Appl Entomol Zool (2014) 49:411–419
DOI 10.1007/s13355-014-0263-1
ORIGINAL RESEARCH PAPER
Survival of Bactrocera latifrons (Diptera: Tephritidae) adults
under constant and fluctuating low temperatures
Shun-ichiro Takano
Received: 25 September 2013 / Accepted: 27 March 2014 / Published online: 12 April 2014
Ó The Japanese Society of Applied Entomology and Zoology 2014
Abstract Bactrocera latifrons (Hendel) is believed to
have originated in Southeast Asia but has invaded Hawaii
and most recently East Africa. This insect has also been
recorded on Okinawa Island, the far south of Kyushu
Island, Japan. To assess the overwintering ability of B.
latifrons adults, survival at constant temperatures (8, 10,
12, 14, 15 °C) and under fluctuating thermal regimes
(FTRs) was investigated. At 14 or 15 °C, more than 30 %
of females survived for 90 days. Time required to kill 95 %
of B. latifrons at 8 °C was estimated to be 13 days; at
10 °C, 29 days; and at 12 °C, 38 days for females, and 8,
17, and 24 days at the same above temperatures, respec-
tively, for males, suggesting low cold tolerance of this
species. The results show that females survive cold tem-
peratures better than males. Under an FTR of 11 °C (22 h)/
20 °C (2 h) (average 11.8 °C) survival of females drasti-
cally increased compared to that at a constant temperature
of 12 °C, whereas the survival of males increased slightly.
Survival under FTRs indicates that adult B. latifrons may
not overwinter in the north of Tanegashima Island, located
30 km south of Kyushu Island, Japan.
Keywords Cold tolerance  Distribution  Fluctuating
thermal regime  Overwintering  Tropical
S. Takano
Yokohama Plant Protection Station, 1-16-10 Shin-yamashita,
Naka-ku, Yokohama, Kanagawa 231-0801, Japan
Present Address:
S. Takano (&)
Faculty of Agriculture, Kyushu University, 744 Motooka,
Nishi-ku, Fukuoka 819-0395, Japan
e-mail: takanoshun@agr.kyushu-u.ac.jp
Introduction
Northward range expansion has recently been reported in
many insect pests that originate in tropical or subtropical
regions, and some of these cause damage to plants in their
new habitats (Halbert and Manjunath 2004; Matsukura
et al. 2009; Morse and Hoddle 2006; Yukawa et al. 2007).
Several factors may affect the poleward range expansion of
phytophagous insects, such as availability of host plants,
temperature, photoperiod, natural enemies, and competing
species (Kiritani 1998; van Lenteren et al. 2006). For
tropical insects, winter cold is a critical factor that may
limit their poleward range expansion, and their level of
cold tolerance has been studied to assess their overwin-
tering ability in potential distribution areas (Ashihara 2007;
Kandori et al. 2006; McDonald et al. 2000; Nakata 2006;
Takano et al. 2012).
Cold tolerance and the overwintering ability of insects
have been studied in several ways. Survival of insects at
constant low temperatures in laboratories has been most
commonly used as an index of cold tolerance (Ashihara
2007; Kandori et al. 2006; McDonald et al. 2000; Nakata
2006; Takano et al. 2012). Although a study on constant
temperatures is of fundamental importance and rather
easy to compare with previous studies, it may be difficult
to extend the interpretations to field situations because
there are diurnal fluctuations, and temperature is not
constant in natural habitats. Therefore, survival of insects
at fluctuating low temperatures that simulate diurnal
fluctuations in temperature, or survival of insects in the
field have been also studied (Ashihara 2007; McDonald
et al. 2000). It is known that insects survive longer at low
temperatures when they are exposed to a fluctuating
thermal regime (FTR) than to constant temperatures
(Coulson and Bale 1996; Hanč and Nedvěd 1999; Koštál
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et al. 2007; Renault et al. 2004). In tropical insect species,
Renault et al. (2004) showed that survival drastically
increased under an FTR of 5 °C (22 h)/20 °C or higher
(2 h). Although average daily maximum temperatures in
winter in temperate regions seldom reach 20 °C, the body
temperature of insects rises in sunlight (Akiyama and
Nishida 2013; Digby 1955; Parry 1951) indicating that
FTRs may affect the survival of tropical insects in the
field in temperate zones.
