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Larvae of the firefly Pyrocoelia pectoralis (Coleoptera: Lampyridae) as possible
biological agents to control the land snail Bradybaena ravida
Xinhua Fu
a,
⇑
, V. Benno Meyer-Rochow
b,c
a
Hubei Insect Resources Utilization and Sustainable Pest Management Key Laboratory, College of Plant Science and Technology, Huazhong Agricultural University, Wuhan 430070,
Hubei, China
b
Jacobs University Bremen, Faculty of Engineering & Sciences, D-28725 Bremen, Germany
c
Department of Biology, University of Oulu, SF-90014 Oulu, Finland
highlights
"
The firefly Pyrocoelia pectoralis was
successfully bred.
"
There are two types of larvae occur
after the third instar.
"
Environmental conditions affected
larval development and were
studied.
"
The effectiveness as a predator on
the pest snail Bradybaena ravida was
assessed.
"
Use as a biological control agent is
discussed.
graphical abstract
.
article info
Article history:
Received 25 July 2012
Accepted 15 February 2013
Available online 28 February 2013
Keywords:
Development
Feeding capacity
Land snails
Firefly
Pyrocoelia pectoralis
abstract
Rearing experiments with the firefly Pyrocoelia pectoralis demonstrated that the species can be success-
fully bred under laboratory and field conditions and that there are two types of larva: overwintering and
non-overwintering. Comparisons showed that the differentiation between the two larval types occurred
after the third larval stage. In the field, non-overwintering larvae pupate in September, emerge in October
and produce a second annual generation, while overwintering larvae begin to grow more slowly from the
3rd instar onward, then overwinter to ultimately reach a larger size than the non-overwintering larvae
and to pupate in September. Adults emerge in October. Larval development at 15, 20, 25, 30 and 35 °C
was investigated under a photoperiod of L:D= 12:12. At 15 °C all larvae died as 4th instars, but from
20 °Cto30°C larval phases became increasingly shorter, while at 35 °C they lengthened again. Larval
feeding capacity increased with higher temperature up to 30 °C, but decreased at 35 °C. Under three pho-
toperiods, i.e., L:D= 16:8, 12:12 and 8:16 at 25 °C, the larval period was shortest under L:Dof 16:8 and
longest under L:D= 8:16. Feeding capacity of the larvae exhibited a positive correlation with the duration
of the dark period. Larvae under longer periods of illumination pupated considerably earlier than those
kept one under shorter periods of light exposure. No significant differences in the numbers of overwin-
tering larvae were found in connection with different temperatures and photoperiods.
Ó2013 Elsevier Inc. All rights reserved.
1049-9644/$ - see front matter Ó2013 Elsevier Inc. All rights reserved.
http://dx.doi.org/10.1016/j.biocontrol.2013.02.005
⇑
Corresponding author.
E-mail address: fireflyfxh@mail.hzau.edu.cn (X. Fu).
Biological Control 65 (2013) 176–183
Contents lists available at SciVerse ScienceDirect
Biological Control
journal homepage: www.elsevier.com/locate/ybcon
1. Introduction
Terrestrial snails are known to be destructive agricultural pests,
capable of causing severe damage to vegetables, ornamentals and
other plants. Pest snail attacks of plant seeds, seedlings, under-
ground tubers, leaves and fruits can lead to major losses (Baker,
1989; Barker, 2002, 2004). The terrestrial species Bradybaena rav-
ida (Pfeiffer, 1850) is a common land snail widely distributed
throughout China, Japan, Korea and Russia (Chen, 2004; Wang,
2008). Recently numerous cases of damage to vegetables, peaches,
grapes, and corn were attributed to this species. In 2006 over 3000
hectares of corn were damaged by it in Qi County, Kaifeng City, He-
nan Province alone (Wang, 2008; Liu and Wu, 2011).
