Journal of Chemical Ecology, Vol. 14, No.

1, 1988

UPWIND FLIGHT ORIENTATION TO PHEROMONE IN

WESTERN PINE BEETLE TESTED WITH ROTATING
WINDVANE TRAPS 1'2

JOHN A. BYERS

Department of Animal Ecology, University of Lund

S-223 62 Lund, Sweden
Department of Entomological Sciences, University of California
Berkeley, California 94720

(Received September 3, 1986; accepted January 20, 1987)

Abstract--In the first trap design, a rotating windvane was connected to a

30 x 30 x 30-cm "square box" sticky trap enclosing a synthetic pheromone
source (exo-brevicomin, frontalin, and myrcene) at the windvane's rotation
axis. A second design used the windvane attached to two tubular (19-cm-
diam. x 30-cm) sticky traps each suspended 120 cm from the same phero-
mone source and opposingly aligned "downwind" and "upwind" of the
windvane. Significantly more beetles of each sex of Dendroctonus brevicomis
LeC. (Coleoptera: Scolytidae) were caught on the downwind side compared
to the upwind side of the square-box design. Even larger differences in catch,
four times more males and 3.4 times more females, were found on the down-
wind tubular trap compared to the upwind one. The windvane trap design
provides rigorous evidence that insects, especially bark beetles, orient up-
wind to pheromone sources (from at least 1.2 m downwind until reaching the
source).

Key Words--Dendroctonous brevicomis, Coleoptera, Scolytidae, phero-

mone, anemotaxis, upwind orientation, exo-brevicomin, frontalin, myrcene.

INTRODUCTION

U p w i n d flight o r i e n t a t i o n b y i n s e c t s to p h e r o m o n e h a s b e c o m e a w e l l - k n o w n
fact o r p a r a d i g m . T h i s k n o w l e d g e is b a s e d l a r g e l y o n c o r r e l a t i v e e v i d e n c e b e -

1Dendroctonus brevicomis LeC. (Coleoptera: Scolytidae).

2Supported in part by grants from USA and Sweden, NSF(INT-8503520), STU (84-3937), NFR,
FRN, and SJFR.

189

0098-0331/88/0100-0189506.00/0 Q 1988 Plenum Publishing Corporation

190 BYERS

tween wind direction and observations of flying insects or trap catches in the
field. There are many studies on Lepidoptera and other insects orienting to pher-
omone sources in wind tunnels (Kellogg et al., 1962; Farkas and Shorey, 1974;
Kennedy, 1977, 1983; Kennedy et al., 1980, 1981; Card6, 1984). In some
larger insects it is possible to correlate wind direction measurements using
windvanes and simple observations of wing fanning (Elkinton et al., 1984).
With many smaller insects, such as bark beetles, it is difficult to document their
upwind flight orientation response to pheromone. Consequently, "very little
(olfactory) research has investigated orientation by flying beetles, nor has it
addressed the intricacies of orientation behaviour" (Borden, et al., 1985).
One reason this is so is that bark beetles have been difficult to fly in wind
tunnels. Flight exercise is required in at least some bark beeries before they will
take off flying upwind in pheromone-bearing wind (Scolytus multistriatus:
Choudhury and Kennedy, 1980) or before they will ignore light and respond
instead to attractant odors (Trypodendron lineatum: Graham, 1959). Even with
flight exercise, preliminary studies (Byers, Schlyter, and L6fqvist, unpublished)
on Ips typographus in a wind tunnel have been largely unsuccessful. The beeries
fly towards lights and seem to have difficulty orienting upwind without leaving
the plume and striking the side of the tunnel. The possibility that bark beetles
are "less accurate" than moths when orienting to a pheromone source may be
explained by differences in their requirements: male moths need to find females
while bark beetles must find only an aggregation spread widely over a tree.
Thus, it is possible that orientation mechanisms in flying bark beeries may be
different from those in better-understood insects.
Due to the inherent difficulties of wind-tunnel experiments and flight ob-
servations on small beetles, close-range orientation studies have been done in
the laboratory on walking bark beetles. The open arena bioassay has indicated
they walk upwind and chemotactically orient to pheromone sources (Wood and
Bushing, 1963; Birch and Wood, 1975; Byers and Wood, 1981) or to host odors
(Byers et al., 1985). The beetles could also do so under natural conditions,
although we have little or no evidence of this.
In the field, there are several studies on bark beetles that indicate, with
various degrees of credibility, that beetles fly upwind to pheromone sources,
presumably when stimulated by pheromone molecules within a plume. How-
ever, these studies were not truely rigorous, in that they relied on descriptive
observation or they utilized correlations between wind direction and trap catches.
For instance, McMullen and Atkins (1962) and Rudinsky (1963) observed that
Douglas-fir beetles flew "upwind" and directly to a pheromone source although
some "flew by and circled back before landing." Gara (1963) stated Ips par-
aconfusus take off in all directions, but usually with the wind. Upon encoun-
tering pheromone-laden air, they oriented against the wind and flew back to-
ward the source of attraction.
Later studies have attempted to quantify the orientation behavior by catch-
UPWIND FLIGHT ORIENTATION TO PHEROMONE 191

ing beetles in traps placed at or near a pheromone source (Chapman, 1962;

