Forensic Science International 172 (2007) 94–97

www.elsevier.com/locate/forsciint

Flight activity of the blowflies, Calliphora vomitoria and

Lucilia sericata, in the dark
J. Wooldridge, L. Scrase, R. Wall *
School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, UK
Received 14 April 2006; received in revised form 7 December 2006; accepted 20 December 2006
Available online 30 January 2007

Abstract
Many species of insects are able to fly at night or in very low light intensities. The question of whether calliphorid blowflies are also able to do
this to locate a corpse and oviposit nocturnally is of considerable forensic importance. However, to date studies of this behaviour have been
contradictory. Here, the activity and number of Calliphora vomitoria L. and Lucilia sericata (Meigen) (Diptera: Calliphoridae) caught on sticky-
traps were examined in a slow-speed wind tunnel, at different intensities of artificial light. The traps were either unbaited or baited-with liver. The
number of both species caught, decreased incrementally as light intensity was reduced. While the responses of the two species were broadly similar,
L. sericata were significantly more active than C. vomitoria, especially at higher light intensities. The number of flies of both species that were
caught was higher in the presence of liver bait, but the presence of the liver bait did not change the shape of the relationship between catch and light
intensity. Hence, light intensity acts as an independent exogenous stimulus for activity and although liver volatiles increase activity levels, they are
not necessary as an activation stimulus. Comparison of the numbers caught in small or large enclosures suggests that any flies caught in darkness
probably alighted on the trap by chance and that in darkness, while flies may be activated by carrion odours, they do not appear to be able to
navigate effectively to the source of that odour. The results presented here suggest that in darkness, the probability of oriented flight leading to
oviposition on a corpse by either species, is relatively low.
# 2007 Elsevier Ireland Ltd. All rights reserved.

Keywords: Activity; Dark; Diptera; Forensic entomology; Post-mortem interval

1. Introduction oviposition was observed by both C. vicina and Lucilia sericata

In forensic entomology, when the age of insect larvae is used
to estimate the minimum post-mortem interval (PMI), knowl-
edge of the factors that are likely to delay the time between death
and oviposition will clearly be of critical importance. One of the
key questions relates to the effects of periods of darkness. Most
blowflies are usually considered to be largely diurnal and
relatively inactive at night [1]. Hence a body left exposed at night
would not attract flies until the next morning [2,3]. However, a
number of anecdotal observations suggest that nocturnal
oviposition may occur, particularly with species of Calliphora.
For example, Calliphora vicina R.-D. were observed laying eggs
in a slaughterhouse at night [4] and on a cadaver in a dark cave
[5]. However, experimental studies which have attempted to test
these observations have given contradictory results. Nocturnal
(Meigen) on carrion both placed directly on the ground [6] and on
carrion raised off the ground to ensure that only flying insects
were likely to reach it [7]. However, studies in the dark and
artificially illuminated areas during the night found no evidence
of nocturnal oviposition [8], neither did 2-year study of carrion
colonisation in Indiana, USA [9]. Oviposition was found to be
synchronised with the onset of the photophase following a period
of darkness [3].
Clearly whether or not flies are active at night is an important
factor which may significantly affect the estimate of PMI. The
aim of the present work therefore was to undertake a
comparative investigation of the question of whether blowflies
are active in darkness, using two species of forensic
importance, Calliphora vomitoria L. and L. sericata.
2. Methods

* Corresponding author. Tel.: +44 117 928 9182. Fly behaviour was investigated in the laboratory using a slow-speed wind
E-mail address: richard.wall@bristol.ac.uk (R. Wall). tunnel, constructed of aluminium with clear reinforced-glass sides and ceiling

0379-0738/$ – see front matter # 2007 Elsevier Ireland Ltd. All rights reserved.
doi:10.1016/j.forsciint.2006.12.011
 J. Wooldridge et al. / Forensic Science International 172 (2007) 94–97 95

