ECOLOGYANDBEHAVIOR

Flight and Locomotion Activity of the Sweetpotato Weevil

(Coleoptera: Brentidae) in Relation to Adult Age, Mating Status, and
Starvation
SEIICHI MORIYA1 ANDSATOSHI HIROYOSHI2

Okinawa Prefectural Agricultural Experiment Station

4-222 Sakiyama-cho, Naha, Okinawa 903, Japan

J. Econ. Entomol.91 (2): 439-443 (1998)

ABSTRACT The flight and walking performance of Cylas formicarius (F.) adults were evaluated
with respect to their age, mating status, and the degree of starvation by using a flight mill and an
actograph system. Males showed a much higher Bight ability than females in all ages tested between
o and 42 d old after spontaneous exiting from the sweetpotato root. Locomotion activity of the male
was also higher than that of the female for 0- to 28-d-old adults, but the difference was not as large
as that of flight activity. Except for the effects of a 3-d starvation period on locomotion, mating status

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and starvation with only water for 1-3 d had little affect on flight or locomotion ability. No distinct
signs of migratory behavior of adults were detected. Females seemed to disperse mainly by walking
because of their extremely low flight ability.

KEY WORDS sweetpotato weevil, flight mill, actograph, age, mating, starvation

THE SWF.ETPOTATO WEEVIL,Cylas formicarius (F.), is the
most destructive pest of sweet potato, Ipomoea batatas
(L.) Lamarck, in tropical and subtropical areas (Chal-
fant et aI. 1990, Jansson and Raman 1991). A series of
detailed studies were conducted to develop a phero-
mone trap monitoring system for the weevil in the
southern United States (Jansson et al. 1992a, b, 1993).
In Japan, the sweetpotato weevil is only found in the
Ryukyu archipelago, the southernmost part of the
territory (Yasuda and Kohama 1990, Setokuchi 1990),
and is designated as a plant quarantine pest by Japan's
Plant Protection Law. No live weevils or any part of
fresh host plants are allowed to be brought into main-
land Japan without permission of the Plant Quarantine
Board. Therefore, an eradication program for this
weevil is on-going using the sterile insect technique or
a male annihilation method (Moriya 1995b, Setokuchi
et al.I996). Assessment of adult dispersal ability is one
of the basic requirements in evaluating a pheromone
trap monitoring system (Mason et aI. 1990) and is also
needed to promote the eradication program of the
weevil (Sugimoto et al. 1994, Miyatake et al. 1995).
However, relatively limited data are available, all of
which were analyzed on the basis of male adult
catches by the pheromone trap in the field. Therefore,
information on the quantitative aspects of the dis-
persal are needed. This is especially true for females
because no female dispersal data can be obtained
I Current address:Japan InternationalResearchCenter for Agri-
cultural Sciences.1-2 Ohwashi.Tsukuba,Ibaraki 305. Japan.
~Current address: Okinawa Prefectural Fruit Fly Eradication
Project Office.123 Maji,Naha, Okinawa902, Japan.
using a sex pheromone trap that attracts only males in
the field.
The current studies measured the effects of adult
age, mating status, and starvation on dispersal activity,
flying and crawling, of sweetpotato weevils using a
simple flight mill and an actograph.
Materials and Methods
Insects. Weevils used in this experiment were
reared on sweetpotato roots ('Beniazuma') at 25 ± 1°C
and a photoperiod of 14:10 (L:D) h. The original pop-
ulation of the stock culture was collected in sweet-
potato fields in Yomitan Village, in the central part of
Okinawa Island, Japan, in September 1988 and reared
at the Okinawa Prefectural Fruit Fly Eradication
Project Office adjacent to the Okinawa Prefectural
Agricultural Experiment Station where the experi-
ments were performed. Adult age was defined by the
days after adult eclosion in the experiment on mating
status, or by the days after the spontaneous exiting of
sweetpotato roots in other treatments. Most of the
individuals stay within sweetpotato roots for 6-9 d
after their adult eclosion (Sutherland 1986).
Measuring Devices, Both flight and locomotory ac-
tivities of the adults were automatically measured for
=23 h by the simple flight mill and actograph systems,
with photoelectric switches described previously
(Moriya 1994, 1995a). Flight activity was represented
in terms of the total distance flown (meters) calcu-
lated from the total number of rotor revolutions re-
corded by the flight mill system. In the actograph
system, locomotory activity was evaluated by the total
number of times a weevil crosses an infrared beam.