Gara, 1963; Coster and Gara, 1968; Seybert and Gara, 1970; Gray et al., 1972;
Tilden et al., 1979; Byers, 1983; Helland et al., 1984; Schlyter et al., 1987).
The conclusions of these studies, however, were all dependent on correlations
of trap catch with wind direction measured nearby (usually at one place) and
time-averaged. Thus, sometimes the wind direction may not have been aligned
with the "prevailing wind direction," and it was not known precisely what the
relative frequencies of beetle attraction/wind direction were over certain periods
during the test. Therefore, the objective of this study was to obtain direct evi-
dence of upwind orientation to a pheromone source which was both quantitative
and not dependent on correlative wind data. A pair of rotating traps on a wind-
vane which aligns with the wind provides a novel design for testing anemotactic
orientation to odor sources in insects. The hypothesis is that the "downwind"
trapping surfaces will intercept and catch significantly more beetles than the
control traps placed " u p w i n d . "

METHODS AND MATERIALS

A windvane was constructed that held either a box-type sticky trap or two
tubular sticky traps suspended on opposing arms. A 5-mm metal shaft served
as the axle which rotated inside two ball bearings fixed near the top of a 1.5-m
metal pole that was driven vertically into the ground. A polycarbonate vane was
attached to a support about 16 cm from the axle and counterbalanced by metal
weights (Figure 1). The dimensions of the vane were approximately 2 mm thick
x 15 cm top x 38 cm bottom • 39 cm and 45 cm sides. In the first trap design,
a square box was constructed from four, 30 • 30-cm, 6.3-mm wire-mesh
screens coated with Stikem Special, | Seabright Enterprises, Emeryville, Cali-
fornia. The box rested on 4-mm fiber board and was aligned so that the down-
wind side was perpendicular to the vane (Figure 1).
In the second trap design, 19-cm-diam. x 30-cm-high tubular sticky traps
were constructed from the same materials and suspended equidistant from the
axle (120 cm) on the ends of a 2 x 2-cm wooden beam (Figure 2). The two
opposing traps were aligned with the vane so that one trap would be consistently
downwind while the other was upwind of the pheromone source. The phero-
mone components of D. brevicomis, exo-brevicomin and frontalin (both
> 96 %), and the host component, myrcene ( > 98 %), all from Chem. Samples
Co., Cleveland, Ohio, were released individually from glass tubes (two for each
component) inside a dispenser (Byers and Wood, 1980; Tilden and Bedard,
1985). The release rates varied significantly with temperature but were esti-
mated to be about 3 mg/day of each component.
Beetles were collected from the sides of the box trap once or twice daily
(seven replicates) during its placement in the Sierra National Forest near Bass
 L)

160

"~120
(0
(J
8O

.O
E 4O
z

0
D L R U
Trap Side
FIG. 1. Upwind orientation of D. brevicomis to a pheromone source within a " b o x "
sticky trap (30 cm on a side) placed upon a windvane at 1.4 m height. The pheromone
components, exo-brevicomin, frontalin and myrcene, were each released at 3 mg/day
from within the box trap. Letters designate trap sides orientation: D = downwind, R =
right and L = left of downwind, and U = upwind.

@- \
beetles wind

300
@
D U
7:
t~ 200

!
t)

e~
100
E
-I

z
0 W
Downwind Upwind
Trap
FIG. 2. Upwind orientation of D. brevicomis to a pheromone source placed at the ro-
tation axis of a windvane (1.4 m height). Opposing tubular sticky traps (30 cm x 19
cm diam.) were aligned with the wind direction and each was 1.2 m from the rotation
axis. The pheromone components, exo-brevicomin, frontalin, and myrcene, were each
released at 3 mg/day in the forest.
UPWIND FLIGHT ORIENTATION TO PHEROMONE 193

Lake, Madera County, California, at 1000 m elevation (August 19-23, 1985).

The opposing traps of the second design were similarly picked of beetles (six
replicates) during testing in the same area (August 24-27, 1985). Wind speeds
were measured with a fan anemometer for several 1-min periods during general
alignment with the windvane in order to obtain an estimate of environmental
conditions. Effects of position on the distributions of catch between the down-
wind side and the other three sides of the box trap and also between the oppos-
ing traps were determined with the Wilcoxon matched-pair test. Binomial con-
fidence limits for sex ratios were calculated (Dyers and Wood, 1980) and
compared for significant differences with chi square.