Fu and Meyer-Rochow (2012) emphasized that larval aquatic
fireflies have the potential of serving as biological control agents
in the fight against disease transmitting freshwater snails. There-
fore, it is now relevant to investigate if terrestrial snail-consuming
firefly larvae might not also be usable in controlling pest land
snails. Support for this notion comes from observations on terres-
trial fireflies by Kaufmann (1965), Buschman (1984, 1988), Barker
(2004) and Wang et al. (2007), which state that their larvae con-
sume primarily land snails and therefore can be considered as
the snails’ natural enemies. Wang et al. (2007) reported that specif-
ically larval Pyrocoelia pectoralis, which distributed in Hubei Prov-
ince feed on the snail B. ravida and indicated that the larvae might
be a possible biological agent to control the snail B. ravida. Tests to-
wards this end were, however, never conducted.
Depending on the species of firefly, some have one, others have
two or even three generations per year (Balduf, 1935; Schwalb,
1961; Naisse 1966; Wootten, 1976; Fu et al., 2006; Ho et al.,
2010). To obtain a sufficient number of larvae to be of potential
use as biological agents to control the pest snail, a short larval per-
iod and successful larval development are key factors. The larval
developments of several Old World lampyrids have been de-
scribed, e.g., Lampyris noctiluca L. (Balduf, 1935; Schwalb, 1961;
Naisse, 1966; Wootten, 1976), Lamprohiza splendidula (LeConte)
(Balduf, 1935; Schwalb, 1961), Lamprigera tenebrosus (Walker)
(Bess, 1956), Luciola discollis Laporte (Kaufmann, 1965), Luciola cru-
ciate Motschulsky (Minami, 1961; Yuma, 1981), Aquatica leii Fu and
Ballantyne (Fu et al., 2006). The firefly Pyractomena lucifera (Melsc-
heimer) was reared successfully under laboratory and field condi-
tions (Buschman, 1988) and Ho et al. (2010) bred the aquatic firefly
A. ficta at temperatures ranging from 18 °Cto30°C. However, very
few studies other than that by Buschman (1988) focus on quanti-
tative assessments of larval feeding capacity during larval
development.
The firefly P. pectoralis was chosen, because it is a common local
firefly and known to be an effective predator of the pest land snail
B. ravida. Our investigation provides details on the larval develop-
ment of a Chinese terrestrial lampyrid maintained under labora-
tory conditions and explores larval feeding capacity under
different conditions of photoperiod and environmental
temperature.
2. Materials and methods
2.1. Egg treatments
Wang et al. (2007) had reported that eggs of P. pectoralis enter
diapause to overwinter and hatch in April the next year. We con-
firmed that the egg period lasts for about six months. In order to
accelerate hatching of the eggs and to obtain newly-hatched first
instar larvae for some observations on growth and development,
we exposed the eggs to a temperature of 6 ± 1 °C for 40 days. Addi-
tional small larvae, measuring no more than 10 mm in length, were
collected in May, 2010 from Xianjian Village, Hongshan District,
Wuhan City (Hubei Province), revealing themselves to the human
collector by their own glow. Head widths, body lengths and body
widths of the collected larvae were measured and compared with
larval developmental parameters available from the earlier growth
experiments, demonstrating that the collected larvae represented
2nd instars. Twenty larvae per transparent plastic box
(30 cm 20 cm 5 cm) were maintained on moist filter paper
with living specimens (usually 5) of the land snail B. ravida added
as prey.
After eclosion, males and females were allowed to mate in
the plastic box, where the females laid their eggs on the moist
filter paper available to them. Three days later, the eggs were re-
moved with the aid of a moist Chinese brush pen and placed
into a similar plastic box lined also with moist paper. For
40 days the box was kept at a temperature of 6 ± 1 °C under nat-
ural photoperiod. Then the eggs were transferred to a climate
chamber set at 25 °C, a photoperiod of 12:12 (L:D) and humidity
of 75% for hatching.
2.2. Confirmation of and comparisons between the two types of larva
It was reported that both larval P. pectoralis and their eggs over-
wintered from November to next April (Wang et al., 2007). Thus,
there would be two types of larva in the wild, namely overwinter-
ing larvae and non-overwintering larvae, the latter appearing from
May to September. To explore how the two types of larva might
arise, three groups of 50 newly-hatched larvae reared in the labo-
ratory at a temperature of 25 °C, photoperiod of 12:12 L:Dand 75%
humidity were used. Larval duration and feeding capacity (fresh
body mass of land snails) at each larval stage were recorded. Body
length, body width, head width, and body mass of each instar were
measured. Because both overwintering as well as non-overwinter-
ing larvae could pupate as 4th, 5th and 6th instar larvae, we com-
pared only larvae of the two types when they pupated as 6th
instars.
