Journal of Medical Entomology, 53(2), 2016, 296–303

doi: 10.1093/jme/tjv202
Advance Access Publication Date: 15 December 2015
Direct Injury, Myiasis, Forensics Research article

Entomofaunal Succession Patterns on Burnt and Unburnt

Rabbit Carrion
Ashraf M. A. Mashaly1,2,3

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1
Department of Zoology, College of Science, P.O. Box 2455, King Saud University, Riyadh 11451, Saudi Arabia (mmashely@ksu.
edu.sa), 2Department of Zoology, Faculty of Science, Minia University, El-Minia, Egypt, and 3Corresponding author, e-mail:
mmashely@ksu.edu.sa

Received 7 August 2015; Accepted 24 November 2015

Abstract
The influence of burning on the decomposition of rabbit carcasses and on insect succession was investigated in
three different habitats (agricultural, desert, and urban) in order to provide data for estimating the postmortem
interval (PMI). Each site had six carcasses divided into two groups of three rabbits, with the carcasses in one
group being partially burned, while the others were not burned. Carrion reached the dry stage within 5 d in the
desert and urban habitats and 13 d in the agricultural habitat. The unburnt and burnt carcasses also decom-
posed at a similar rate in the three study habitats. Adult dipteran and coleopteran insects were collected daily.
A total of 24 species and 2,381 specimens were collected; of these, 732 specimens from 21 species were taken
from the partially burnt carcasses and 1,649 specimens from 21 species from the unburnt carcasses. There
were significantly distinct insect communities between the agricultural habitat and the desert and urban habi-
tats. There were also significant differences in the insect communities between the decay stage and other
stages, with fresh and dry stages recording the lowest number of insects. There were some species which only
presented themselves during the decay stage of decomposition, namely, Platypalpus sp., Desmometopa vari-
palpis Malloch, Atherigona orientalis (Schiner), Atherigona yorki Deeming, Musca sorbens Weidemann, and
Onthophagus nitidulus Klug. In addition, there were significant distinctions in the insect communities between
unburnt carcasses and burnt carcasses. The presence of these distinctions means that it is possible to estimate
the PMI from partially burnt rabbit carcasses.

Key words: burnt, decomposition, forensic entomology, insect succession

Animal carcasses have been used as a model for forensic entomologi-
cal research in many parts of the world, including Saudi Arabia,
where rabbits were first used for these purposes in the work of
Abouzied (2014) in the south-western mountains. Entomological
succession varies according to different factors but, under similar sit-
uations, it follows a specific pattern, which can help in the estima-
tion of the postmortem interval (PMI; Smith 1985). Even within the
same region, the succession pattern of species may vary according to
the state of the carcass, and the conditions around the body, which
may lead to a delay in the duration of the decay stages (Micozzi
1986, Mann et al. 1990, Catts 1992, Goff 2000, Amendt et al.
2007). These changes may have an effect on the activity of each spe-
cies participating in the decomposition process (Avila and Goff
1998).
There have been a number of studies that have investigated insect
attraction to burnt remains, and their utility in calculating an accu-
rate PMI for these bodies (Avila and Goff 1998, Introna et al. 1998,
Pai et al. 2007, Chin et al. 2008, Vanin et al. 2013). McAlpine et al.
(1981) demonstrated that the insect fauna collected from unburnt
and burnt carcasses include a large number of similar species during
each of the decomposition stages. Avila and Goff (1998) and Kolver
(2009) confirmed this finding, showing no difference in the insect
fauna that visited normal and burnt carcass, although they did find
differences in the succession pattern and process of decomposition,
with the process being between 1 and 4 d faster in the burnt car-
casses. Byrd and Castner (2001), meanwhile, found that the level of
burning affects the decomposition rate and then the succession pat-
tern. Moreover, if the succession pattern for burnt carcass is used to
estimate the PMI of unburnt carcasses, the PMI will be overesti-
mated (Pai et al. 2007). Chin et al. (2008), however, reported that
the rate of decomposition and the insect succession pattern were not
affected by burning, and therefore does not influence the PMI
estimation.
Postmortem burning of remains as a means of destroying physi-
cal evidence has been documented in various areas of forensic litera-
ture (Barillo and Goode 1996, Sledzik and Micozzi 1997, Fanton
et al. 2006, DeHaan 2008). Identification of the specific pattern and
rate of decomposition in charred remains would be useful for foren-
sic investigators (Gruenthal et al. 2012). Thus, in the present study,
we hypothesized that the burning of rabbit bodies would affect