RESULTS

The catches of D. brevicomis on the downwind, left and right of down-

wind, and upwind sides of the box sticky trap situated on the windvane are
shown in Figure 1. As might be expected, if the beetles were flying upwind just
prior to landing, the downwind side caught more beetles (32.3%) than either
the left (22.8%) or right sides (25.7%) or the upwind side (19.3%), and the
upwind side caught the fewest beetles. The downwind side caught significantly
more males and females than did the upwind side (1.5 and 1.9 times more,
respectively, (P < 0.05), and the sex ratios caught on the various sides were
not significantly different (Table 1). It is apparent, however, that each of the
four sides caught a significant proportion of the total catch.

TABLE 1. NUMBER OF D. brevicomis CAUGHT AND RATIOS (WITH 95 % BINOMIAL

CONFIDENCE LIMITS) ON WINDVANE TRAP WITH BOX TRAP OR OPPOSING TRAPS (SEE
FIGURES 1 AND 2)

Number caught Sex ratio

Male Female M/F" 95 % BCL

Box trap
Downwind side 74 92 0.80(0.59-1.09)
Left side b 53 64 c 0.83(0.58-1.19)
Right side b 56 C 76 0.74(0.52-1.04)
Upwind side 50 c 49 c 1.02(0.69-1.51)
Opposing traps
Downwind trap 156 182 0.86(0.69-1.06)
Upwind trap 39 c 53 C 0.74(0.49-1.11)

~Sex ratios were not significantly different (P > 0.05, X2).

bWhen one is facing upwind.
cWilcoxon matched-pair test showed trap surfaces had significantly different distributions of catch
for a particular sex compared to the corresponding downwind side or downwind trap (P < 0.05).
194 BYERS

In order to further test the anemotactic orientation hypothesis, two oppos-

ing traps were placed 120 cm from the pheromone source such that one trap
was always aligned downwind while the other was upwind (Figure 2). Here it
can be seen that the trap catches were even more different since four times more
males and 3.4 times more females were caught on the downwind trap compared
to the upwind trap (Figure 2). The differences in catch were statistically signif-
icant for both males and females (P < 0.05), while the sex ratios caught on
each trap were not significantly different (Table 1). The wind was observed to
change direction rather frequently (about once every minute) by as much as
180 ~ This was apparently more variable than observed earlier (Byers, 1983).
Wind speeds varied between 0.5 and 1.5 m/see with brief periods of calm (wind
changing). All tests were conducted during favorable flight weather (warm and
sunny, midday temperature 25-32~

DISCUSSION

The catches of D. brevicomis on the respective sides of the box sticky trap
on the windvane indicate that both sexes orient upwind to pheromone sources,
at least in the final landing stage. The fact that significant proportions apparently
were caught on sides other than the downwind side can be explained in several
ways. One possibility is that the wind direction changed more abruptly than
could be followed by the windvane so that anemotactic beetles were caught by
inappropriate sides. It is almost certain that the zigzag flight path of bark beetles
as they traverse a narrowing plume (as in other insects, Kellogg et al., 1962;
Traynier, 1968) would cause significant catches on the sides. The observation
of earlier scientists also indicate that beetles that fly out of the odorous air
attempt to turn and even circle back to the source, resulting in catches on the
upwind side. After the experiments were concluded, I became aware of another
example where a similar windvane trap (as in Figure 1) indicated that onion
flies oriented upwind to host-plant odors, at least within 20 cm of the source
(Dindonis and Miller, 1980). A box trap of much larger dimensions would have
probably given much larger differences in catch among the sides.
Tests of anemotaxis on a larger dimension were attempted by separating
the trapping surfaces about 120 cm from the source in both the downwind and
upwind directions. The higher catches on the downwind trap indicated that bee-
tles were responding anemotactically at least 1.2 m from the pheromone source.
The fact that some beetles were caught on the upwind trap can be explained by:
(1) the windvane may have sometimes lagged behind the change in wind direc-
tion and (2) not all anemotactic beetles were caught by the downwind trap and,
when they were not trapped at the source, they dispersed at random on even
continued more or less upwind until they were caught by the upwind trap. It
UPWIND F L I G H T ORIENTATION TO PHEROMONE 195

