (14 total) completing a rotation approximately
every 18 min. Insect target counts were low and similar in the two areas for the
first two counting periods, but soon after a significant number of drones were seen
leaving the colonies (1330 hr, MST) more targets were being counted at the DCA's
than in the control area (Table 1). The ratio of counts between the two areas
continued to increase with time maximum at 1447 hr, just as the colony counts
showed a large number of drones had flown out. The maximum insect count in
the DCA's was 68; insect counts in the control area also maximized in that same
period at only 8. Insect target counts in the control area returned to zero by 1619
hr while targets in the DCA's, although decreasing in number, continued until
1655 hr. Insect targets in the control area could have been either workers, drones
or other species of insects. It is interesting to note that whatever insects were
flying in the control area, there were almost as many in the higher elevation as
in the lower (21 vs. 27). In the DCA's, there were 2.5 times more insects at the
lower elevation, supporting the data obtained in Kansas (Loper et al., 1987) which
showed that in a DCA, drones congregate within an area that can be described
as an inverted cone, with fewer and fewer drones at the higher elevations.
From many radar sites, both flyways and DCA's were observed, which appeared
only in the afternoons during documented drone flight times.
Flyways: Insect targets with linear movements within long and relatively nar
row (50-100 m) areas were documented on the radar movies (Fig. 1).When a
queen or a lure with 9-ODA was elevated in these areas, the insect targets re
sponded by converging near and downwind of the lure. Visually, and by use of a
net drone trap (Taylor, 1984), the insects were found to be drones. By comparing
the location of these insect targets on the pictures with other pictures (background)
taken either before or after drone flight times, the location of drone activity relative
to terrestrial could be accurately determined
features (?9-15 m depending on the
range). By taking pictures while increasing the radar beam elevation (i.e., from 2?
to 8? above the horizontal) estimates were made of the height at which the drones
were flying. The linear areas in which unidirectional movement of drones was
observed were generally 50-100 m wide and these "flyways" formed at various
distances (60-150 m) from, and parallel to, tree lines or roadways. In the nearly
?
flat terrain (-0.38% slope, S N), the flyways formed adjacent to the most

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VOLUME 65, NUMBER 3 227

Table 1. Comparison of insect targets counted in DCA's and a control area as observed by radar
with counts of drone flight from the nearby apiary. March 23, 1988.

Elapsed time Radar observations

Colony observations0
Insect targets/radar sweep No. drones/2 min
#9+10
DCA Control"Ratio
Time-DCA/ % of
(min)
(MST) 3?a 5? Total 3? Total
5? control Time Total

1310-1317 7 4 0 4 2 1 3 1.3 1312 0

1329-1348 19 2 0 2 0.1 1 1.1 1.8 1331 0.98
1348-1358 10 2 1 3 0.2 0.3 0.5 6.0 1349 1.17
1405-1408 3 18 6 24 2 2 4 6.0 1407 6.67
1428-1447 19 30 25 55 4 2 6 9.2 1428 19.87
1447-1458 10 40 20 60 3 2 5 12.0 1447 16.78
1504-1507 3 50 18 68 4 4 8 8.5 1505 15.91
1526-1543 17 40 10 50 5 3 8 6.2 1525 19.04
1553-1558 5 35 5 40 3 4 7 5.7 1548 13.08
1610-1617 7 25 10 35 4 2 6 5.8 1607 7.61
1619-1632 13 15 10 25 0 0 0 oo 1631 0.22
1632-1653 21 527000oo
1651-1655 4 303000OO
Totals 269 107 376 27.3 21.3 48.6
a
During each observation period, radar dish was operated at both 3? and 5? elevation above the
horizontal-observing areas at approx. 16.4 m and 25 m above ground, respectively.
b
Control area of radar scan was the same distance (244 m) south of the radar (and closer to the

apiary) as DCA #9 & 10 were north of the radar.

c
The drones leaving colonies were counted for 2 min/colony, 2 observers each rotated among 7
colonies (14 colonies total) approximately every 18 min.

prominent visual features available?tree lines in the washes or small tree groves
outside the washes. Generally the greater the height differential between the trees
and the adjacent shrubs, the closer the flyway would be to the tree line (although
rarely closer than 60 m). Flyways adjacent to structural features with low "ho
rizons" such as the dirt road, were located further (up to 150 m) from these
features. In this study area, flyways were predominant; from one advantageous
radar site, a continuous flyway 1700 m long interconnecting 3 DCA's was ob
served. Drones appeared to avoid flying over large barren areas. In one case the
flyway detoured around a barren area (100 x 150 m) and another flyway ter
minated in a DCA rather than continuing along the major tree line which ap
proached a large barren area (Fig. 1, #1). We also filmed two flyways that crossed
the N-S dirt road (and power line), others that crossed a tree line, and another
that passed between two converging tree lines that came within 7 m of each other
(Fig. 1, #22-24).
DCA 's: In addition to the flyways, other areas of drone concentration formed
within theflyways, in which drones were maneuvering (no linear movement doc
umented) and flying higher (DCA's). These areas were usually 70-100 m in di
ameter and often formed where terrain features were interrupted (i.e., terminated,
turned sharply, or where another tree line was visible within 200-250 m) (Fig. 1,
#2, 8, 9, 11, 12, 26). At these places, the flyway either terminated, changed
direction or branched to form two or more flyways. Several DCA's had the
appearance of a "Y" junction (e.g., #9 and 14). One DCA (#1) had the appearance

