Forensic Science International 211 (2011) 67–75

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Forensic Science International

journal homepage: www.elsevier.com/locate/forsciint

Decomposition and insect succession of clothed and unclothed carcasses in

Western Australia
Sasha C. Voss *, David F. Cook, Ian R. Dadour
Centre for Forensic Science M420, The University of Western Australia, 35 Stirling Highway, Crawley, Western Australia 6009, Australia

A R T I C L E I N F O A B S T R A C T

Article history: The effect of clothing on carcass decomposition and patterns of insect succession onto remains were
Received 24 September 2010 investigated in two separate years during autumn in Western Australia. The progression of
Received in revised form 12 April 2011 decomposition differed between clothed and unclothed carcasses in both years of the study. The
Accepted 18 April 2011
presence of clothing markedly prolonged the wet decay stage in both years with larval feeding occurring
Available online 24 May 2011
across the moist skin surface underneath clothing, as well as within and under the carcasses. Ambient
temperatures were higher in the second year of the study and corresponded to marginally faster rates of
Keywords:
decay throughout decomposition. Within years, insect arrival and oviposition were largely consistent
Forensic entomology
Succession
between clothed and unclothed carcasses with a few notable exceptions. The green blow fly, Lucilia
Decomposition sericata Meigen (Diptera: Calliphoridae) oviposited one day earlier on clothed than unclothed carcasses
Clothing in both years of the study. The black carrion fly, Australophyra rostrata Robineau-Desvoidy, (Diptera:
Post-mortem interval Muscidae) colonised clothed carcasses in two distinct waves of succession but only one wave of
ovipoistion was observed on unclothed carcasses in either year. Correspondingly, clothed carcasses
supported larval feeding by A. rostrata for a longer duration than unclothed carcasses. Finally, dipteran
larval masses were more widely distributed across the carcass surface and were present for a longer
period of time on clothed carcasses than on unclothed carcasses in both years. Forensically relevant data
detailing the seasonal pattern of insect succession onto clothed and unclothed decomposing remains in
Western Australia are reported.
ß 2011 Elsevier Ireland Ltd. All rights reserved.

1. Introduction
The establishment of a post-mortem interval (PMI) of victims of
unexplained death is a vital step in many forensic investigations
[1,2]. Knowledge of the biology, behaviour and distribution of
insect species found in association with decomposing remains has
proven invaluable to investigators as a tool in helping establish
PMI and/or indicating post-mortem movement of the body [3–5].
Decomposing remains represent a temporary, changing habitat,
offering both food and shelter resources to numerous arthropod
species. The activity of the insect species that utilise this resource
gradually alters the state of the carcass, such that different species
are attracted to, and colonise remains at different time periods and
stages of decomposition [6,7]. The timing of insect colonisation,
development and eventual departure from decomposing remains
is a predictable and orderly process for a given set of conditions and
is closely linked to the progression of carcass decomposition
[5,7,8]. Entomological estimates of PMI are typically based on
known patterns of insect succession and the developmental age of
* Corresponding author. Tel.: +61 8 6488 7286; fax: +61 8 6488 7285.
E-mail addresses: sasha.voss@uwa.edu.au, varisvoss@gmail.com (S.C. Voss).
immature insects collected from the body [4]. As such, under-
standing the factors that affect decomposition and the associated
arrival and departure of insects colonising remains is key to the
accuracy of entomological estimates of time since death [9].
Many abiotic and biotic factors influence the rate of decompo-
sition and insect succession onto remains including geographic
location [9–11], climatic conditions [12,13], season [14,15], habitat
[16–18], the physical state of the remains [19] and the
decomposition environment [20–23]. Therefore, entomological
estimates of PMI require baseline reference data detailing the
expected pattern of insect succession onto decomposing remains
for a given set of parameters. Problems arise when the research
conditions under which succession data are collected do not match
the specifics of the investigated death scene. Given the wide range
of death scene scenarios encountered by forensic investigators
there is a clear dissociation between the available succession data
and the applicability of these data to forensic casework.
One of the main factors contributing to variation in the pattern
of insect succession onto remains is the degree of insect access to
the body [24]. When a body decomposes in an enclosed
environment or is wrapped in some way, carcass attendance by
individual insect species may be delayed or prevented, altering the
timing and expected pattern of insect succession [21,23,25]. The