Bactrocera latifrons (Hendel) (Diptera: Tephritidae) is
believed to have originated in Southeast Asia, but in the
1980s it invaded Hawaii (Vargas and Nishida 1985), and
then Tanzania in 2006 and Kenya in 2007 (Mwatawala
et al. 2007; Mziray et al. 2010). In Japan, B. latifrons was
first found in 1984 on Yonaguni Island, to the far south of
Kyushu Island (it was eradicated in 2011) (Fukugasako and
Okamoto 2012). Thereafter, it was found on Okinawa
Island, 500 km northeast of Yonaguni Island, in 2010 and
further spread is of concern (The Ministry of Agriculture,
Forestry and Fisheries of Japan 2013).
Bactrocera latifrons attacks the fruit of Solanaceae and
Cucurbitaceae (Mziray et al. 2010; Vargas and Nishida
1985). Although occurrences of B. latifrons have been
reported only in tropical or subtropical regions to date, fruit
of Solanaceae or Cucurbitaceae, potential host plants of
this fruit fly, are also available in temperate zones such as
Kyushu, Shikoku, and Honshu Islands, Japan. Therefore, if
the cold tolerance of B. latifrons is high, it may expand its
distribution further poleward.
A few studies have been conducted on the cold tolerance
of B. latifrons. Vargas et al. (1996; 1997) showed that B.
latifrons completed its development and reproduced at
16–32 °C, but the responses of this fruit fly to temperatures
below 16 °C have not been studied. Shimizu et al. (2007)
suggested that B. latifrons may overwinter as the adult
stage on Yonaguni Island, Japan.
In this study, survival of adult B. latifrons at constant
temperatures (8, 10, 12, 14, 15 °C) and FTRs (5 °C/20 °C,
8 °C/20 °C, 11 °C/20 °C, 11 °C/14 °C: 22 h for low tem-
peratures and 2 h for warm temperatures) were investi-
gated to evaluate the overwintering potential in Japan.
Because prior exposure to cold temperature induces cold
tolerance in other tropical insects (Kandori et al. 2006;
McDonald et al. 2000), the effects of cold-acclimation on
cold tolerance were also examined.
Materials and methods
Insects
Bactrocera latifrons used in this study were from a stock
culture initiated with insects collected on Okinawa Island,
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Appl Entomol Zool (2014) 49:411–419
Japan, in December 2010. They were maintained as
described by Hirabayashi et al. (2012) with some modifi-
cations at 27 ± 1 °C and 60 ± 10 % relative humidity
(RH) under 14 L:10 D. Eggs were collected using ovipo-
sition devices (Ishida et al. 2005). Hatched larvae were
reared on an artificial diet based on wheat bran. Emerged
adults were maintained in a wire-mesh cage (220 mm
long 9 150 mm wide 9 150 mm high) and provided with
water and a 4:1 (weight) mixture of sugar and protein
hydrolysate AY-65 (Asahi food & Healthcare, Tokyo,
Japan). Copulation and oviposition were observed within
10 days from adult emergence.
Effect of cold acclimation
To determine the effect of cold acclimation on the survival
of B. latifrons, the survival rates of flies after exposure to
various acclimation conditions were examined.
Acclimation of immature and adult stages
Eggs collected from a stock culture at 27 °C were trans-
ferred to the acclimation conditions (20 ± 1 °C,
60 ± 10 % RH, 12 L:12 D) within 6 h of oviposition.
Larvae and adults were maintained in the acclimation
conditions in the same manner as the stock culture. For the
tests, 20- to 30-day-old adults were used.
Acclimation of adult stages
Adult flies (10- to 15-day-old) obtained from the stock
culture were maintained in a wire-mesh cage (220 mm
long 9 150 mm wide 9 150 mm high) with water and
a mixture of sugar and AY-65 and were exposed to 15
or 17 °C for 2, 4, or 7 days with a 12 L:12 D light
regime.