2.3. Larval development and predation on land snails
2.3.1. Impact of temperature on larval development and predation on
land snails
Newly-hatched larvae were reared at 15, 20, 25, 30, and 35 °Cin
an artificial climate chamber (SPX-250IC: Shanghai Boxun Corpo-
ration), with relative humidity of 70 ± 10% and 12:12 L:Dphotope-
riod. Each treatment contained 45 larvae in a fully randomized
setup. Each larva was reared in a transparent plastic box
(7 cm 7cm6 cm) with moist filter paper covering bottom
and sides. The filter paper was replaced every two days. The fol-
lowing parameters were evaluated: number of instars; duration
spent as instars at each of the larval periods; fresh body mass of
snails before and of empty shells after predation; survival at differ-
ent larval stages; body length, body width, head width, and body
mass of every larva at each larval instar; number of overwintering
larvae and number of larvae turning into pupae. Vernier calipers
were used in morphometric assessments and an analytical balance
(AUY120, Shimadzu, Japan) was used for weighing the snails before
and after feeding to determine feeding capacity. With 1st to 3rd in-
stars, we used a small piece of snail meat put on a square plastic
sheet to feed them. As usual plastic sheet and the food on it were
weighed before and after feeding.
2.3.2. Impact of photoperiod on larval development and predation on
land snails
Wang et al. (2007) reported that the optimum temperature for
larval activity of P. pectoralis was 18–30 °C. Thus, the newly-
hatched larvae were reared at 25 °C in an artificial climate chamber
X. Fu, V. Benno Meyer-Rochow / Biological Control 65 (2013) 176–183 177
(SPX-250IC: Shanghai Boxun Corporation), with relative humidity
of 70 ± 10% and photoperiods of 8:16, 12:12 and 16:8 (L:D). Each
treatment involved 45 larvae in a fully randomized setup. Rearing
methods were same as above. The same parameters as above were
monitored and recorded.
2.3.3. Larval development and predation on land snails under natural
conditions
Fifty newly-hatched larvae were reared under natural condi-
tions. Each larva was kept in a transparent plastic box
(7 cm 7cm6 cm) lined with moist filter paper. The boxes were
Fig. 1. Comparison between development of non-overwintering and overwintering larvae (mean ± SD, Significant differences between wintering and non-wintering larval
development and feeding capacity are labelled with different letters and based on Duncan’s multiple comparison test).
178 X. Fu, V. Benno Meyer-Rochow / Biological Control 65 (2013) 176–183
placed into the high 15–20 cm tall grass outside the lab. Rearing
methods and parameters recorded were the same as above.
2.4. Statistical analysis
Significant differences between wintering and non-wintering
larval development, determined by using Duncan’s multiple com-
parison test at A= 0.05, affected body length, body width, body
weight, diapause duration and feeding capacity. To compare feed-
ing capacity and larval duration under different temperatures or
photoperiods, One-Way analyses of variance (ANOVA) were used;
means were then compared by ANOVA, followed by Duncan’s test
at A= 0.05. To compare the number of larvae pupating at different
instars and overwintering under different photoperiods at a tem-
perature of 25 °C, the Kruskal–Wallis test was used. All analyses
were done with SAS software (The SAS System for Windows V8).