0022-0493/98/0439-0443$02.00/0 © 1998 EntomologicalSocietyof America

440 JOURNAL OF ECONOMIC
The beam is set between photoelectric switches at-
tached to the container (31.5 by 31.5 by 12.5 mm). A
detailed configuration and dimension of each system
was described previously (Moriya 1994, 1995a). De-
viation of the sensitivity of each set of photoelectric
switches was carefully adjusted to the same level be-
fore each run according to the results of Moriya
(1994). The 2 systems were installed separately in
walk-in chambers set at 27 ± 1°C and a photoperiod
of14:l0 (L:D) h. Up to 8 and 12 adults were measured
simultaneously by the flight mill and acto graph sys-
tems, respectively.
Measurement of Activities. Adult activities were
evaluated in relation to 3 parameters: adult age, mating
status of the individuals, and degree of starvation. In
the flight mill experiment, 32-70 male and female
adults were tested at 0, 7, 14, 21, 28, and 42 dafter
exodus in relation to their age, and 30 male and female
adults were tested in relation to mating status and
degree of starvation in 0, 1, and 3 d. In the actograph
experiment, 78-114 males and females were treated at
0,7,14, and 28 d after exodus, and the other treatments
were same as in the flight mill experiment. All the
individuals were measured only once and they were
set on the devices 2-5 h prior to scotophase. Only
accumulated activities during =23 h were recorded in
these systems.
Adults emerging from roots were collected daily
and both sexes were maintained in a petri dish with a
slice of sweetpotato root. Adults were used for the
experiment when they reached respective ages of 0, 7,
14,21,28, and 42 d. To obtain virgin individuals, insects
that pupated were removed from roots and sexes were
separated just after adult eclosion. Four virgin adults
were reared per container. Two days before testing,
combinations of 1 male with 3 females and 1 female
with 3 males were made, respectively, from the virgin
adults and then each single male and female among
combinations were selected for study. After treat-
ment, females were dissected to check whether they
had mated (e.g., presence of sperm in their sper-
matheca). Only the males whose partners were in-
seminated or the females that had sperm in their body
cavity were regarded as mated individuals and all
others were discarded. Adults aged 28 d after adult
eclosion were chosen in the experiment for mating
status.
Effect of starvation was evaluated by providing only
water for 1 or 3 d before measurement. Adults tested
were 14d old and were allowed to mate freely until the
test was initiated. Preliminary experiments showed
that starvation for up to 3 d did not affect adult survival
rates; all of 30 starved males and females were alive
after 5 d and there was no significant difference on the
survival rate between treated and untreated individ-
uals even after 10 d (P < 0.05; Fisher exact proba-
bility test, Abacus Concepts 1994).
Data Analysis. Flight distance (calculated in
meters) in the flight mill and number of crosses to an
infrared beam in the actograph were transformed by
IOglO(X + 1) before analysis. Data were ana-
lyzed using 2-factor factorial analysis of variance
ENTOMOLOGY Vol. 91, no. 2
1.0 -+- Male
..•. Female
'+
E
Qj"
o
c:
~ 0.5
'0
E
.2>
;;::
g;
-'
f,
0
".
......
J.. .
J " .
0 7 14 21 28 42
Age in d
Fig.!. Flight distance in meters of C.formicarius adults
measured by the flight mill system during 23 h in relation to
their age at 27 :':: 1°C and a photoperiod of 14:10 (L:D) h.
Error bars = standard error of the mean.
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(ANOVA) to determine the difference between treat-
ments, and the means were separated by the Fisher
protected least significant difference (LSD) (Abacus
Concepts 1994). For comparing the data between the
sexes in relation to their ages, the Mann-Whitney
V-test was performed (Sokal and Rohlf 1981).
Results
Effect of Adult Age. Weevil flight ability, expressed
by distance flown, was significantly different both be-
tween sexes and among ages (Fig. 1; P < 0.001); that
is, males flew much longer than females and O-d-old
males flew less actively. The interaction between sex
and age was also significant according to ANOVA (P =
0.014), which was derived mainly from the low flight
ability of male adults at age 0 d. Comparing flight
activity of males with that of females in each age class,
males had a significantly higher ability to fly than
females in all age classes from 0 to 42 d (P < 0.05 in
o d and P < 0.0001 in other treatments). It was noted
that 41.9%of the males and 90.6%of the females never
flew, especially females aged 0, 7, and 42 d (Fig.l). The
theoretically calculated average distance flown in
males and females were 53.6 and 2.3 m for 23 h, re-
spectively, based on the number of revolutions of the
flight mill rotor and its length. The maximum distance
was estimated to be 1,687 m in males and 330 m in
females by the same procedure.
Degree of locomotion activity was significant be-
tween sexes (P < 0.01) and among ages in days (P <
0.001) and the interaction between sex and age was
also significant (P < 0.05) (Fig. 2). These were
caused by relatively low activity of young females as
shown in Fig. 2. Because males moved more actively
than females, only in age 0 d (P < 0.01), the differ-
ence between sexes was much smaller compared with
that for flight ability. No peak period was observed in
the flight and locomotion activities in relation to adult
ages (Figs. 1 and 2).
April 1998 MORIYA AND HrnOYOSHI: FuCHT AND LocOMOTION OF SWEETPOTATO WEEVIL 441