0379-0738/$ – see front matter ß 2011 Elsevier Ireland Ltd. All rights reserved.
doi:10.1016/j.forsciint.2011.04.018
68 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75
rate of decomposition can also be affected by a reduction in insect
numbers due to the slower removal of total biomass through larval
feeding [24].
A common death scenario is the decomposition of clothed
human remains, yet the influence of clothing on decomposition has
received limited attention and is mainly focused on soil burial as
opposed to surface decomposition [26,27]. The data that are
available are geographically specific and often inconsistent.
Reports indicate that the presence of wrapping or clothing can
have different effects on decomposition rate, either slowing or
accelerating decomposition. Clothing can act as a barrier to
colonising insects and thus delay insect succession and slow
decomposition [26]. For instance, Goff [21] reported that wrapping
a body in layers of blankets can cause a delay of up to several days
in blow fly invasion of a corpse by reducing access to the body.
Conversely, clothing can also accelerate the decay process by
slowing down postmortem body cooling and providing a favour-
able micro-environment for insect development [9]. Clothing can
offer protection from ambient conditions (wind, direct sunlight,
and rain) and predators [24]. The micro-environment beneath
clothing can be warmer and more humid than ambient conditions
of exposed carcasses, resulting in greater insect diversity,
abundance and feeding activity and consequently increasing the
rate of decomposition [9,24,28].
Insect succession patterns associated with clothed carcasses
can also change as a consequence of the altered carcass state.
Carcasses can remain moist beneath clothing for longer periods
than exposed carcasses and as such may be attractive to certain
insect species that prefer wet environments [28,29]. Further, the
fluid retention properties of most garments can also increase larval
abundance and thus decomposition rate as clothing soaked in
decomposition fluid can increase ovipositonal opportunities for
blow flies [26]. Conversely, a recent work by Kelly et al. [29] in
central South Africa reported no difference in insect succession
patterns or the timing of blow fly oviposition between clothed and
unclothed carcasses during warmer seasons (autumn and sum-
mer). Clothing and wrapping did, however, influence the rate of
decomposition and biomass loss resulting in a prolonged period of
advanced decay and reduced biomass loss [29].
Although there is the potential for variation in insect succession
patterns on clothed and unclothed decomposing remains, at
Fig. 1. Four defined stages of pig decomposition observed during the study: fresh (A), bloat (B), wet (C) and dry (D).
present, there are no available reference data for Australia detailing
the affect of clothing on decomposition and insect succession. As
the species that colonise decomposing remains and their behav-
iour can be geographically specific, data for the Australasian region
are needed to address this gap in current knowledge. In particular,
there is a paucity of data detailing the succession patterns of
synanthropic flies associated with clothed carcasses. As such, this
study aimed to investigate the effect of clothing on carcass
decomposition and the linked succession of calliphorid flies during
autumn in Western Australia. The decomposition and succession
data associated with clothed and unclothed pig carcasses are
compared.
2. Materials and methods
2.1. Study site
A study of pig (Sus scrofa Linneaus) decomposition was conducted at a wildlife
reserve situated 23 km south of Perth, Western Australia (328 100 S, 1158 500 E). The
site encompassed 253 ha of coastal bushland, consisting of predominately jarrah
(Eucalyptus marginate Donn ex Smith), marri (Corymbia calophylla Lindley),
paperbark (Melaleuca spp.) and grass trees (Xanthorrhoea spp.).
The study was conducted during autumn and was repeated in two separate
years, 2001 and 2003. Trials commenced on the 5th April 2001 and the 13th March
2003. The date of death was designated as day 0 and each trial ran for 98 days. Daily
records of ambient temperature at the study site were recorded throughout the
study using a Datataker DT50 data logger (Hinco Instruments, Canning Vale, WA,
Australia). Records of ambient temperature, rainfall and humidity were also
obtained from the closest Australian Bureau of Meteorology weather station to the
study site (approximately 7 km; 328 100 S, 1158 880 E).
2.2. Animal model
Ten freshly killed, 40–45 kg domestic pig carcasses (S. scrofa) were used as
models for human decomposition. The domestic pig is the internationally preferred
animal model for human decomposition [30]. Five carcasses were used in each year.
At the start of each trial, pigs were sacrificed by captive head bolt. Following death,
the head wound was immediately closed using a silicon sealant so that the wound
did not present a focus for insect activity. Three carcasses were clothed and two
were left unclothed per trial. Clothed carcasses were dressed in light-weight, loose