[10]. The wind-tunnel had a working section length of 2 m, width of 1.3 m and
height of 1 m and was illuminated by one 1.5 m, 32 kHz, diffused fluorescent
light. Air was drawn into the wind tunnel through a vertical bed of activated
charcoal by a centrifugal fan and blown upwards into a secondary horizontal
bed of activated charcoal filter. The airflow was then rotated through 908 by a
series of aluminium vanes and straightened by passing through two sheets of
steel mesh before entering the working section of the wind tunnel. Air exited the
wind tunnel via a sheet of 2 mm diameter nylon mesh covering the downwind
end of the working section. The wind tunnel was maintained in a small room at
25 8C (range 1 8C).
Larval C. vomitoria and L. sericata to be used in experimental trials were
obtained from a local fishing-bait shop and kept in the laboratory to pupate. The
adults that emerged were maintained in a cage (30 cm  30 cm  30 cm)
supplied with granulated sucrose and water ad libitum, and at 24–25 8C and
under a 16 h light:8 h dark cycle. Female adults only were used in experiments,
when 6 days-of-age. Prior to use they were briefly cooled, to allow then to be
counted and sexed.
Sticky-traps were constructed from a square of cardboard (10 cm  10 cm)
covered by a white plastic sheet coated with polybutene-based, high-tack, non-
setting adhesive (AgriSense BCS Ltd., Pontypridd). The cardboard base and
sticky plastic had a hole in its centre, under which was placed a plastic beaker
(4 cm  6 cm) covered with mesh. The beakers were either empty or contained Fig. 1. The mean number (S.E.) of Lucilia sericata females caught on a white
50 g of liver. The liver used was bought from a local retailer, cut into pieces the sticky-trap under a range of different light intensities (W m2) in the presence
same day and left for 3 days at ambient laboratory conditions prior to use. (*) or absence of (&) liver bait.

2.1. Small cage trials P < 0.001; Fig. 1). There was no significant interaction between
light intensity and the presence or absence of liver bait (F = 2.70,
Batches of 30 adult female C. vomitoria or L. sericata were transferred to
P = 0.05), indicating that the responses of flies to light intensity
the wind tunnel in a plastic cup and placed into each six mesh cages
(35 cm  35 cm  35 cm) positioned in the centre of the wind tunnel in two followed the same pattern in both liver and control treatment
rows of three, perpendicular to the air flow. One sticky-trap had been placed in groups. Tukey multiple range tests showed a significant
the centre of each mesh cage prior to the release of the flies, which were then left difference between the 100% (345 W m2) and 32%
for 3 h before the number of flies caught on each sticky-target was counted. (110 W m2) treatments and between the two higher and the
Light intensities within the wind tunnel were changed using neutral density
two lower light intensities, but no significant difference between
filters wrapped around the fluorescent light source, to give light intensities of
approximately 100%, 32% and 12%. The experiment was also run with lights the number caught at the lowest light intensity or in darkness.
off. The precise irradiances were confirmed using a light intensity meter and
found to be: 345 W m2 (full illumination), 110, 40 and 0 W m2 (darkness). Four 3.2. C. vomitoria
replicates of this experiment were performed for the four light intensities for
each species, two with liver, and two without.
A greater number of flies was caught when traps were baited
2.2. Large scale trial
with liver than when unbaited (F = 22.66, P < 0.001; Fig. 2)
and more flies were caught at higher light intensities
In the dark in the small cages, it was possible that if flies were spontaneously
active they might have alighted on the sticky-traps at random rather than having
demonstrated any specific orientation towards the liver bait. To allow this
possibility to be considered further, in a second trial 200 female C. vomitoria or
L. sericata were released directly into the entire wind tunnel with a larger sticky-
trap (30 cm  30 cm) and 100 g liver as bait. Flies were again left for 3 h before
the numbers caught were counted.

2.3. Data analysis

All data were tested for normality and, where required, log10 transformation
was used to normalise the residuals. The log10 (+1) of the number of flies caught
was then subjected to analysis of variance with the light intensity and bait
presence or absence as fixed factors. Tukey pairwise comparisons were used to
compare the means of within treatment groups.