Ten adults obtained from each acclimation treatment
were transferred to one plastic cup (6.0 cm diame-
ter 9 3.5 cm high) containing a filter paper soaked in
water (a total of 50 females and 50 males were used for
each treatment), and exposed to 8 °C for 7 days. After
exposure, they were warmed to 27 °C to assess mortality.
Survival was confirmed by touching adults with forceps 6 h
after removal from the low temperature. Adults that could
not turn over when they lay on their backs were regarded as
dead because such weakened adults were doomed to die.
In the experiments on survival under constant low
temperatures and FTRs described below, non-acclimated
insects were used because acclimation conditions tested in
this study did not enhance the cold tolerance of B. latifrons
(see ‘‘Results’’).
Appl Entomol Zool (2014) 49:411–419
Survival under constant low temperatures
To determine the lethal time of non-acclimated insects
under constant low temperatures, adult flies (10- to 15-day-
old) were exposed to constant temperatures (8, 10, 12, 14,
15 °C). Survival of adults was examined in two ways
because at 8, 10, or 12 °C, flies did not move and it was
difficult to confirm their survival at these temperatures,
whereas at 14 and 15 °C flies moved and their survival was
confirmed at these temperatures.
At 8, 10, or 12 °C
Ten adults (10- to 15-day-old) in a plastic cup (6.0 cm
diameter 9 3.5 cm high), containing sugar and a filter
paper soaked in water, were exposed to 8 °C for 3, 5, 7, 8,
10, and 15 days (total, 380 adults); 10 °C for 7, 10, 12, 15,
20, and 25 days (total, 400 adults); and 12 °C for 7, 10, 12,
15, 17, 20, 25, 30, and 35 days (total, 560 adults). To
determine the effects of food resources on survival, adults
in plastic cups containing only water were also examined at
8 °C for 3, 5, 7, 10, and 15 days (total, 460 adults); 10 °C
for 4, 7, 10, 12, and 15 days (total, 320 adults); and 12 °C
for 5, 7, 10, 12, and 15 days (total, 290 adults). After
exposure, flies were warmed to 27 °C and mortality was
assessed as described above in the experiments on the
effects of cold acclimation.
At 14 or 15 °C
Fifteen females and males (10- to 15-day-old) obtained
from a stock culture were maintained at 14 or 15 °C (12
L:12 D) in a wire-mesh cage (220 mm long 9 150 mm
wide 9 150 mm high) with water and a mixture of sugar
and AY-65. Ninety insects were used for each temperature.
Their survival was monitored every 2–3 days until they
died.
Survival under FTRs
Survival under FTRs was investigated using the same
methodology as experiments for survival under constant
low temperatures at 14 or 15 °C. Each FTR consisted of
22 h (1400–1200) of low temperature and 2 h (1200–1400)
of high temperature, i.e., 5 °C (22 h)/20 °C (2 h) (average
6.3 °C), 8 °C (22 h)/20 °C (2 h) (average 9 °C), 11 °C
(22 h)/20 °C (2 h) (average 11.8 °C), 11 °C (22 h)/14 °C
(2 h) (average 11.3 °C). Survival was confirmed between
1200 and 1400 (at 14 or 20 °C). A total of 90 insects were
used for each FTR. To assess the effects of food resources
on survival, survival of adults provided only with water
was also examined.
413
Reproductive abilities of surviving females
To assess the effects of low temperatures on the repro-
ductive ability of females, egg production and hatchability
of females were examined. After exposure to the constant
low temperatures of 8, 10, and 12 °C for different periods
of time and the FTR of 11 °C (22 h)/20 °C (2 h) for
90 days, adult females were maintained in a wire-mesh
cage (220 mm long 9 150 mm wide 9 150 mm high)
with water and a mixture of sugar and AY-65. Adult flies
used in those experiments (10 to 15-day-old), were
assumed to be mated before exposure to low tempera-
tures. When male survivors were available, they were
maintained with the females. Within 10 days of removal
from the low temperatures, females were individually
transferred to a plastic cup (15.0 cm diameter 9 9.0 cm
high) for oviposition. An oviposition device (Ishida et al.