cotton T-shirts and shorts with elastic waist bands. The two unclothed carcasses
acted as controls. Carcasses were placed directly on the soil surface on their side,
head facing east, legs facing north, in a partially shaded site, approximately 10 m
apart. An abdominal probe inserted into the centre of the abdomen of each carcass,
post-mortem, recorded changes in internal temperature of the cadaver via a
connection to a Tinytag Plus1 data logger (Hastings Data Loggers, Port Macquarie,
NSW, Australia). All data loggers were programmed to record the minimum and
maximum temperature at 30 min intervals. Pigs were protected from large animal
 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75
scavenging by individual metal cages, covered with 2.5 cm wire mesh, which was
placed over the entire pig carcass. The cage sides were extended 30 cm into the
ground to prevent scavengers burrowing beneath the cage. All carcasses were
exposed to insect activity within 15 min of death. Unfortunately, during 2003 the
cage protecting one of the unclothed carcasses was not sufficient to prevent native
animal scavenging [31] and considerable disturbance to the carcass occurred. As
such, data from the replicate were not included in the results of this study.
2.3. Sampling procedure
Decomposition is a continuous process that is commonly characterised into
discrete stages for descriptive purposes. Five different stages of decomposition have
previously been defined for pig decomposition; fresh, bloat, wet, dry and skeletal
[23]. For the purposes of this study, the fresh stage occurred from the moment of
death to the onset of bloat (Fig. 1). Bloat was considered completed and wet
decomposition commenced, upon deflation of the carcass. Escaped body fluid and
liquefaction were evident during the wet stage (Fig. 1). The dry stage commenced
when the majority of the carcass flesh and fluid had been removed and only dry
constituents, skin, cartilage and bone remained (Fig. 1). The skeletal stage is defined
by the absence of soft tissue with only hair and bone remaining.
During sampling, replicates were observed and photographed for assessment of
the decomposition state of the carcass, the presence of maggot masses and the
insect species present. Soil temperatures associated with each carcass were also
measured using a handheld Digithermo temperature probe (Oregon Scientific,
Hung Hom, Kowloon, Hong Kong). Measurements were taken at the soil interfaces
on four sides of the carcass (head, abdomen, rear and backline) and used to provide
an average soil temperature for each carcass. A control temperature was also
recorded 1 m away from each carcass.
Sampling of insect fauna was conducted between 0900 and 1200 h daily for the
first 14 days, and then every second day until the carcasses had progressed well into
the dry decomposition stage. Subsequently sampling was conducted once a week
Fig. 2. Autumn succession onto clothed carcasses (2001). Species occurrence reported where specimens were present on a sample day. Differing line widths (adults – black,
immatures – grey) indicate specimen presence on one, two or three replicates. Decomposition and life stage identified by the following:
adults, (E) eggs, (L) larvae and (P) pupae. Indicates maggot masses (>500) present.
69
until day 98. The selected sampling days encompassed time periods of significant
change in carcass decomposition. Representative samples of dipteran eggs, larvae
and pupae from each replicate were also collected. Insects associated with carcasses
were sampled through aerial sweeps and manual collection (paintbrush/forceps). In
the advanced stages of decomposition, the area immediately surrounding each
carcass, extending outward for approximately 1 m, was carefully hand sieved to a
depth of 5 cm for the presence of adult insect taxa and immature specimens.
Emphasis was placed on the collection and identification of Diptera and Coleoptera
specimens rather than a catalogue of all arthropod species present. Collected
specimens were labelled in relation to replicate number, sampling time and
collection site, placed in ventilated vials and stored at 6  2 8C during transport to
the laboratory. Adult specimens were killed at 20 8C and pinned for later
identification. Half of all larval samples were killed in hot water and preserved in
70% ethanol (egg samples were directly preserved in 70% ethanol). The remaining
specimens were reared to adulthood for identification. Adult flies were identified based
on published keys [32,33] and with the aid of reference specimens previously
identified by the Department of Food and Agriculture, Western Australia.
3. Results
3.1. Rate of decomposition
The progression of decomposition differed between clothed and
unclothed carcasses in both years of the study, notably in respect to
the duration of wet decay (Figs. 2–5). Within years there was no
difference in the onset or duration of bloat between clothed and
unclothed carcasses. All carcasses were in full bloat within 24 h of
death. Carcasses from both treatments remained in bloat for 4–5
bloat, wet, dry, (A)
70 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75