would have been interesting to have compared these results with a test where a
box trap was also placed over the source in order to reduce the possibility of
the second explanation above. Of course, many trap configurations are possible
in combination with the windvane concept.
Lindel6w and Weslien (1986) sum up our knowledge of L typographus
flight behavior during the dispersal flight: "when released, beetles always flew
downwind for at least 10-15 m, after which time they were impossible to ob-
serve." However, it is sometimes possible to observe beetles a few meters
before landing at a pheromone source during the late afternoon as sunlight is
transmitted through their wings. Others have used white sheets to offer a con-
trast (Chapman, 1962; Seybert and Gara, 1970). These observations are not
quantitative and are only correlated with wind direction, which is often imper-
fectly known.
Several quantitative studies have provided evidence for upwind orientation
to attractive sources in bark beetles but one or more drawbacks are apparent:
(1) the tests used marked beetles, often selected, that were subsequently re-
leased (unnaturally), (2) the tests were performed in unnatural settings (cages,
grassland), (3) the tests used fixed-position traps that may not have been opti-
mally aligned with the prevailing wind direction, (4) the wind direction was
averaged and could only be correlated on a gross scale since instantaneous catch
and wind direction were not determined, and (5) the wind measurements were
usually only at one location and somewhat distant from the traps, and in some
cases measurements were from weather stations several kilometers away.
Chapman (1962) found that more Trypodendron lineatum were caught on
the downwind sides of sticky traps placed over attractive logs (drawbacks 3-5
above). Gara (1963) performed an interesting experiment in which a line of
nine traps, placed parallel to the wind direction, released pheromones from each
in increasing and then decreasing amounts. Marked Ips paraconfusus were then
released at either the upwind or downwind areas of the line. The suggestion
was that since the percent recoveries on the traps were virtually identical, this
must mean that beetles released upwind flew with the wind until turning upwind
in response to pheromone, while beetles released downwind flew immediately
upwind in response to pheromone. Ignoring anemotactic theories and personal
observation of flying beetles, however, one could argue that there is a more
simple explanation of Gara's results. This is that beetles do not utilize wind
direction when orienting to pheromone as their release either upwind or down-
wind made no difference on their distribution of recapture along the trap line.
However, the windvane trap results presented here allow us to believe more
strongly in Gara's explanation.
Coster and Gara (1968) released marked D. frontalis in a grassland area
from four cardinal directions that were 30.5 m from a pheromone source. The
beetles released from the downwind direction were recaptured in the highest
196 BYERS

proportion, but the numbers were probably too low for reliable statistics (draw-
backs 1-5). Seybert and Gara (1970) found that caged L pini, artificially in-
duced to fly, flew proportionately more to the upwind wall (75%) when the
cage was placed 6 m downwind of a pheromone source (no statistics, drawbacks
2-4). In a similar experiment with caged D. ponderosae, Gray et al. (1972)
showed a statistically significant preference for upwind flight to attractants
(drawbacks 2-4). Tilden et al. (1979) used sticky traps in four cardinal direc-
tions at 1.5- and 4.5-m distances from a pheromone source to monitor the at-
traction of D. brevicomis. They found that the downwind trap consistently
caught more beetles than the upwind trap (drawbacks 3-5). Byers (1983) used
three trap rows, each 21 m long of 15 sticky traps, placed perpendicular to the
prevailing wind and each succeeding row was 4.6 m further downwind from a
pheromone source in order to observe the upwind orientation of L paraconfu-
sus. Both sexes were caught in a narrowing and concentrating pattern as they
approached the source, but the male catch at the source was significantly re-
duced compared to female catch (also compared to the sex ratios on the rows
of traps), indicating that male orientation to pheromone is less focused than that
of females (drawbacks 3-5).
Helland et al. (1984) released marked L typographus in grassland from
four cardinal directions either 12 or 20 m from a central pheromone source.
Beetle catch was monitored in 64 traps, each capable of catching beetles from
four cardinal directions with respect to the source, stationed in eight 24-m-long
" s p o k e s " radiating from the source. The pattern of catch was consistent with
an upwind orientation to pheromone, for beetles released downwind, and there
was evidence that beetles " s w a r m e d " around the source and did not always
move straight into the trap (drawbacks 1-5). In many respects their results are
reminiscent of those obtained by Gara (1963) above. Schlyter et al. (1987)
placed barrier traps in four cardinal directions each 3 m from a central phero-
mone source and found that L typographus was again caught in a pattem con-
sistent with upwind anemotaxis (although the wind measurements were taken
several kilometers away from the experiment, drawbacks 3-5).
The present study provides a method for continuously integrating wind
direction and flight intensity in a natural setting with naturally responding bee-
tles. Thus, the method of coupling a windvane with beetle traps circumvents
the five experimental drawbacks encountered in the past. However, much more
remains to be understood about close-range flight orientation (observed here)
and as well longer-range orientation that often was probed in the experiments
discussed above.

Acknowledgments--I am grateful to the USFS and W.D. Bedard, P.E. Tilden, and M.I.
Haverty for providing the facilities for research in the field. I am especiallygratefulto P.E. Tilden
for helpful assistanceand information. I thank D.L. Wood, Departmentof EntomologicalSciences,
University of California, Berkeley, and J. Lrfqvist, Departmentof Animal Ecology, Universityof
Lund, Sweden, for providing me with research facilities and funding.
UPWIND FLIGHT ORIENTATION TO PHEROMONE 197

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