2.0 2.5

~ Male
2.0
T
0 Female
T
.L .L

0.5

Age In d
Fig. 2. Index of locomotion activity of C. formicarius
adults measured by the actograph system during 23 h in
relation to their age at 27 ± l°e and a photoperiod of 14:10
(L:D) h . Error bars = standard error of the mean.
o
Mated Virgin
Fig. 4. Effect of mating status on index of locomotion
activity of 28-d-old (after adult eclosion) C. fOl7nicarius
adults measured by the actograph system during 23 h at 27 ±
l°e and a photoperiod of 14:10 (L:D) h. Error bars = stan-
dard error of the mean.

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Effect of Mating Status. Neither mating status (P =
0.750) nor interaction between mating status and sex
(P = 0.862) significantly affected the flight activity
of28-d-old adults (Fig. 3). MalesfIew much more than
females (P < 0.0001) as was expected from the
results shown in Fig. 1. Males flew and females rarely
flew regardless of their mating experience.
Mating status did not affect the locomotion activity
of adults (Fig. 4; P = 0.195) and the effect of sex and
interaction between sex and mating status were not
significant. Thus, mating experience had no effect on
the flight and locomotion activities of the weevils.
ERect of Starvation. Starvation for 1 and 3 d also had
no effect on the flight activity of 14-d-old adults (Fig.
5; P = 0.604) and no interaction was detected be-
tween sex and starvation period (P = 0.591). Flight
ability of males was higher than that of females (P <
O.000 1) , which agrees with results of effect of mating
status. These data indicate that males were active
flyers even when deprived of food for 3 d.
Conversely, starvation had a significant effect on
the locomotion activity of weevils (Fig. 6; P < 0.001),
but there was no interaction between sex and period
1.5
fZl Male
of starvation (P = 0.319). Although the locomotion
activity was not different between control (0 d) and
1-d starvation period (P = 0.428), the differences
between control and 3-d starvation and between l-
and 3-d starvation were both significant (P < 0.01).
The results show that a starvation period of 3 d sig-
nificantly lowered the locomotion activity of the
adults. A significant difference was also detected be-
tween the sexes (P < 0.01). However, this was not
consistent with the other result already shown in Fig.
2. No definitive conclusion could be drawn from these
results.
Discussion
It is very clear that male weevils flew more actively
than females, regardless of their age. This result sup-
ported direct observations from the field and labora-
tory (Reinhard 1923, Cockerham et al. 1954) and light
trap records (Cockerham et a1.1954). Although there
1.5
--- Male
..•.. Female