Fig. 3. Autumn succession onto unclothed carcasses (2001). Species occurrence reported where specimens were present on a sample day. Differing line widths (adults – black,
immatures – grey) indicate specimen presence on one or two replicates. Decomposition and life stage identified by the following: bloat, wet, dry, skeletal,
(A) adults, (E) eggs, (L) larvae and (P) pupae. Indicates maggot masses (>500) present.

days in 2001 and 2–3 days in 2003 (Figs. 2–5). While insect activity
and larval feeding contributed to skin breakage in all carcasses,
extensive abdominal splitting was common on clothed carcasses.
The presence of shorts caused constriction around the abdomen
during bloat and lead to abdominal splitting above the waistband
of the shorts. This resulted in fluid seepage across the entire front of
the T-shirt on clothed carcasses. In comparison, skin breakage
occurred mainly along the carcass regions directly in contact with
the soil on unclothed carcasses. Within years, the duration of the
wet decay stage was considerably longer for clothed than
unclothed carcasses (6 days longer). Clothed carcasses did not
dry out as quickly and supported maggot masses for a greater
period of time than unclothed carcasses. The presence of clothing
markedly prolonged the wet decay stage in both years with larval
feeding occurring across the moist skin surface underneath
clothing as well as within and under the carcasses. In contrast,
on unclothed carcasses, moisture and larval activity was focused
within and under the carcass only.
All carcasses eventually progressed into the dry remains stage
with the majority of tissue removed such that carcasses were
nearly resting flat against the soil. In some cases the head and
lower extremities were largely skeletal while the upper legs and
abdominal core retained desiccated tissue. Skeletisation of the
entire carcass was not evident by the end of the 98-day study
period for any of the clothed carcasses in either year of the study
(Figs. 2–5). Desiccated tissue remained beneath clothing, appear-
ing partially mummified and often supporting Australophyra
rostrata Robineau-Desvoidy (Diptera: Muscidae) (previously
Hydrotaea rostrata). Skeletal remains of unclothed carcasses in
2001 and 2003 were evident by days 98 and 91 respectively
(Figs. 2–5).
Decomposition rates differed between years. Ambient tem-
peratures were typically higher throughout the study period
during 2003, with a daily average ( standard deviation) of
17.4  4.0 8C as opposed to 15.7  3.3 8C in 2001, corresponding to
marginally faster rates of decay (2 days difference) throughout the
fresh to dry stages of decomposition (Figs. 2–5). Average relative
humidity and total rainfall were, however, slightly higher during the
study in 2001 compared to 2003 (64.9  14.7% and 253.8 mm;
59.8  15.4% and 249.8 mm respectively).
3.2. Insect succession
The succession patterns of representatives of the orders Diptera
and Coleoptera onto clothed and unclothed carcasses are
presented in Figs. 2–5. Nineteen species were collected and
identified throughout the study consisting of eleven Diptera and
eight Coleoptera. Of the predominant Diptera taxa, Calliphora dubia
Macquart (Diptera: Calliphoridae) consistently colonised all
carcasses within the first 24 h in both years of the study on both
clothed and unclothed carcasses. Other early calliphorid coloni-
sers, arriving during either the fresh or bloat stages of decomposi-
tion, included Calliphora albifrontalis Malloch, Chrysomya
megacephala Fabricius, Lucilia sericata Meigen and Chrysomya
rufifacies Macquart. Although C. albifrontalis was an abundant
primary coloniser of both clothed and unclothed carcases in 2001
 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75
Fig. 4. Autumn succession onto clothed carcasses (2003). Species occurrence reported where specimens were present on a sample day. Differing line widths (adults – black,
immatures – grey) indicate specimen presence on one, two or three replicates. Decomposition and life stage identified by the following:
adults, (E) eggs, (L) larvae and (P) pupae. Indicates maggot masses (>500) present.
the species was rarely observed to frequent remains and did not
oviposit on carcasses in 2003. The muscid flies Musca domestica
Linnaeus and Musca vetustissima Walker were observed regularly
on carcasses throughout decomposition but no offspring were
found or collected during sampling in either year of the study
(Figs. 2–5).
Later dipteran colonisers included Sarcophagidae, Chrysomya
varipes Macquart, A. rostrata and Piophila casei Linnaeus. When
present, fly larvae of the family Sarcophagidae typically larvipos-
ited towards the end of bloat and start of the wet decay. C. varipes
and A. rostrata adults were generally observed in association with
carcasses during the bloat stage of decomposition however, eggs
and/or larvae were not noted until the wet decay stage or later.
Representatives of the Piophilidae family, P. casei Linnaeus
attended carcasses predominantly during the wet decay and dry
stages of decomposition. Immature specimens were, however,
only observed from clothed and unclothed carcasses in 2001 and
clothed carcasses in 2003 after carcass decomposition had
progressed into the dry remains stage (Figs. 2–5).
The arrival and oviposition of blowflies was consistent within
years between clothed and unclothed carcasses with only a few
notable exceptions. The green blow fly, L. sericata, oviposited one
day earlier on clothed than unclothed carcasses in both years of the
study. This 24 h difference was consistent between all clothed (6)
and unclothed (3) replicates. A. rostrata, was observed to re-strike
clothed carcasses in both years of the study, initially at the onset of
71
bloat, wet, dry, (A)
the wet decay stage and then again towards the end of the wet
decay stage. This secondary wave was not observed on unclothed
carcasses in either year. Further, clothed carcasses continued to
support A. rostrata larval feeding throughout the duration of the
study while larvae were no longer observed on unclothed carcasses
after day 77 in either year (Figs. 2–5). Finally, while the presence of
maggot masses was commonly observed in association with all
carcasses throughout the wet decay stage, maggots remained in
masses for a longer period of time on clothed carcasses, were more
visible and occurred across a wider area of the carcass surface than
on unclothed carcasses. Larval aggregations were widespread on
clothed carcasses, often observed between the clothing and the
skin’s surface, along the carcass/soil interface and inside the
carcass itself. Daily counts of the location of maggot masses (skin’s
surface, carcass/soil interfaces and inside the carcass) on clothed
and unclothed carcasses indicated that location was dependant on
treatment in both years of the study (2001, x2 = 7.712, df = 2,
P < 0.05; 2003, x2 = 6.151, df = 2, P < 0.05). Significantly more
maggot masses occurred on the skin’s surface on clothed (under
clothing) than unclothed carcasses while maggot masses were
equally common inside body cavities and along the carcass/soil
interface in both treatments.
Coleoptera taxa were present on carcasses as early as day 2,
although numbers were initially limited to just one or two
individuals. Beetle abundance increased considerably during the
wet decay and dry remains stage in both years and Coleoptera were
72 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75