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V
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Journal of Medical Entomology, 2016, Vol. 53, No. 2
Table 1. Summary of weather data at the sites over the duration of the experiment
Site Daily temperatures ( C)
High Mean
Agricultural 29.3 6 1.7 23.6 6 1.2
Desert 43.2 6 2.1 36.4 6 4.7
Urban 39.9 6 1.8 34.3 6 3.3
insect succession and the rate of decomposition in three different
habitats in Riyadh, Saudi Arabia.
Materials and Methods
Site Description
The study was conducted in three different ecological habitats: an
agricultural area (24 44’36.54” N, 46 33’45.12” E) with many
palm trees and grasses; a desert area (24 47’41.61” N, 46
32’38.47” E) with rocky ground; and an urban area (24 43’16.51”
N, 46 37’2.45” E) represented by the roof of the building of the
College of Science, King Saud University. At each of the three sites,
there were very few human dwellings. The study was conducted dur-
ing June 2014. Temperatures and relative humidity (Table 1) were
recorded using a Lascar EL-USB-2 data logger.
Experimental Procedures
Eighteen live mature rabbits (Oryctolagus cuniculus L.) were pro-
cured locally, euthanized with chloroform, weighed (mean ¼
1.57 6 0.44 kg), and divided into three groups of six in each habitat.
At each study site, rabbit carcasses were arranged in a regular, sym-
metrical pattern of two parallel rows, with three carcasses in each
row spaced 2 m apart. Sampling occurred daily after the carcasses
were set down. In order to simulate a crime scene, one group of rab-
bits in each site was partially burned using wood without accelerant,
soon after their death, while the other group was left unburnt
and served as the control. Each carcass was put inside an iron cage
(55 by 40 by 24 cm3) in order to avoid attack by vertebrate scaven-
gers. Four pitfall traps (10 cm in diameter) were placed around each
rabbit carcass. On each sampling day, each trap was filled up with a
solution of water, soap, and salts. Collections were made daily dur-
ing the sampling period, and only adult specimens were included in
the count of collected insects. The adult insects collected were then
killed by freezing, labeled by date and hour of collection, sorted into
families, and pinned. Fly specimens were identified by Prof. Magdi
El-Hawagry (Entomology Department, Faculty of Science, Cairo
University, Egypt), while beetle specimens were identified by Prof.
Aly Maghrabi (King Saud University, College of Food and
Agricultural Sciences, Riyadh, Saudi Arabia).
Data Analysis
In order to compare the mean durations of the decomposition stages
between the three different habitats, ANOVA was applied followed
by Bonferroni correction. SPSS software (Version 15; SPSS,
Chicago, IL) was used for the statistical analyses. A significance level
at P  0.05 was used in all tests.
Results
The duration of the decomposition stages was found to be highly af-
fected by the ambient temperature. The rate of decomposition was
297
Daily relative humidity (%)
Low High Mean Low
20.5 6 1.9 45.6 6 3.2 39.2 6 1.8 30.4 6 3.3
34.1 6 3.6 19.5 6 1.4 17.2 6 2.2 13.4 6 2.6
32.6 6 2.4 22.3 6 2.7 19.1 6 1.3 15.6 6 1.1
13 d to reach the dry stage in agricultural habitats and 5 d in desert
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and urban habitats. In addition no difference in the rate of decompo-
sition was recorded between burnt and unburnt carcasses.
Figure 1A indicates the total number of insects attracted to rab-
bit carcasses in the three different habitats, based on the mean fig-
ures recorded in Table 2. The agricultural habitat had the highest
abundance of insects (1,861) compared to desert and urban habitats,
where 125 and 395 individuals were identified, respectively. This
difference was statistically significant (P < 0.05).
Figure 1B shows the number of insects attracted to the different
decomposition stages, also based on the mean figures in Table 2.
The decay stage showed the highest abundance of flies (about 1,308
individuals) followed by the bloat stage (727 individuals). The fresh
and dry stages had the lowest value (42 and 40 individuals, respec-
tively). Concerning beetles, the same pattern of colonization was de-
tected, where decay stage had the greatest abundance of beetles (107
individuals) followed by the bloat stage (60 individuals), the fresh
stage (53 individuals), and the dry stage (44 individuals). This differ-
ence was statistically significant (P < 0.05). In general, unburnt car-
casses had more flies and beetles (174 individuals) than burnt
carcasses (Table 2).
Eight species of flies belonging to five families were collected
from the urban habitat, 17 species from seven families were col-
lected from the agricultural habitat, and seven species belonging to
four families were collected from the desert habitat. Species and
their abundance are summarized in Table 3. Concerning beetles, five
species of flies belonging to three families were collected from the
urban habitat, eight species from five families were collected from
the agricultural habitat, and five species belonging to three families
were collected from the desert habitat (Table 4).
Among the 24 species collected (as shown in Tables 3 and 4),
18 colonized both types of carcasses. Platypalpus sp. (Hybotidae),
Desmometopa varipalpis Malloch (Milichiidae), and Megaselia
scalaris Loew (Phoridae) were collected only from the unburnt car-
cass, whereas Atherigona yorki Deeming and Musca sorbens
Weidemann (Muscidae) and Onthophagus nitidulus Klug
(Scarabaeidae) were present only on the burnt carcasses. Insects
were found in all decomposition stages in the three different habi-
tats, apart from the fresh and dry stages of both unburnt and burnt
carcasses in the urban habitat (Tables 3 and 4). Moreover, no flies
were found in the dry stage of burnt carcasses in the desert habitat
(Table 3).
Figure 2 explores the differences in the number of insect species
attracted to carcasses in the three different habitats. In the agricul-
tural habitat, 17 fly species from seven families were collected. In
the desert habitat, seven species of flies from four families were col-
lected. In the urban habitat, however, 10 species of flies from five
families were collected (Fig. 2A). In general, the results in Fig. (2A)
indicated that the desert habitat had the lowest abundance of flies.
Lucilia sericata was the most common species in the agricultural
and desert habitats, but in the urban habitat, M. domestica were the
298 Journal of Medical Entomology, 2016, Vol. 53, No. 2

