Sounds Associated with the Mating Behavior of a Mutillid Wasp*

HAYWARD G. SPANGLER AND DONALD G. MANLEY2

USDA, SEA, Bee Research Laboratory, 2000 East Allen Road, Tucson, AZ 85719
ABSTRACT
Pairs of Dasymutilla joxi produced a coordinated female, he produced 2-4 honks/sec. If the female was
sound sequence while mating and attempting to mate. not receptive the male would seize her and continue to
Two distinct sounds are made by these insects. Males honk while the female stridulated 8-12 times/sec. The
and females produce stridulation, sounds made by rub- male appeared to immobilize the female partially with
bing abdominal parts together. Males can also honk by vibration and caused her to alter her stridulatory chirp.
vibrating their wings. If mating was successful, the We suggest that this sound sequence plays an essential
male and female both stridulated, alternating with each role in mating.
other at ca. 1-sec intervals. As the male approached a

Male and female velvet ants (Mutillidae) produce
frictional sounds by rubbing abdominal segments to-
gether, referred to as stridulation. Males vibrate their
wings while advancing toward and grasping females
with their legs (Brothers 1972). When the wings are
both at the field site and in a sound-proof room. A
Bruel and Kjaer 12.7-mm Condenser Microphone
Type 4134 was used with a General Radio 1560-P42
preamplifier/power supply. Recordings were made
with a Uher 4000 L tape recorder. The recorded

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vibrating, a different type of sound called "honking"
is produced.
Stridulatory organs typically consist of a "file" with
many more or less regular ridges and a "scraper"
with one or more ridges or teeth. In the Mutillidae
the stridulatory file is on the base of the 3rd visible
abdominal tergite. The scraper is a sharp ridge on
the underside of the 2nd visible abdominal tergite
(Hinton et al. 1969). When stridulating, the wasp
draws the 3rd abdominal segment in and out so that
the file passes under and against the scraper to pro-
duce vibration.
The function of stridulation in Mutillidae has never
been clearly determined. It may serve to get the
sexes together for mating (Manley 1976, Mickel
1928), although other observations indicate that the
attraction between the sexes is primarily olfactory
and visual (Linsley et al. 1955). Vibration from a
stridulating mutillid might also aid in dislodging soil
particles and assist the wasp's movement through
tight passages in the soil (Spangler 1973).- There is
also a possibility that stridulation has a defensive
role by startling and otherwise repelling predators
(Hinton et al. 1969, Schmidt and Blum 1977).
We report here sounds produced by both the male
and the female of the Mutillid Dasymutilla joxi
(Cockerell) during mating activity.
sounds were analyzed with a Kay Electronic 7029 A
spectrum analyzer, a Spectrum variable band pass
filter, a General Radio 1523 level recorder, and a
Tektronix 564B oscilloscope.
RESULTS AND DISCUSSION
During copulation and attempts at copulation both
the male and female mutillids made sounds. The fe-
male stridulated, varying her "chirp" considerably de-
pending on the situation. Though the male could
stridulate, he could also "honk" by vibrating the tho-
rax and wings with the flight muscles with the wings
resting over the abdomen, much as the queen honey
bee does when piping (Simpson 1964).
When mating, the male wasp mounted the female
and grasped her about the thorax with his legs. Gen-
erally the 2 wasps then fell to one side and lay on
the ground with the tip of the male abdomen curved
ventrally over the tip of the female abdomen. The
female typically began to stridulate as the male
grasped her and continued as long as the male con-
tacted her.
As a male approached the female,- and often before

MATERIALS AND METHODS

An aggregation of Dasymutilla joxi was found near
a livestock watering tank ca. 35 mi north of Tucson,
Ariz, around the nesting site of Diadasia sp. bees.
Both males and females were present, numerous at-
tempted matings were observed, but only a few suc-
cessful matings were seen.8
Recordings of the sound produced by the wasps
while they mated or attempted to mate were made
FIG. 1.—Oscillogram of sounds produced by a mating
1
Received for publication Oct. 25, 1977. Mention of a pro- pair of velvet ants (upper trace). Sound patterns direct-
prietary
2
product does not constitute an endorsement by the USDA. ly over the blocks on the lower trace were produced by
Dept. of Entomology, Univ. of Arizona, Tucson 85719.
8
We thank Stephen Taber, III for discovering and leading the female. Those between the blocks were produced by
us to the aggregation of mutillid wasps. the male. Oscilloscope writing speed—1 sec/division.
389
390 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA [Vol. 71, no. 3