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228 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

of a "cul-de-sac"?a DCA with a single flyway terminating at the DCA. From

some radar sites, as many as 4 DCA's with connecting flyways were observed.
As mentioned, most DCA's appeared to be nearly circular at the lower elevations
but with much smaller diameters at higher elevations. However, two sets of DCA's
(#9-10 and #28-29) seemed to be "connected." At the lowest altitudes they had
the appearance of being a single, large, irregularly shaped DCA but, at higher
altitudes, each had two focal points of drone activity separated by at least 100 m.
There was flyway activity between them at higher altitudes than in adjacent
flyways. The area of #28-29 was over 300 m long and 150 m wide.
The height of drone activity varied during the afternoon. The average upper
limit of drones in flyways was 21 m although transient "bubbles" within flyways
were observed in which drones were flying higher. The altitude of drone flight
within a DCA was usually as high as 30 m, but in some "larger" DCA's (either
in diameter or drone density) drones were seen up to 55 m. As a result of mea
surements obtained from film and 35 mm pictures of the radar screen, our defi
nition of DCA a is: 1) that site at which drones maneuver and fly higher and are
more numerous than within the adjacent flyway, and 2) an area approximately
100 m in diameter which has a concentration of moving targets the location of
which does not drift with the wind and occurs in the same location day after day
and year after year.
The response of drones to queen pheromones elevated within DCA's was tested
on 3 occasions and also within flyways on 3 different occasions. After an initial
radar documentation of drone activity, a queen or lure was elevated via a helium
filled balloon. In each situation, the radar documented that most of the drones
converged on the lure, coming basically from downwind. The lure was then moved
(at a slow walking pace) either upwind or cross wind, stopping every 70-80 m for
at least 5 min. When moving upwind, many, but not all drones followed the lure
at least as far as 300 m. However, when the lure was moved cross wind, most
drones followed as far as the first 70 m, but at distances in excess of 100 m, only
a few (20-30) followed, and during the 5 min stop, most of these left the lure
although there were almost always 2-3 drones continually around the lure. As
the lure was being moved beyond the 100 m distance (cross wind) drone activity
in either the flyway or DCA returned to "normal" dimensions with typical un
disturbed flight activity.
All DCA's occurred at the same sites each day and year; although only the
larger DCA's could be detected on days with low drone density. Occasionally,
small, transient clusters of drones were detected by radar within both flyways and
DCA's at times when queen pheromone was not being exposed experimentally.

Discussion
Use of the X-band radar has permitted us to objectively locate drone flyways
and DCA's without the use of pheromones. The location of flyways and DCA's
along with observations of drone targets on the radar screen lead to several hy
potheses concerning drone orientation and honey bee mating behavior. The fly
ways appear to form in relation to visual cues; at certain places the flyways branch,
again often in relation to additional visual cues. At these branch points, the radar
indicates that drones maneuver rapidly with frequent changes of direction and
altitude thus creating a DCA. We suggest that the drones are reorienting and

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 VOLUME 65, NUMBER 3 229

perhaps flying higher to regain a sense of direction or search for additional cues.
Our study area was almost flat, the nearest hills were more than 7.5 km distant,
and vegetation was less than 5 m tall. It is apparent that orientation to low points
on the horizon and other geographic cues are not necessary for DCA formation.
The occasional small transient clusters within flyways and DCA's could be the
result of either biological or physical effects. Biological explanations could include:
1) drones chasing a virgin queen, 2) chasing and examination of "false targets,"
such as worker bees, other drones or other insects, or 3) a non-random distribution
or clumping of drones based on a time-pulse phenomenon in which large numbers
of drones leave and return to their colony in short periods of time (5-10 minutes,
Loper, pers. obs.).

Additionally, transient clusters may be explained on the basis of physical in

fluences suchas atmospheric turbulence. For example, small scale (<200 m)
thermals were observed to shift a large number of drones completely out of their
flight paths. Even smaller scale local wind convergences, eddy currents or local
thermal updrafts could influence drone flight in small areas (<20 m) which would
be detectable by the radar. Our studies tend to refute two of the hypotheses
suggested by Tribe (1982) but definitely support and give some definition to the
relationship between the "barrier" or vertical relief and drone flight behavior.
These findings suggest that virgin queens on mating flights could easily en
counter many drones in flyways and that matings might occur outside of DCA's.
We suggest that the conventional idea that DCA's are points where drones ac
cumulate for the purpose of mating needs to be reevaluated. In our study area,
even though there are many DCA's, flyways with thousands of drones are a very
prominent feature in the flight space. We acknowledge that these observations
were obtained near a large apiary; in other areas where only scattered feral colonies
and low drone populations exist, the formation of DCA's may be crucial to
successful matings.
It is clear that an X-band radar can detect large scale movements of drones
(and even honey bee swarms, Loper et al., unpubl.). A Doppler radar system
would be better in terms of interpreting flight dynamics and could reveal the
existence of minor pathways which are difficult to detect with X-band radar. At
this point in time, the biggest gap in our knowledge of the mating behavior of
honey bees is in not knowing the flight behavior of virgin queens and what
stimulates her to be receptive to drones. To obtain information on the flight paths
of individual targets, a different system using a transmitter small enough to fit on
a drone or queen together with a sophisticated tracking system is required.

Acknowledgment
We acknowledge the grant funding this research provided by the USDA-ARS,
Pilot Test/Integrated Pest Management Committee, Beltsville, Md.

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