2000
A
1800

1600 Flies Beetles

Mean number of insects ± SE

1400

1200

1000

800

600

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400

200

0
Agricultural Desert Urban
Habitats

1200
B
Mean number of insects ± SE

1000
Fresh Bloat Decay Dry

800

600

400

200

0
flies beetles flies beetles flies beetles
Agricultural Desert Urban
Habitats

Fig. 1. (A) Total number of insects attracted to the rabbit carcasses in each habitat. (B) The number of insects in the decomposition stages.

Table 2. Abundance of insects in the decomposition stage according to the carcass type in the three different habitats

Habitat Carcass Mean no. of flies 6 SE

Fresh Bloat Decay Dry

Agricultural Unburnt 7.24 6 0.61 512.13 6 7.32 653.43 6 6.34 3.31 6 0.41
Burnt 15.13 6 1.62 198.11 6 4.21 329.51 6 7.11 21.21 6 2.41
Desert Unburnt 6.23 6 0.54 7.45 6 1.1 2.13 6 0.31 6.12 6 0.57
Burnt 1.33 6 0.1 5.23 6 0.91 4.23 6 0.82 060
Urban Unburnt 060 2.33 6 0.32 277.46 6 8.12 060
Burnt 13.32 6 2.1 3.21 6 0.53 43.34 6 3.37 10.27 6 1.31
Mean no. of beetles 6 SE
Agricultural Unburnt 15.5 6 2.1 21.71 6 1.8 28.12 6 1.6 19.67 6 1.2
Burnt 060 4.62 6 0.76 22.23 6 1.6 9.56 6 1.1
Desert Unburnt 31.2 6 3.1 17.5 6 1.2 060 10.45 6 1.4
Burnt 6.23 6 0.9 7.34 6 1.3 18.41 6 1.7 4.21 6 0.77
Urban Unburnt 060 5.3 6 0.72 23.91 6 1.7 060
Burnt 060 3.17 6 0.64 15.16 6 2.01 060
Journal of Medical Entomology, 2016, Vol. 53, No. 2 299

Table 3. Flies succession on rabbit carcasses in three different habitats

Family Species Urban habitat

Fresh Bloat Decay Dry

Unburnt Burnt Unburnt Burnt Unburnt Burnt Unburnt Burnt

Calliphoridae Lucilia sericata H H

Lucilia cuprina H
Hybotidae Platypalpus sp. H
Muscidae Atherigona theodori H H H H
Hydrotaea capensis H
Musca calleva H H H