2005) was placed in the plastic cup for 24 h. Collected
eggs were moved to petri dishes (9.5 cm diame-
ter 9 2.0 cm high) with moistened filter paper to observe
hatchability.
Data analysis
The differences in survival rates between non-acclimated
flies and flies with acclimation treatments were compared
using Dunnett’s test after arcsine transformation. The dif-
ferences between males and females were compared using
Fisher’s exact test. The time required to kill 95 % of the
population at a temperature, Ltime95, was estimated using
logistic regression at 8, 10, or 12 °C. Differences between
lethal time estimates were considered to be statistically
significant if 95 % fiducial limits did not overlap. Fisher’s
exact tests were used to compare the percentages of sur-
viving females that laid fertile eggs between the rearing
temperature (27 °C, control) and low temperatures, and
between sugar-provided females and only water-provided
females at the same exposure temperature and duration.
These analyses were conducted with version 2.15.2 of the
R software (R Core Team 2012).
To assess the potential overwintering area of adult B.
latifrons, outside air temperature in the winter, from
December to March (1.5 m above the ground, mean from
1980–2010) for the six sites was obtained from the website
(Japan Meteorological Agency 2013a; Fig. 1; Table 1).
Results
Effect of cold acclimation
The survival rate of adult flies following exposure to 8 °C
for 7 days did not increase for all acclimation treatments
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414 Appl Entomol Zool (2014) 49:411–419

Fig. 1 Distribution map of the

sites of the temperature data
collected (1980–2010)

Table 1 Average daily Sites The number of days (°C)

temperatures in the winter
(1980–2000, December to City Latitude Longitude \8 B8 \10 \11 B12 \14 \15
March) at six sites
Ibusuki 31°150 N 130°380 E 0 15 45 70 102 113 121
0 0
Tanegashima 30°43 N 130°59 E 0 0 0 0 51 81 97
Yakushima 30°230 N 130°400 E 0 0 0 0 49 82 97
Nakanoshima 29°500 N 129°520 E 0 0 0 0 57 90 105
Amami 28°230 N 129°300 E 0 0 0 0 0 0 12
Okinoerabu 27°260 N 128°420 E 0 0 0 0 0 0 0

tested (Fig. 2). The survival rate of females reared at 20 °C
from eggs to adults did not significantly differ from that for
non-acclimated females (Dunnett’s test, p [ 0.05),
whereas the survival rate for males was significantly lower
than for non-acclimated males (Dunnett’s test, p \ 0.05;
Fig. 2). Survival rates decreased with increasing acclima-
tion period (2–7 days) for acclimation at 15 or 17 °C
except there was no decrease from 4 to 7 days at 17 °C
(Fig. 2). Females showed significantly higher survival rates
than males in all treatments except for the non-acclimated
insects and insects treated with 15 °C for 7 days (Fisher’s
exact test, p \ 0.05; Fig. 2).
Survival under constant low temperatures
At 8, 10, or 12 °C
As the exposure temperature increased, Ltime95 also
increased in insects that were provided with water and
sugar (Fig. 3). In insects provided with only water,
Ltime95 increased slightly but there was no significant
difference between Ltime95 at 10 and 12 °C (Fig. 3). At
8 °C, Ltime95 did not increase following provision of
sugar to the flies. Females showed significantly greater
Ltime95 than males when water and sugar were provided,

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Appl Entomol Zool (2014) 49:411–419
Fig. 2 Survival of Bactrocera latifrons after exposure to 8 °C for
7 days with different acclimation temperatures and periods. Error
bars indicate SE. Survival rates with the same capital letters do not
differ significantly between sexes (Fisher’s exact test, p [ 0.05).