Fig. 5. Autumn succession onto unclothed carcasses (2003). Species occurrence reported where specimens were present on a sample day (adults – black, immatures – grey).
Decomposition and life stage identified by the following: bloat, wet, dry, skeletal, (A) adults, (E) eggs, (L) larvae and (P) pupae. Indicates maggot
masses (>500) present.

the dominant taxa during the late dry remains stage. Dominant
coleopteran species included Dermestes spp., Saprinus spp.
Creophilus erythrocephalus Fabricius and Necrobius rufipes De Geer.
Dermestidae were observed to breed on both clothed and
unclothed carcasses with larvae typically noted in high numbers
on carcasses beginning to progress into the dry remains stage
(Figs. 2–5). The beetles, Helea castor Pascoe, Omorgus tatei
Blackburn, Onthophagus binodis Thunberg and Ptomaphilia lacry-
mosa Schreibers, were less common and typically present after
carcasses had entered the dry remains stage.

3.3. Soil and internal carcass temperature

Average internal carcass temperature of both clothed and

unclothed carcasses was significantly higher than ambient
temperature throughout the study in both years (Friedman
Repeated Measures ANOVA on Ranks; 2001, x2 = 68.49, df = 2,
n = 37, P < 0.001; x2 = 63.95, df = 2, n = 37, P < 0.001). Differences
between ambient and internal carcass temperatures were greatest
throughout the wet decay stage (Fig. 6). There was no significant
difference, however, between clothed and unclothed carcasses in
either year of the study (Tukey Multiple Comparison Test, P < 0.05;
Fig. 6). Similarly, soil temperature beneath both clothed and
unclothed carcasses was significantly elevated above control soil
temperature (taken 1 m from carcasses) in both 2001 (x2 = 46.52,
df = 2, n = 31, P < 0.001) and 2003 (x2 = 37.23, df = 2, n = 31,
Fig. 6. Ambient temperature and average internal carcass temperature for clothed
P < 0.001). There was no significant difference between the soil and unclothed carcasses in 2001 (A) and 2003 (B). Recordings were stopped when
temperatures of clothed and unclothed carcasses (Tukey Multiple the first probe was exposed through larval feeding and tissue loss to ambient
Comparison Test, P < 0.05; Fig. 7). temperature (day 38).
 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75
Fig. 7. Average control soil temperature (1 m from carcasses) and soil temperature
associated with clothed and unclothed carcasses during decomposition in 2001 (A)
and 2003 (B).
4. Discussion
This study considered the impact of clothing on insect
succession and carcass decomposition and assessed the potential
implications for entomological estimation of PMI. Focus was
placed on the succession of Diptera and Coleoptera, as represen-
tatives of these orders were considered the primary indicator
species for estimating time since death. Considerable differences
were evident in the decomposition rates of clothed and unclothed
carcasses and minimal, but not discountable, differences were
evident in the associated pattern of insect succession.
Our findings largely agree with that reported by Kelly et al. [29]
in that there was no evidence of a delay in Calliphoridae arrival or
oviposition on clothed carcasses. This contrasts with Goff’s [21]
findings, where fly access to wrapped and securely tied carcasses
was delayed by 2.5 days. This latter difference is likely attributed to
the tighter constriction level of the wrapping as opposed to the
clothing used in this study. The presence of fairly loose clothing
such as T-shirts and shorts appeared to offer no barrier to fly access
to decomposing remains. In this study, oviposition by L. sericata
occurred 24 h earlier on clothed carcasses compared to unclothed
carcasses and this trend was consistent, within years, between all
replicates. Kelly et al. [29] reported that oviposition by Lucilia spp.
occurred simultaneously on clothed and unclothed carcasses in
South Africa. Individual species were not identified or separated in
respect to arrival time and oviposition suggesting that differences
in the succession of individual Lucilia species cannot be discounted.
Alternatively, differences in the succession of L. sericata onto
clothed remains in South Africa and Australia could be attributed
to differences in the attraction response of the two geographically
separate populations [34]. In either case, the observed earlier
colonisation of clothed carcasses by L. sericata introduces a
potential for error in the estimation of time since death when
PMI is based on the arrival and developmental time of this primary
colonising species.
Blow fly location of a suitable breeding resource involves a
series of steps including initial activation of interest, orientation
73
and site assessment culminating in oviposition itself [35]. Previous