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Musca domestica H H H H H
Sarcophagidae Ravinia pernix H H
Ulidiidae Physiphora alceae H H H
Agricultural habitat
Calliphoridae Lucilia sericata H H H H H H
Lucilia cuprina H H H H H
Milichiidae Desmometopa varipalpis H
Muscidae Atherigona orientalis H
Atherigona theodori H H H H
Atherigona yorki H
Hydrotaea capensis H H H H H H
Musca calleva H
Musca domestica H H H H
Musca sorbens H
Muscina stabulans H H H H H H
Phoridae Megaselia scalaris H
Sarcophagidae Ravinia pernix H H H H H H
Sarcophaga hirtipes H H H
Sepsidae Sepsis lateralis H H H H
Ulidiidae Physiphora alceae H H H H H
Physiphora smaragdina H H
Desert habitat
Calliphoridae Lucilia sericata H H H H H
Lucilia cuprina H
Muscidae Atherigona theodori H
Hydrotaea capensis H
Musca domestica H H H
Sarcophagidae Ravinia pernix H H
Ulidiidae Physiphora alceae H H

H—Presence of flies.

most abundant. Concerning beetles, in the agricultural habitat, six
species from four families were collected, three species from three
families in the desert habitat, and two species from two families in
the urban habitat. The results indicated that the urban habitat had
the lowest abundance of beetles. Saprinus moyses was the most com-
mon species in the agricultural and desert habitats, but in the urban
habitat, Dermestes maculates were the most abundant (Fig. 2B). In
general, the results in Fig. 2 indicate that L. sericata and S. Moyses
were the most common species from flies and beetles, respectively,
and can be used to estimate the PMI.
Figure 3 shows that the unburnt carcasses attracted 21 species of
insects (16 flies species belonging to eight families and five beetles
species belonging to three families), compared to burnt carcasses
which attracted 24 species of insects (15 flies species belonging to
five families and six beetles species belonging to four families).
L. sericata and M. domestica were the most abundant flies on
both burnt and unburnt carcasses (Fig. 3A). The most abundant bee-
tles on both types of carcasses were S. moyses and D. maculates.
Two species from the 18 species of flies were not attracted
to unburnt carcasses, namely, A. yorki and M. sorbens. In contrast,
three species were not attracted to the burnt carcasses,
Platypalpus sp., De. varipalpis, and Me. scalaris. O. nitidulus was
the only species of beetle that was not attracted to unburnt carcasses
(Fig. 3B). These results in Fig. 3 indicate that there is a correlation
between the carcass type (burnt and unburnt) and the species that
were attracted.
Figure 4 explores the differences in the species numbers attracted
to the different stages of decomposition. The decay stage had a sig-
nificant species abundance compared to other stages of decomposi-
tion, with 17 species of flies and six species of beetles being
represented (Fig. 4A and B). The bloat stage had the second highest
level of abundance, with 12 species of flies and six species of beetles.
The fresh stage had a lower abundance, with nine species of flies
and five species of beetles followed by the dry stage, which had six
species of flies and five species of beetles. L. sericata was the most
abundant species in the decay stages compared to the other species
in the same stage. Saprinus moyses was the most abundant beetle in
the decay stage (Fig. 4A and B).
300 Journal of Medical Entomology, 2016, Vol. 53, No. 2

Table 4. Beetles succession on rabbit carcasses in three different habitats

Family Species Urban habitat

Fresh Bloat Decay Dry

Unburnt Burnt Unburnt Burnt Unburnt Burnt Unburnt Burnt

Cleridae Necrobia rufipes H H H

Dermestidae Dermestes maculatus H H H H
Histeridae Saprinus sp. H
Saprinus semipunctatus H
Saprinus moyses H
Agricultural habitat

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Cleridae Necrobia rufipes H
Dermestidae Dermestes maculatus H H H H H H
Histeridae Saprinus sp. H H H H
Saprinus semipunctatus H H H H
Saprinus moyses H H H H H H
Scarabaeidae Onthophagus nitidulus H H H
Maladera insanabilis H
Scolytidae Coccotrypes sp. H
Desert habitat
Cleridae Necrobia rufipes H H H
Dermestidae Dermestes maculatus H H H H H H
Histeridae Saprinus sp. H
Saprinus semipunctatus H
Saprinus moyses H H H H H

H—Presence of beetles.

800
A
700
Agricultural Desert Urban
600
Mean no. of flies ± S.E.

500

400

300

200

100

Fly species

70

60 Agricultural Desert Urban B

50
Mean number of beetles ± SE

40

30

20

10

Beetle species

Fig. 2. Abundance of insect species according to the habitat. (A) Flies. (B) Beetles.
Journal of Medical Entomology, 2016, Vol. 53, No. 2 301

700
A
600 Unburn
nt Burnt

ies ± SE
500

of flies
400

Mean number offl 300

200

100

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0

Fly species

70
Unburnt Burnt B
60
eetles ± SE
obeetles

50

40
Mean number of

30

20

10

Beetle speciies

Fig. 3. Abundance of insect species according to the carcass type. (A) Flies. (B) Beetles.