3. Results

3.1. L. sericata

A greater number of flies was caught when traps were baited Fig. 2. The mean number (S.E.) of Calliphora vomitoria females caught on a
with liver than when unbaited (F = 20.19, P < 0.001; Fig. 1) and white sticky-trap under a range of different light intensities (W m2) in the
more flies were caught at higher light intensities (F = 43.08, presence (*) or absence of (&) liver bait.
96 J. Wooldridge et al. / Forensic Science International 172 (2007) 94–97
Fig. 3. The mean number (S.E.) of female Lucilia sericata (^) and Calli-
phora vomitoria (^) landing on a white sticky-trap in the presence or absence
of a carrion bait (3-day-old liver), with all light intensity treatments pooled.
(F = 18.63, P < 0.001; Fig. 2). Again, there was no significant
interaction between light intensity and the presence or absence
of liver bait (F = 1.26, P = 0.29). Tukey multiple range tests
showed that there was no significant difference in the number
caught at the two highest light intensities (3.45 and 1.1 W m2.
However, there was a significant difference between the
numbers caught at two higher and the two lower light intensity
treatment groups and between at the lowest light intensity and
darkness.
3.3. Comparison between species
Fewer C. vomitoria were caught than L. sericata overall
(P < 0.05) and in both control and liver-bait treatment groups
(Fig. 3).
3.4. Large scale trial
When flies were released into the entire wind tunnel in the
dark, rather than the small mesh cages, a significantly lower
mean percentage of flies was caught than when they were
released in the dark into the smaller cages (x2 = 30.01;
P < 0.005). Means of 1.04% and 24.22% of the L. sericata
were caught and 3.0% and 14.72% of the C. vomitoria were
caught, in the wind tunnel and cages, respectively.
4. Discussion
Many insects, such as moths, flies, beetles and bees, are
well known to be able to fly at night in very low light
intensities [11,12]. Nocturnal flight activity is often strongly
related to ambient temperature. However, to date, the debate
over the ability of calliphorids to locate a corpse and oviposit
nocturnally has been inconclusive. Here, the number of L.
sericata and C. vomitoria caught, which it is assumed reflects
underlying levels of flight activity, decreased as light intensity
was reduced. It is notable however, that in the small cages
used, captures were not reduced to zero even in, what was
assumed to be total darkness. The room in which the wind
tunnel was placed had no windows and no other form of
illumination such as infra-red lighting; however, the possi-
bility of some low level illumination, for example, leaking
from under the laboratory door, can not be completely ruled
out. If this was the case it was certainly at a level that was to
low to be measured by the light intensity meter used here.
Nevertheless, the graded response to light intensity, particu-
larly when the liver bait was present, is surprising. From an
adaptive perspective, it might have been anticipated that flies
would remain active down to a threshold below which their
eyes could not function effectively, and that an all-or-nothing
relationship between activity and light intensity would have
been observed. The ommatidia of compound eyes are capable
of complex photoreceptor dark adaptation, although resolu-
tion may decrease as the signal to noise ratio increases at lower
light intensities [13,14].
The responses of the two species were broadly similar, but L.
sericata were significantly more active than C. vomitoria,
especially at the higher light intensities and particularly at
110 W m2. As a result, the decline in activity of L. sericata
appeared to show something of a step response, with a
substantive decline in activity between 110 and 40 W m2
whereas the decline in activity seen in C. vomitoria appeared to
be more gradual. This difference may be related to their
different behaviours in the field; C. vomitoria occur more
commonly in shaded and edge-habitats found in woodland and
hedgerows. These environments have a lower level of
illumination than the open, field habitats where L. sericata
are more commonly caught [15–17].
The numbers of flies of both species that were caught was
higher in the presence of liver, but the presence of the liver did
not change the shape of the relationship between activity and
light intensity. This shows that light intensity acts as an
exogenous independent stimulus for activity and that although
liver odours increase the level of activity of the flies, they are
not necessary as an activating stimulus. Hence fly activity can
be modulated directly by light intensity, without the need for
volatile odours to trigger activation. The fact that L. sericata
were more active than C. vomitoria both in the presence and
absence of the liver bait also suggests that the difference in
activity level is not causally related to any difference in the way
these two species respond to odour cues.
In the small cages, the fact that flies continued to be caught
in the dark, particularly on the liver-baited sticky-trap,
suggested that either that there was undirected flight and that
flies, moving at random, eventually alighted by chance on the
sticky surface, or that the flies were capable of oriented flight
towards the odour source in darkness. To attempt to distinguish
between these alternative possibilities, trials were carried out in
the much larger volume of the entire wind tunnel. In these trials,
captures were reduced to almost zero, suggesting that the
former alternative was most likely; in darkness, while flies may
be activated by carrion odours, they do not appear to be capable
of the oriented flight necessary to bring them to the source of
 J. Wooldridge et al. / Forensic Science International 172 (2007) 94–97
that odour. Captures in darkness, were probably therefore the
result of random movements.
The results presented here do not exclude the possibility that
these species can locate an oviposition site and oviposit
nocturnally. An especially full moon or artificial lighting might
provide a strong enough source of illumination, particularly in
urban areas. Nevertheless, the probability of oriented flight
leading to oviposition in the absence of such illumination
would appear to be relatively low.
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