Survival rates with the same lower case letters do not differ
significantly between non-acclimated flies and flies with acclimation
treatments within the same sex (Dunnett’s test, p [ 0.05)
Fig. 3 Time (days) (± 95 % fiducial limits) to kill 95 % of non-
acclimated insects of adult Bactrocera latifrons at different exposure
temperatures
while there was no difference between sexes when only
water was provided.
At 14 or 15 °C
After 30 and 60 days from transfer to each temperature,
more than 50 and 30 % of adults, respectively, survived at
both 14 and 15 °C, showing higher survival rates than that
at 27 °C (Fig. 4). After 90 days from transfer, 30 % of
females and 32 % of males survived at 15 °C, while 43 %
of females and 11 % of males survived at 14 °C.
Survival under FTRs
Under an FTR of 11 °C (22 h)/20 °C (2 h), survival of
female B. latifrons drastically increased compared to a
constant temperature of 12 °C, whereas the increase in
415
Fig. 4 Survival of Bactrocera latifrons at a constant temperature of
14, 15, or 27 °C
survival was slight in the males. Under this FTR, 67 and
13 % of female and male insects, respectively, survived
after 40 days from introduction (Fig. 5). Under an FTR of
11 °C (22 h)/14 °C (2 h), survival showed a similar pattern
to that at a constant temperature of 12 °C, in both females
and males, with 98 % mortality after 40 days (Fig. 5).
Under an FTR of 8 °C (22 h)/20 °C (2 h), 91 % of females
and 87 % of males died after 14 days, and 93 % of females
and 98 % of males died after 30 days. One female adult
survived more than 90 days under this FTR (Fig. 5).
Without sugar, survival of insects under the FTRs of
11 °C (22 h)/14 °C (2 h) and 11 °C (22 h)/20 °C (2 h)
decreased faster than when sugar was provided to the
insects. Under FTRs of 5 or 8 °C, survival showed a sim-
ilar tendency in insects provided with sugar and those
insects without sugar (Fig. 5).
Reproductive abilities of surviving females
The percentage of surviving females that laid fertile eggs
did not differ between the rearing temperature (27 °C,
control) and low temperatures, and between sugar-provided
and only water-provided females (at 8 °C for 3, 5, or
7 days) (Fisher’s exact test, p [ 0.05). Female survivors
after exposure to 12 °C for 35 days, 10 °C for 20 days,
8 °C for 7 days, or 90 days under FTRs of 11 °C (22 h)/
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Fig. 5 Survival of Bactrocera
latifrons in fluctuating thermal
regimes: 5 °C (22 h)/20 °C
(2 h), 8 °C (22 h)/20 °C (2 h),
11 °C (22 h)/20 °C (2 h), 11 °C
(22 h)/14 °C (2 h). The curves
shown in the figure indicate
logistic regression for the data
on survival of adult B. latifrons
at a constant 12 °C
20 °C (2 h) produced fertile eggs, whereas no fertile eggs
were produced by females after exposure to 8 °C for 8 or
more days (Table 2).
Average air temperatures
Table 1 shows average daily air temperatures at the six
sites. On Tanegashima, Yakushima and Nakanoshima
Islands, average daily maximum temperatures were
between 14 and 16 °C, when the average daily temperature
was 12 °C or lower.
Discussion
Bactrocera latifrons adults showed lower tolerance to cold
temperatures than other tropical fruit flies. When adults of
B. latifrons were maintained at 14 or 15 °C, they survived
for long periods of time (Fig. 4). At 12 °C or lower,
however, the survival rate decreased with decreasing
temperatures, from 38 days Ltime95 at 12 °C to 13 days
Ltime95 at 8 °C (Fig. 3). In other tropical fruit flies,
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Appl Entomol Zool (2014) 49:411–419
Koidsumi and Shibata (1964) showed that adults of B.
cucurbitae (Coquillett) and B. dorsalis (Hendel) could
survive more than 100 days at 9 °C.