3. Results
3.1. Confirmation of and comparisons between the two types of larva
Under rearing conditions of 25 °C and L:D= 12:12, 130 larvae
out of a total of 150 individuals survived in three groups of larvae
surveyed. 114 larvae pupated in the same year and enclosed, while
16 larvae overwintered. Four subsequently died before pupation,
but 12 eventually yielded adults the following year. Thus, 10.7%
of the larvae stopped to develop beyond the third instar in the first
year and overwintered to eventually mature the following year. No
significant differences were found between non-overwintering and
overwintering larvae from first to third instar with regard to feed-
ing capacity, body length, body width, and body mass (Fig. 1;Ta-
bles 1 and 2) and predictions, which of the larvae might winter
over were not possible. Yet, from fourth instar onward, overwinter-
ing larvae grew larger than their non-overwintering counterparts
and outperformed them in feeding capacity, body length, body
width, and body mass (p< 0.01) (Fig. 1;Tables 1 and 2). The dura-
tions of 117.76 ± 12.33, 104.32 ± 11.79 and 99.34 ± 10.10 days
spent as fourth, fifth and sixth instar larvae, respectively, were
much longer in the overwintering than in the non-overwintering
types, in which the respective larval instar durations amounted
to no more than 25.00 ± 3.83, 41.06 ± 4.57 and 52.00 ± 5.40 days
(Fig. 1;Tables 1 and 2). Amongst the overwintering larvae, only
two larvae pupated as 5th instars, while the remaining ten pupated
as 6th instar larvae. However, non-overwintering larvae also pu-
pated as 4th instar larvae in addition to those pupating as 5th
and 6th instars.
3.2. Larval development and predation on land snails
3.2.1. Impact of temperature on larval development and predation
activity
Under a photoperiod of 12:12 (L:D) environmental temperature
affected larval development and feeding capacity. At 15 °C, larval
mortality was highest with all individuals dying as 3rd instar indi-
viduals. However kept at 20 °C, 25 °C and 30 °C, larvae survived
and showed a tendency to have shorter larval periods with increas-
ing temperatures. Larval periods of individuals kept at 25 °C and
30 °C were shorter than those recorded in larvae kept at 20 °C
(p< 0.05, df = 130) (Fig. 2A), but when reared at 35 °C larval periods
were longer than those kept at 20 °C, 25 °C and 30 °C(Fig. 2A). Lar-
val feeding capacity also increased with increasing temperature
from 20 °C via 25 °Cto30°C, but thereafter at 35 °C decreased
slightly (Fig. 2B).
Although highest at 30 °C of the four temperatures studied, lar-
val feeding capacity showed no statistically significant difference
to that seen at 25 °C(p< 0.01, df = 130) (Fig. 2). With the exception
of 15 °C at which no larvae survived beyond stage three instars, lar-
vae turned into pupae as 4th, 5th, and 6th instars at the other four
temperatures (Table 3). At 35 °C, 33 larvae (82.3%) pupated as 4th
instars, which represented the largest percentage and was statisti-
cally significantly different from the pupal percentages seen at the
other temperatures (p< 0.01, df = 144) (Table 3). At 20 °C, 12 lar-
vae, representing a percentage of 41.4%, pupated as 6th instars (Ta-
ble 3). Of the 5 levels of temperature tested, larval survival was
highest, i.e. 93.3%, at 35 °C. With respective percentages of 16.7%
and 4.8% overwintering larvae were most common at 30 °C and
least common at 35 °C, but the difference was not statistically sig-
nificant (Table 3).
3.2.2. Impact of photoperiod on larval development and predation on
land snails
Reared at a constant temperature of 25 °C, different photoperi-
ods affected larval development and food consumption. Larvae un-
der a photoperiod of 16:8 (L:D) grew faster than those reared under
12:12 and 8:16 (p< 0.01, df = 100) (Fig. 3A). Yet at a photoperiod of
8:16, because of the longer period spent in the larval stages, the
larvae oversall consumed the largest number of snails, while at
16:8 consumption was least (Fig. 3B).
Table 1
Development of non-overwintering larvae under 25 °C and 12:12 (L:D) photoperiod (mean ± SD).