Discussion
In this study, the decomposition stages of burnt and unburnt car-
casses were represented by four stages of decomposition as observed
by Rodriguez and Bass (1983) and Avila and Goff (1998). The de-
composition process took 13 d in the agricultural habitat but just 5 d
in the desert and urban habitats. The process was apparently mainly
affected by the high temperature and humidity percentage, as has
been described before by Goddard and Lago (1985) and Anderson
(2009). The present study also found no difference in the rate of de-
composition between burnt and unburnt carcasses, as described by
Chen et al. (2008) and Gruenthal et al. (2012).
Matuszewski et al. (2010) said that the succession pattern of in-
sects differs depending on the geographical region. In this study, the
agricultural habitat exhibited a higher abundance of insects due to
the lower temperature compared to desert and urban habitats.
In addition, a significant difference was recorded between the
faunal succession on the burnt and unburnt carcasses (Fig. 3A and
B, P < 0.05) including the number of species and individuals repre-
sented. Whereas the unburnt carcasses attracted more flies, the
burnt carcasses attracted more beetles. These results agreed with
data reported by Chin et al. (2008) in his study on pig carcasses. He
found that the unburnt carcass attracted more flies than the burnt
carcass. Vila and Goff (1998), meanwhile, noted that the burnt car-
casses attracted much more fly oviposition than unburnt carcasses.
In the fresh stage, eight species of flies and four species of beetles
were recorded, in broad agreement with Goff (2010), who said that
the first wave of colonization by necrophagous insects includes adult
sarcophagids, generally arriving in the first minutes to hours of the
fresh stage. In this study both types of carcass attracted flies in the
fresh stage, as shown by Vila and Goff (1998) but in contrast to
Chin et al. (2008) and Oliveira-Costa et al. (2013), who each stated
that only unburnt carcasses attract flies in the fresh stage. Rivers and
Dahlem (2014) reported that, depending on the season, adult beetles
from the family Silphidae would arrive at any time during the fresh
stage.
In general, the burnt carcasses attracted more flies than the
unburnt ones during this initial fresh stage. This may be because the
high temperature at the study sites caused the smoke residue from
302 Journal of Medical Entomology, 2016, Vol. 53, No. 2

600
A
500 Fresh Bloat Decay Dry

Mean number of flies ± SE

400

300

200

100

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0

Fly species

45
40 Fresh Bloat Decay Dry B
35
Mean number of beetles ± SE

30
25
20
15
10
5
0

Beetle species
Fig. 4. Abundance of insect species according to the decomposition stage. (A) Flies. (B) Beetles.

burning to evaporate quickly, and no gasoline was used in the burn-
ing process. Chin et al. (2008), for example, concluded that the
lower incidence of insects on the fresh stage of burnt carcasses was a
consequence of the smoke and the gasoline odor exhaled for some
hours by the burnt carcass and of the temperature of the carcass.
In the bloat stage, there were generally more insects on the
unburnt carcasses than on the burnt carcasses. Avila and Goff
(1998) concluded that the abundant fluids exuded in the bloat stage,
due to the premature exposure of the viscera, might attract more
flies. Also, in the decay stage, a significant increase (Fig. 4A and B,
P < 0.05) in the number of insects was recorded. Moreover, a higher
number of insects were collected from the unburnt carcasses than
from the burnt carcasses. In the dry stage, however, the number of
insects was greatly decreased. In general, due to the effect of high
temperature, the remains of the freshly burned carcass were much
drier than those of the unburned carcass. Wardle (1921) and
Oliveira-Costa et al. (2014) indicated similar results.
In conclusion, in this study, burning has been shown not to have
an effect on the rate of decomposition but to have an effect on insect
succession. The insect succession showed a different pattern in the
different habitats (agricultural, desert, and urban) and between
burnt and unburnt carcass. Also, the decay stage showed the highest
value of insect abundance while the fresh and dry stages recorded
the lowest number of insects. These differences mean that PMI can
still be estimated from partially burnt rabbit carcasses.
Acknowledgments
I extend my appreciation to the Deanship of Scientific Research at King Saud
University for funding the work through the research group project number
RGPVPP- 028.
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