The results showed that presence of sugar, food
resources, increased the survival times of B. latifrons at
10 °C or higher but not at 8 °C (Fig. 3). Koidsumi and
Shibata (1964) showed that B. cucurbitae and B. dorsalis
adults had extended survival times when water was pro-
vided, compared to insects without water at 10 °C, but such
extension did not occur at 8 °C or lower. They suggested
that both flies did not feed at temperatures of 8 °C or lower.
Although survival times of B. latifrons at 8 and 10 °C were
much shorter than those of B. cucurbitae and B. dorsalis
(Koidsumi and Shibata 1964), the critical temperature at
which feeding ceases appears to be common to the three
fruit flies.
The cold acclimation treatments did not enhance the
cold tolerance of adult B. latifrons (Fig. 2). In other trop-
ical insects such as Cylas formicarius (Fabricious), Thrips
palmi (Karny), Alphitobius diaperinus Panzer, and Bron-
tispa longissima (Gestro), acclimation to 10 or 15 °C (for
several days to 1 month) extended survival times in a
Appl Entomol Zool (2014) 49:411–419 417

Table 2 Percentages of Exposure Food (sugar) Exposure n Percentage of n Percentage of females

surviving Bactrocera latifrons temperature (with: ?, without: time (days) surviving which laid fertile eggs
females that laid fertile eggs (°C) -) females
after exposure to different low
temperatures 8 ? 3 30 80.0 8 37.5
- 3 30 73.3 14 7.1
? 5 40 62.5 9 44.4
- 5 40 80.0 12 41.7
? 7 40 50.0 10 60.0
- 7 60 46.7 7 14.3
- 8 30 6.7 2 0
- 10 30 6.7 1 0
? 15 40 7.5 2 0
10 - 4 30 76.7 2 50.0
- 7 40 77.5 13 38.5
- 10 30 43.3 4 25.0
- 12 30 10.0 3 33.3
? 15 30 33.3 8 50.0
? 20 40 45.0 10 20.0
12 - 5 20 75.0 10 30.0
- 7 30 60.0 13 15.4
- 10 30 43.3 9 55.6
- 12 30 3.3 1 0
? 25 30 50.0 10 30.0
? 30 30 50.0 1 0
? 35 40 10.0 4 50.0
11 (22 h)/20 ? 90 45 22.2 10 10.0
(2 h)
15 ? 7 –a – 15 40.0
a 27 ? 10–15 – – 25 36.0
not available

subsequent exposure to low temperatures of 0, 5, or 10 °C
(Kandori et al. 2006; McDonald et al. 2000; Renault et al.
2004; Takano et al. 2012). In this study, prior exposure to
15 or 17 °C negatively affected the survival times of B.
latifrons. Females reared at 20 °C showed similar survival
rates as those reared at 27 °C, while in males, those reared
at 20 °C showed significantly lower survival rates than
those reared at 27 °C. Although B. latifrons adults can
survive for long periods of time at 14 or 15 °C, females
may accumulate ‘‘chill injury’’ at 17 °C or lower, and
males at 20 °C or lower. This appears to be consistent with
the results of the experiments on survival under FTRs,
which demonstrate that females recover from chill injury
sustained at 11 °C following 2 h at 20 °C, but males only
showed slight recovery (see the following paragraph).
Under an FTR regime of 11 °C (22 h)/20 °C (2 h),
survival of female B. latifrons drastically increased com-
pared to that at a constant temperature of 12 °C. Consid-
ering that the average temperature in one cycle of the FTR
(24 h) is 11.8 °C, lower than 12 °C, the FTR may strongly
contribute to the extend survival time of B. latifrons
females. On the other hand, under an FTR of 11 °C (22 h)/
14 °C (2 h), survival showed a similar tendency to that at a
constant temperature of 12 °C. Thus, the interruption
temperature, 14 or 20 °C, is considered to be very critical
in female B. latifrons for increasing survival under FTRs.
Renault et al. (2004) showed similar results in the tropical
beetle Alphitobius diaperinus Panzer, i.e., survival
increased slightly under FTRs 5 °C (22 h)/10 °C (2 h) and
5 °C (22 h)/15 °C (2 h), but it markedly improved at 5 °C/
20 °C or higher. In males of B. latifrons, survival showed
only a slight increase under an FTR of 11 °C (22 h)/20 °C
(2 h). Interruption temperatures above 20 °C were not
investigated in this study; therefore, it is not known whe-
ther B. latifrons males have markedly increased survival
with higher interruption temperatures.