Instar Body length (mm) Body width (mm) Head width (mm) Body mass (g) Feeding capacity (g) Period (d)
1st 7.17 ± 0.11 1.07 ± 0.01 0.77 ± 0.03 0.005 ± 0.001 0.060 ± 0.003 20.4 ± 1.7
2nd 12.27 ± 1.22 2.14 ± 0.05 1.00 ± 0.07 0.009 ± 0.001 0.266 ± 0.018 20.9 ± 2.6
3rd 16.66 ± 1.41 2.94 ± 0.09 1.13 ± 0.02 0.031 ± 0.003 0.276 ± 0.031 18.9 ± 2.2
4th 23.01 ± 1.56 3.98 ± 0.10 1.21 ± 0.02 0.097 ± 0.015 0.574 ± 0.061 25.0 ± 3.8
5th 29.17 ± 1.60 5.20 ± 0.21 1.38 ± 0.03 0.172 ± 0.016 1.168 ± 0.167 41.1 ± 4.6
6th 33.92 ± 2.61 6.20 ± 0.31 1.44 ± 0.05 0.299 ± 0.025 1.478 ± 0.279 52.0 ± 5.4
Table 2
Development of overwintering larvae under 25 °C and 12:12 (L:D) photoperiod (mean ± SD).
Instar Body length (mm) Body width (mm) Head width (mm) Body mass (g) Feeding capacity (g) Period (d)
1st 7.19 ± 0.11 1.07 ± 0.01 0.77 ± 0.02 0.005 ± 0.001 0.060 ± 0.002 21.5 ± 1.8
2nd 11.89 ± 1.41 2.13 ± 0.06 1.15 ± 0.04 0.009 ± 0.001 0.234 ± 0.015 21. 6 ± 1.9
3rd 14.96 ± 1.67 2.61 ± 0.07 1.27 ± 0.07 0.028 ± 0.002 0.389 ± 0.043 38.3 ± 4.7
4th 29.76 ± 1.58 4.88 ± 0.10 1.36 ± 0.05 0.199 ± 0.023 1.129 ± 0.133 117.8 ± 12.3
5th 41.59 ± 2.95 7.20 ± 0.43 1.49 ± 0.08 0.435 ± 0.039 1.542 ± 0.367 104.3 ± 11.8
6th 52.34 ± 5.36 8.59 ± 0.57 1.62 ± 0.09 0.582 ± 0.058 2.449 ± 0.598 99.3 ± 10.1
X. Fu, V. Benno Meyer-Rochow / Biological Control 65 (2013) 176–183 179
Under all the three photoperiods, larvae were able to pupate
as 4th, 5th or 6th instars (Table 4). The largest number of over-
wintering larvae, i.e. 13.2% of the total, occurred in connection
with a photoperiod of 8:16 (L:D), but the figure was not statisti-
cally significantly different from the percentages of overwinter-
ing larvae under the other two photoperiods. Highest larval
survival rate (88.9%) occurred under a photoperiod of 16:8
(Table 3).
3.2.3. Larval development and predation on land snails under natural
conditions
Under field rearing experiments, 33 out of 50 larvae survived
and pupated. The period spent as larvae was 114.27 ± 16.74d
(mean ± SD) and thus similar to larval development in the lab at
30 °C with an L:Dof 12:12; feeding capacity reached 1.76 ± 0.19 g
(mean ± SD), a value close to that seen in lab-reared individuals
at 25 °C and an L:Dof 16:8. Under natural conditions most larvae
Fig. 2. Feeding capacity and larval duration under different temperatures (mean ± SD. Based on One-way analysis of variance (ANOVA) with means compared by ANOVA
followed by Duncan’s test at A= 0.05. Significant differences between the mean data are labelled with different letters).
Table 3
Numbers of larvae pupating at different instars and overwintering under different temperatures.
Temperatures 15 °C20°C25°C30°C35°C
Number of non-overwintering larvae 0 25 32 30 40
Number of non-overwintering larvae pupating at 4th instar – 12 a 12 a 17 a 33 b
Number of non-overwintering larvae pupating at 5th instar – 1 a 18 b 11 b 7 a
Number of non-overwintering larvae pupating at 6th instar – 12 a 2 b 2 b 0 b
Number of overwintering larvae – 2a 4a 6 a 2 a
Percent of overwintering larvae 0 6.9% 11.1% 16.7% 4.8%
Total survival larvae 0 29 36 36 42
Larval survival rate 0 64.4% 80% 80% 93.3%
– Larvae died.