research has indicated that while visual and tactile cues play a role
in this process, the cues used by Lucilia spp. are predominantly
olfactory and can be driven by the volatile compounds produced by
different bacterial strains [35–38]. Clothing on carcasses may have
facilitated bacterial incubation against the skin’s surface and
heightened L. sericata attraction to clothed carcasses, however, this
does not explain why only L. sericata responded differently to
changed bacterial odours due to clothing (and not the other species
recorded). Alternatively, Davies and Hobson [49] demonstrated the
importance of moisture as a stimulant for oviposition by L. sericata
and a similar finding has been noted for other Lucilia species
[39,40]. The micro-environment beneath clothing is potentially
warmer and more humid than the surface condition of skin on
unclothed carcasses, particularly clothing wet by either body fluid
or rainfall [9,24,28]. This altered micro-environment may have
been favoured by L. sericata more than the other species recorded.
As such, an investigation of the micro-ambient conditions
occurring beneath clothing and against the skin of clothed carcass
during decomposition in respect to temperature, humidity and
bacteria would prove useful in understanding clothed carcass
decomposition. Characterisation of such conditions would assist in
identifying potential olfactory attractants and ovipositional cues
presented to blowflies as well as establishing the developmental
conditions experienced during egg development and eclosion on
clothed carcasses.
In general, species composition observed in association with
both clothed and unclothed carcasses were consistent with
previous studies undertaken in Western Australia for the autumn
season [11,23]. Differences in the colonisation of C. albifrontalis
between years were likely related to the timing of the two trials. C.
albifrontalis is a primary coloniser of remains in all seasons except
summer and low population numbers likely account for the
absence of immatures of this species in 2003 which commenced
earlier than the 2001 trial [11].
The relationships between the progression of decomposition,
temperature and insect activity have been well documented
[5,41,42]. Differences in decay rates observed between years can
be attributed to climatic conditions, seasonal variations in insect
populations and insect development. Voss et al. [23] reported a
faster rate of wet decay (9 days) in 2001 for a surface exposed
carcass during early autumn 2001 in Perth. Within season
variation in carcass decomposition has previously been documen-
ted [43,44] and can be further exacerbated by differences in the
timing of decomposition commencement within a season due to
variation in ambient temperature. A vast number of factors
influence carcass decomposition and insect succession onto
remains in the field and while attempts to summarise data by
season and decomposition stage are useful they are nevertheless
artificial. For this reason, the physical appearance of the carcass is a
less reliable indicator of PMI than the succession and development
of insects on decomposing remains.
Within years, the duration of the wet decay stage was
considerably longer for clothed than unclothed carcasses (6
days longer). Maggot masses were present on all carcasses during
wet decay and internal carcass temperatures were consistently
above ambient. Clothed carcasses, however, supported a greater
abundance and distribution of maggot masses during wet decay.
As suggested by Kelly et al. [29] clothing potentially protects
maggots from environmental conditions and facilitates greater
movement across the carcass surface contributing to the greater
visibility and distribution of maggot masses. While the density of
maggot masses on clothed and unclothed carcasses was not
measured, maggot masses were clearly more visible and promi-
nent on clothed carcasses. This was likely an artifact of the greater
distribution of maggots on clothed carcasses. There was no
74 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75
significant difference in internal carcass temperature or soil
temperature between clothed and unclothed carcasses within
years, suggesting that maggot mass density on clothed and
unclothed carcasses were probably comparable.
The presence of clothing markedly prolonged the wet decay
stage in both years and likely contributed to the second
colonisation wave of A. rostrata and the subsequent extended
colonisation period observed. As A. rostrata occurs throughout the
advanced stages of decomposition, the species may have impor-
tance in cases where other indicator species have completed their
development [45]. As such, the appropriate generational wave