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*#*.iMyim,

TIME (seconds)
FIG. 2.—Sonograms of sounds produced by a pair of velvet ants while the male was attempting to mate. Upper
pattern shows female chirps and the lower male honks. Contour lines represent 6dB.
May 1978] SPANGLER AND MANLEY : SOUNDS OF A MUTILLID WASP 391

he grasped her, he began to honk. The honk was
much louder than the sound from the stridulating
female. During the honk the wings and the dorsal
surface of the thorax vibrated rapidly. After grasp-
ing the female while honking, the male pumped his
abdomen and attempted copulation by reaching the
tip of his abdomen toward that of the female. (On
the 3 occasions when a receptive (unmated) female
was observed, copulation began.) Honking by the
male ceased as copulation began and was replaced by
stridulation. Throughout copulation (which lasted
for over one min on one occasion) the male and fe-
male stridulated in alternating sequences, each lasting
ca. 1 sec (Fig. 1). The frequency of sound from the
male's stridulation was slightly lower than the fe-
male's and thus could be distinguished by ear. When
copulation was complete, they ceased stridulating and
separated.
When the female was not receptive to copulation,
500 hz, while that of the female was ca. 600 hz. Both
had considerable energy in overtones extending well
above these frequencies. Each of the 7 intensity steps
on the contour sound spectrograms represent 6dB.
Because of the distinct frequency difference be-
tween the honks and chirps, a filter was used to sepa-
rate the recorded sounds on a sound level recorder.
A band-pass filter set at 20-400 hz passed mostly
male honk energy while 5-20 khz allowed the passage
of some female chirp energy without much interfer-
ence from the male. Fig. 3 shows the time sequence
of these 2 sounds on the same time base. Upward
movement of the traces represents the presence of in-
sect sound energy, while downward or horizontal
movement represents absence of sound. The 1st series
of 13 male honks (upper trace) is the same series
shown in Fig. 2. The female chirping shown in Fig.
2 corresponds to that beginning at the 8-sec point
(lower trace). The male produced between 2 and 4

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the male nevertheless approached her and began to
honk. He grasped her, both typically fell on their
sides, and the male extended and pumped the tip of
his abdomen to attempt copulation. Throughout this
process the male honked and the female stridulated.
The female moved her abdomen apparently to avoid
copulation, and, as she did so she altered her stridu-
latory chirp. A total of 24 pairs consisting of males
and apparently mated females was placed together,
one pair at a time, in the laboratory. In all cases the
male approached the female, honked, and seized her.
In 21 cases the female stridulated, altering her chirps
when the male honked. In the remaining 3 cases the
female stridulated only while the male honked.
In addition to being louder, male honking was
lower in frequency than either male or female stridu-
lation. The male honk (Fig. 2) has its primary
energy around 225 hz. The maximum energy of male
stridulation, on the other hand, appeared to be near
honks/sec while the female produced 8-12 chirps/sec.
When the male honked, changes were heard in the
chirps made by the female. The chirps became ir-
regular, and varied in the intensity, duration, and
time intervals separating them. This altered chirping
is apparent in the lower trace at time when the upper
trace indicates honking by the male.
Males which had recently attempted copulation
with females often continued to honk when moved
to the presence of another male. On 5 occasions these
honking males attempted to mount other males; the
male being mounted stridulated and altered its chirp
much like a female.
We suggest that these sound sequences are impor-
tant in successful mating. Observations indicated that
males which failed to honk repeatedly while attempt-
ing to mount females were unlikely to achieve and
maintain copulatory position. A female had a tend-
ency to stand stationary once a male touched her

20
TIME (seconds)

FIG. 3.—Sound level traces of the sounds produced by a pair of velvet ants while the male was attempting to
mate. Upper trace shows the intensity pattern of male honking while the lower trace shows the intensity pattern
of female chirping. Sound produced by the insects is represented by upward movement of the trace. When
no sound was present the trace drops, becomes horizontal and continues on that plane, then rises during the next
sound.
392 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA
body and honked. The function of the stridulation is
not clear. Perhaps it results from the tendency of
these insects to stridulate whenever prevented from
moving freely. The fact that the female chirp changes
in response to male honks, however, indicates that a
communicative function may be involved.
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