180 X. Fu, V. Benno Meyer-Rochow / Biological Control 65 (2013) 176–183
pupated as 4th instars (n= 19), 10 as 5th instars, 4 as 6th instars;
and 6 (i.e., 12%) individuals overwintered. This result resembled
what was seen in larvae developing at 25 °C under an L:Dcycle
of 16:8 or at 30 °C and an L:Dof 12:12 (Table 5).
4. Discussion
In the firefly P. lucifera, laboratory and field-rearing experiments
showed that some larvae, which hatch in spring, can complete
development by August the same year and thus produce a second
generation. However, body mass distribution in field-collected lar-
vae indicated that most of the larvae did not develop this rapidly
and spend at least one winter as larvae. Most of these overwinter-
ing larvae mature the next year. Larval development was con-
trolled by photoperiodic cues, which inhibited the development
of the larvae into the pupal stage in autum (Buschman, 1988).
The flexibility in the development of P. lucifera was interpreted
by Buschman (1984) as an important factor contributing to the
wide geographic distribution of this insect. A similar situation oc-
curs in the Japanese species L. cruciata (Yuma, 1981). In this case,
some of the overwintering larvae did not complete development
in the spring. This means that some larvae take 2 years to complete
development (Yuma, 1981). Ho et al. (2010) reported that food
shortages and drops in temperature can lead to increased moulting
in A. ficta and reduce larval growth.
Our results on P. pectoralis, under conditions of sufficient food,
show that the numbers of overwintering and non-overwintering
larvae were not statistically significantly altered by either different
temperatures or photoperiods or combinations of the two. It was
observed, however, that larvae at 20 °C moulted more frequently
(although not reaching statistical significance) than at the other
temperatures tested, thus showing a similar tendency to that of
the aquatic firefly A. ficta (Ho et al., 2010).
In order to use P. pectoralis larvae as biological agents to control
pest snails, larvae released into the field should remain in the larval
stages as long as possible and be able to consume large numbers of
snails. For mass rearing of larvae under laboratory conditions the
objectives are different: a maximum number of larvae (to be re-
leased into the field) should be reared in the shortest possible time
with as little feeding as possible. Thus, which combinations of tem-
perature and photoperiod can we recommend to achieve these
goals?
Fig. 3. Feeding capacity and larval duration under different photoperiods (mean ± SD. Based on One-way analysis of variance (ANOVA) with means compared by ANOVA
followed by Duncan’s test at A= 0.05. Significant differences between the mean data are labelled with different letters).
X. Fu, V. Benno Meyer-Rochow / Biological Control 65 (2013) 176–183 181
We found that under the identical photoperiod of 12:12 (L:D),
larval survival rate at 35 °C was the highest, exceeding even that
of all temperatures and the duration spent as a larva was the lon-
gest at the five levels of temperature tested. At 30 °C, however, lar-
val life span was the shortest amongst the five levels of
temperature tested. Considering mass rearing of fireflies and their
larvae, a temperature of 35 °C must be seen as superior to that of
30 °C in order to achieve the highest larval survival rate at a pho-
toperiod of 12:12 L:D, but a temperature of 25 °C, but with differ-
ent photoperiods, larval survival rate in combination with a
photoperiod of 16:8 (L:D) was highest, larval duration shortest,
and feeding capacity lowest amongst the three levels of photope-
riod tested. Therefore, a photoperiod of 16:8 (L:D) at a temperature
of 25 °C seems the most suitable photoperiod for laboratory-based
mass rearings of P. pectoralis.
The largest percentages of overwintering larvae in P. pectoralis
were seen to occur in connection with a temperature of 30 °C
and photoperiods with longer dark phases, but based on our data,
the percentages of overwintering larvae under the different regi-
mens of temperature and photoperiod did not reach statistical sig-
nificance. This suggests that diapause in the overwintering larvae
is not solely controlled by temperature and photoperiod alone,
but that other factors could play a role. P. pectoralis adults emerge
in October and courtship is easily affected by bad weather such as
strong winds and cold rain, both of which frequently occur in the
Wuhan area. The proportion of about 10% of the overwintering lar-
vae in P. pectoralis (recorded by us in the field, cf. Section 3.3.3) can
probably compensate for fecundity losses due to adverse summer
conditions. Since Ho et al. (2010) reported that in the species A. fic-
ta not only a drop in environmental temperature but also food
shortages can lead to increased moulting and reduce larval growth,
we need to consider this possibility for P. pectoralis as well.