must be identified in cases were clothing is present, as inaccurate
identification of the generational wave could lead to substantial
errors in PMI.
Considerable variation between clothed and unclothed car-
casses was evident in respect to the occurrence of A. rostrata larvae
and pupae, and no trend was evident between years. In 2001, the
first pupae were collected 26 and 10 days after the first occurrence
of eggs on clothed and unclothed carcasses respectively. In
contrast, in 2003, the first pupae were collected 9 and 20 days
after the first occurrence of eggs on clothed and unclothed
carcasses respectively. A. rostrata predominately pupates on the
soil surface within the vicinity of the carcass. Potentially, as pupae
are more exposed on the surface, heavy predation by ants and
other foraging insects may have removed specimens prior to
sampling. Alternatively, while sampling was thorough, it is
possible we missed initial oviposition events by A. rostrata. Eggs
and first instar larvae of this species are extremely difficult to
locate since eggs are rarely laid in a mass. Individual eggs are
scattered across the skin’s surface, under extremities or in the folds
of clothing while first instar larvae are rarely aggregated and are
often located beneath the skin making detection difficult.
A. rostrata has previously been reported as a tertiary coloniser
by a number of studies [45–47] but this may be a misconception
related to sampling and detection issues and compounded by the
common occurrence of secondary strike waves by this species. The
authors have recently recorded A. rostrata eggs on guinea pig
carcasses as early as 6 h after exposure (unpublished data) while
Lang et al. [50] reported A. rostrata oviposition onto fresh possum
carcasses as early as day 2 of exposure. Given that uncertainty
remains regarding the timing of succession of A. rostrata onto
carcasses, and the identified importance of this species’ presence
throughout late post-mortem decomposition, further work is
needed to comprehensively detail succession data for this species.
4.1. Implications for forensic entomology
The accuracy of entomological estimates of time since death
rely on the establishment of multiple sets of baseline reference
data encompassing the huge variation evident in forensic death
scenes, for a given geographic region. Studies of different death
scenarios provide valuable data for forensic entomologists and
other practitioners such as pathologists and scene of crime officers
[48]. This study assessed the potential for variation in the rate of
decomposition and the associated pattern of insect succession
between clothed and unclothed carcasses for application to death
scenes in which remains are clothed. Data generated indicate that
reference succession data from unclothed decomposition can
reasonably be applied to clothed death scenarios where clothing is
loose fitting. Care should be taken however, where the key
indicator species of time since death is either L. sericata or A.
rostrata, as the potential for differences in the initial timing of
oviposition between clothed and unclothed carcasses must be
considered. Succession data generated by this work contribute to
the establishment of a succession reference database for Australia.
Such studies will improve the applicability of reference data to the
specifics of the investigated death scenario and ultimately advance
the precision of estimates of time since death in forensic
investigations.
Acknowledgements
We thank Natalie Campman, Filipa Carvalho, Bob Cooper,
Michelle Harvey, and Jeremy Lindsey for their support in the field.
The authors also thank Alexander Larcombe for his useful
comments on the manuscript. This project complies with the
ethical and legal requirements of Australian research.
References
[1] V.J. Geberth, Establishing Time of Death, in Practical Homicide Investigation:
Tactics, Procedures, and Forensic Techniques, CRC Press, Boca Raton, FL, 1996.
[2] B. Morris, I. Dadour, Forensic entomology: the use of insects in legal cases, in: I.
Freckelton, H. Selby (Eds.), Expert Evidence, Law Book Company, Sydney, 2005.
[3] J. Amendt, et al., Best practice in forensic entomology—standards and guidelines,
International Journal of Legal Medicine 121 (2007) 90–104.
[4] E. Catts, M. Goff, Forensic entomology in criminal investigations, Annual Review of
Entomology 37 (1992) 253–272.
[5] M.L. Goff, Estimation of postmortem interval using arthropod development and
successional patterns, Forensic Science Reviews 5 (1993) 81–94.
[6] R. Putman, The role of carrion feeding arthropods in the decay process, Ecological
Entomology 3 (1978) 133–139.
[7] W. Rodriguez, W. Bass, Insect activity and its relationship to decay rate of human
cadavers in East Tennessee, Journal of Forensic Sciences 28 (1983) 423–432.