o Female
'+
E'1.0
~<:
.•
;;
'0
E
'"
SO.5
8'
-'
.......
I""
I o
o
Mated Virgin o 1 3
Starvation period In d
Fig. 3. Effect of mating status on flight distance in meters
of28-d-old (after adult eclosion) C.fonnicarius adults mea- Fig. 5. Effect of starvation period on flight distance in
sured by the flight mill system during 23 h at 27 ± l°e and meters of 14-d-old C. fonnicarius adults measured by the
a photoperiod of 14:10 (L:D) h. Error bars = standard error £light mill system during 23 h at 27 ± l°e and a photoperiod
of the mean. of 14:10 (L:D) h. Error bars = standard error of the mean.
442 JOURNAL OF ECONOMIC
2.0
...••
..• " Female
1.5
.......
.........
'+ I "
II>
Ql
II>
r'" '"I
".
II> ",
b 1.0
c:i
g;c:
-' 0.5
o sibility of movement of weevils between islands where
o 1 3 weevils have to fly without a break (Miyatake et al.
Starvation period in d
Fig. 6. Effect of starvation period on index oflocomotion
activity of 14-d-old C. fonnicarius adults measured by the
actograph system during 23 h at 27 :!:: ec and a photoperiod
of 14:10 (L:D) h. Error bars = standard error of the mean.
were big differences in the flight ability between
sexes, as noted by Moriya (1995a), flight ability of the
weevils as a whole was low compared with other
insects. For example, in the flight mill experiments,
melon flies, Bactrocera cucurbitae Coquillett, fly >2 km
without a break (Nakamori and Simizu 1983), brown-
winged green stink bugs, Plautiastali Scott, fly 5 km on
an average and maximum 54 km/24 h (Moriya 1987),
and the average distance flown by diamondback
moths, PluteUa xylostella (L.), is 5-13 km/48 h (Shirai
1991). Compared with dispersal distances of male
weevils by using sex pheromone traps in the field,
maximum 280 m/16 h (Mason et al. 1990),55-64 mIl
d (Sugimoto et al.1994), and 32-163 mIl d (Miyatake
et aI.1995), our value of53.6 m/23 h falls within these
values.
Locomotion activity was also significantly higher in
males than in females; however, the difference was
much smaller than that of the flight activity. Sakuratani
et al. (1994) observed that females were less active
than males during the hot season. Considering their
low flight activity, it is likely that the females disperse
mainly by walking in the field (Moriya 1995a).
Shimizu and Moriya (1996) showed that the absolute
walking distance could be estimated with consider-
able accuracy from the count number obtained by an
actograph using photoelectric switches in Euscepes
postfasciatus (Fairmaire), which is another important
weevil pest of sweet potato (Yasuda and Kohama 1990,
Moriya 1995b). Because this method might be appli-
cable for C. formicarius (Shimizu and Moriya 1996),
the number of infrared beam crosses in this study
could be a good indicator of the accumulated distance
walked without obstructions and stimuli.
Except for the results on the effect of starvation
obtained by using an actograph, mating status and
starvation period had no effect on flight nor locomo-
tion activities. However, decreased locomotion activ-
ity in starved adults was probably caused by lack of
food. Starvation did not accelerate the locomotion
activity.
ENTOMOLOGY Vol. 91, no. 2
In summary, adult age, mating status, and degree of
Male
starvation had little effect on the movement of the
sweetpotato weevil. The movement of the weevils,
including flying and walking, was almost trivial, and no
migratory movement, a distinct behavioral and phys-
iological syndrome (Dingle 1972), was detected ac-
cording to the definition of Southwood (1962) under
the laboratory conditions in this study.
The current flight mill system was simple and could
not measure the continuous flight distance of the wee-
vil, but could estimate the accumulated flight distance.
However, information on continuous flight is another
important parameter, especially considering the pos-
1997). Therefore, for advanced studies, a flight mill
system should be computer controlled.
In addition, lack of migratory characteristics,
especially low flight ability, appear to be favorable
with regard to the weevil eradication program in the
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Ryukyu archipelago where the nearly 200 small islands
are scattered in the ocean and therefore the control
action would only be applicable to a limited area.
However, because of low migration activity of the
sweetpotato weevil, more dense groupings of phero-
mone traps should be used in the field to catch and
eliminate wild male adults for male annihilation
method and the sterile males also must be released
more in the target area for sterile insect technique,
compared with highly locomotive insects.
Acknowledgments
We thank J. Koyama (Okinawa City, Japan) and I" J.
Mason (Purdue University, West Lafayette, IN) for critically
reading the manuscript. Thanks are extended to the Okinawa
Prefectural Fruit Fly Eradication Project Office (Okinawa
Prefectural Government, Okinawa, Japan) for providing fa-
cilities for the experiment.
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Received for publication 7 April 1997; accepted 29 October
1997.