Thus the strategy of P. pectoralis to allow both eggs and larvae to
overwinter together, appears to safeguard the survival of the pop-
ulation during times of environmental stress. The exact trigger,
however, of what controls the balance between wintering and
non-wintering larvae and the change in development after the
third instar is still not clear as temperature and photoperiod alone
or in combination with each other fail to change the balance be-
tween the two kinds of larva in a statistically significant way.
Although using firefly larvae for the biological control of land
snails is not a new idea, e.g., L. noctiluca L. was earlier imported
to New Zealand from the U.K. for the biological control of Helix as-
persa Mull. (Clausen, 1940), this measure has not received wide-
spread recognition and acceptance. Releasing larval fireflies into
greenhouses and gardens to control snails remains an alternative
to more conventional control methods; besides it has the potential
of attracting people to watch fireflies and their bioluminescent dis-
plays and could thus become a sightseeing attraction. However,
once non-indigenous firefly larvae were to be released into regions
outside their own distribution, some problems could arise: the
introduced firefly larvae could become a threat to some endan-
gered snail species or could compete with locally present species
of fireflies. Research not only will have to address these questions,
but additionally will have to focus on factors affecting the balance
between wintering and non-wintering larvae. Finally sexual com-
petition and choice between wintering and non-wintering adults
will need to be looked at it in order to fully understand the survival
strategies of P. pectoralis.
Acknowledgments
We thank Li Fang, Quan Liu, Tengfei Zhu, Xianying Ji, Xiaoqing
Zhang, Jiali Liu for collecting and helping to rear the insects. This
research was supported by the Chinese National Science Founda-
tion (#30900147 and #31172137) and Fundamental Research
Funds for the Central Universities (Program No. 2011PY054).
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Table 4
Number of larvae pupating at different instars and overwintering under different photoperiods at a temperature of 25 °C (mean data were analyzed by
Kruskal–Wallis test and significant differences between the mean data are labelled with different letters).
Photoperiods (L:D) 16:8 12:12 8:16
Number of non-overwintering larvae 36 32 33
Number of non-overwintering larvae pupating as 4th instars 19 a 12 a 5 b
Number of non-overwintering larvae pupating as 5th instars 14 a 18 a 19 a
Number of non-overwintering larvae pupating as 6th instars 3 a 2 a 9 b
Number of overwintering larvae 4 a 4 a 5 a
Percent of larvae overwintering 10% 11.1% 13.2%
Total surviving larvae 40 36 38
Larval survival rate 88.9% 80% 84.4%
Table 5
Larval development under natural conditions (mean ± SD).
Instar nBody length (mm) Body width (mm) Head width (mm) Body mass (g) Feeding capacity (g) period (d)
1st 33 7.13 ± 0.14 1.05 ± 0.01 0.77 ± 0.02 0.004 ± 0.001 0.060 ± 0.003 22.03 ± 1.40
2nd 33 10.36 ± 0.46 1.87 ± 0.19 0.87 ± 0.07 0.008 ± 0.001 0.239 ± 0.034 23.21 ± 2.58
3rd 33 16.37 ± 0.46 2.93 ± 0.07 1.08 ± 0.07 0.031 ± 0.003 0.281 ± 0.030 18.55 ± 1.28
4th 33 22.97 ± 0.42 3.97 ± 0.11 1.08 ± 0.04 0.095 ± 0.015 0.570 ± 0.046 24.79 ± 2.20
5th 14 27.41 ± 3.28 4.91 ± 0.44 1.22 ± 0.10 0.154 ± 0.022 1.032 ± 0.156 45.36 ± 5.42
6th 4 33.85 ± 1.29 6.19 ± 0.23 1.44 ± 0.06 0.298 ± 0.022 1.424 ± 0.212 53.00 ± 3.83
182 X. Fu, V. Benno Meyer-Rochow / Biological Control 65 (2013) 176–183
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