Sex-specific differences in response to cold tempera-
tures were demonstrated in this study. Female B. latifrons
showed significantly greater Ltime95, fewer negative
effects following prior exposure to 15, 17, or 20 °C (cold
acclimation), and a greater chance of survival under FTRs
than males. These results indicate that females are more

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tolerant of cold temperatures than males. Although the
opposite trend is reported in Drosophila melanogaster
Meigen, where male cold resistance is higher than that of
females (Jensen et al. 2007; Kelty and Lee 2001; Sejerkilde
et al. 2003), the mechanisms underlying the differences
between the two sexes are still unclear. Female survivors of
B. latifrons after exposure to 12 °C for 35 days or to FTRs
of 11 °C (22 h)/20 °C (2 h) for 90 days could produce
fertile eggs. In the case of FTRs, female flies produced
fertile eggs without copulation after exposure because all
males died before the end of the 90-day exposure. This
may indicate that females alone can overwinter and
reproduce without mating in the spring.
On Okinoerabu and Amami-Oshima Islands, the average
daily temperature was always equal to or higher than
14 °C. More than 30 % of B. latifrons females survived
longer than 90 days at 14 °C suggesting that they could
probably overwinter in these locations. On the other hand,
on Tanegashima, Yakushima, and Nakanoshima Islands,
there were more than 40 days where the average daily
temperature was 12 °C or lower (average daily maximum
temperatures were 14–16 °C). Therefore, it seems unlikely
that B. latifrons will overwinter on these islands because
the Ltime95 at 12 °C was estimated to be 38 days, and
under an FTR of 11 °C (22 h)/14 °C (2 h) the mortality
after 40 days was 98 % (Fig. 5). When the effect of sun-
light on body temperature is considered, however, we
cannot rule out the possibility of overwintering of B. lati-
frons on Tanegashima, Yakushima and Nakanoshima
Islands. The body temperature of insects including small
dipteran species may rise in sunlight and it may become
10 °C higher than that of the surrounding air (Akiyama and
Nishida 2013; Digby 1955; Parry 1951; Willmer and
Unwin 1981). Although the winter behavior of B. latifrons
is unknown, dipteran species often bask in the sun in
winter, suggesting that B. latifrons adults may also raise
their body temperature by basking. If basking occurs, they
may overwinter on these islands, because 67 % of females
survived for 40 days under an FTR of 11 °C (22 h)/20 °C
(2 h) (Fig. 5). In Ibusuki, the probability of overwintering
is low, even if diurnal fluctuations and the effect of sunlight
are considered. There were 15 days when the average daily
temperature was 8 °C. Although a few females survived
more than 90 days under an FTR of 8 °C (22 h)/20 °C
(2 h), most of them (91 %) died before 15 days (Fig. 5).
Global warming is causing poleward range expansion in
many insects and further warming trends are predicted to
occur in the future (Parmesan and Yohe 2003; Root et al.
2003). In Japan, the average temperature in the winter on
the east coast of Kyushu has been predicted to rise by 3 °C
by 2100 (Japan Meteorological Agency 2013b). In this
scenario, the average winter temperature on Tanegashima
would never drop below 14 °C, indicating a high
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Appl Entomol Zool (2014) 49:411–419
probability of overwintering, and on Ibusuki it would never
drop below 11 °C, indicating conditional probability of
overwintering.
Acknowledgments I thank Kenji Ohto for his technical suggestions
on this study. I thank Toshihisa Kamiji, Ryoko T. Ichiki, and Mika
Murata for their technical assistance. I thank Quint Newcomer for
helpful comments on the manuscript. I am grateful to members of the
Research Division of Yokohama Plant Protection Station for their
kind assistance and members of the Naha Plant Protection Station for
providing the insects.
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