[8] J.A. Payne, A summer study of the baby pig, Sus scrofa Linnaeus, Ecology 46 (1965)
592–602.
[9] C.P. Campobasso, G. Di Vella, F. Introna, Factors affecting decomposition and
Diptera colonization, Forensic Science International 120 (2001) 18–27.
[10] A. Galloway, et al., Decay rates of human remains in an arid environment, Journal
of Forensic Sciences 34 (1989) 607–616.
[11] S.C. Voss, H. Spafford, I.R. Dadour, Annual and seasonal patterns of insect
succession on decomposing remains at two locations in Western Australia,
Forensic Science International 193 (2009) 26–36.
[12] M. Archer, Rainfall and temperature effects of the decomposition rate of exposed
neonatal remains, Science and Justice 44 (2004) 35–41.
[13] B. Shean, L. Messinger, M. Papworh, Observations of differential decomposition on
sun exposed v. shaded pig carrion in coastal Washington State, Journal of Forensic
Sciences 38 (1993) 938–949.
[14] M. Johnson, Seasonal and micro seral variations in the insect populations on
carrion, American Midland Naturalist 93 (1975) 79–90.
[15] K. Tabor, C. Brewster, R. Fell, Analysis of the successional patterns of insects on
carrion in southwest Virginia, Journal of Medical Entomology 41 (2004) 785–795.
[16] T. Eberhardt, D. Elliot, A preliminary investigation of insect colonisation and
succession on remains in New Zealand, Forensic Science International 176 (2008)
217–223.
[17] L. Goff, A. Omori, K. Gunatilake, Estimation of postmortem interval by arthropod
succession, American Journal of Forensic Medicine and Pathology 9 (1988)
220–225.
[18] J. Davis, L. Goff, Decomposition patterns in terrestrial and intertidal habitats on
Oahu Island and Coconut Island, Hawaii, Journal of Forensic Science 45 (2000)
836–842.
[19] F. Avila, M.L. Goff, Arthropod succession patterns onto burnt carrion in two
contrasting habitats in the Hawaiian Islands, Journal of Forensic Sciences 43
(1998) 581–586.
[20] N. Centeno, M. Maldonado, A. Oliva, Seasonal patterns of arthropods occurring on
sheltered and unsheltered pig carcasses in Buenos Aires province (Argentina),
Forensic Science International 126 (2002) 63–70.
[21] L. Goff, Problems in estimation of postmortem interval resulting from the
wrapping of a corpse; a case study from Hawaii, Journal of Agricultural Entomol-
ogy 9 (1992) 237–243.
[22] J.A. Payne, E.W. King, Insect succession and decomposition of pig carcasses in
water, Journal of the Georgia Entomological Society 7 (1972) 153–162.
[23] S.C. Voss, S. Forbes, I.R. Dadour, Decomposition and insect succession on cadavers
inside a vehicle environment, Forensic Science Medicine and Pathology 4 (2008)
22–32.
[24] R. Mann, W. Bass, L. Meadows, Time since death and decomposition of the human
body: variables and observations in case and experimental field studies, Journal of
Forensic Sciences 35 (1990) 103–111.
[25] J.L.O. Pohjoismäki, et al., Indoors forensic entomology: colonization of human
remains in closed environments by specific species of sarcosaprophagous flies,
Forensic Science International 199 (2010) 38–42.
[26] G.S. Anderson, Factors that influence insect succession on carrion, in: J.H. Byrd, J.L.
Castner (Eds.), Forensic Entomology: The Utility of Arthropods in Legal Investgia-
tions, CRC Press, Boca Raton, FL, 2010, pp. 201–250.
[27] R.C. Janaway, Decomposition of materials associated with buried cadavers, in: M.
Tibbett, D.O. Carter (Eds.), Soil Analysis in Forensic Taphonomy: Chemical and
Biological Effects of Buried Human Remains, CRC Press, Boca Raton, FL, USA, 2008,
pp. 153–201.
 S.C. Voss et al. / Forensic Science International 211 (2011) 67–75
[28] L.C. Dillion, Insect Succession on Carrion in Three Biogeoclimatic Zones in British
Columbia in Department of Biological Sciences, Simon Fraser Univeristy, Bur-
naby, British Columbia, 1997.
[29] J.A. Kelly, T.C.v.d. Linde, G.S. Anderson, The influence of clothing and wrapping on
carcass decomposition and arthropod succession during the warmer seasons in
Central South Africa, Journal of Forensic Sciences 54 (2009) 1105–1112.
[30] K. Schoenly, et al., Comparative performance and complementarity of four
sampling methods and arthropod preference tests from human and porcine
remains a the Forensic Anthropology Centre in Knoxville, Tennessee Journal of
Medical Entomology 44 (2007) 881–894.
[31] R.C. O’Brien, et al., Forensically significant scavenging guilds in the southwest of
Western Australia, Forensic Science International 198 (2010) 85–91.
[32] J. Wallman, Third-instar larvae of common carrion breeding blowflies of the
genus Calliphora (Diptera: Calliphoridae) in south Australia, Invertebrate Taxon-
omy 15 (2001) 37–51.
[33] J.F. Wallman, A key to the adults of species of blowflies in southern Australia
known or suspected to breed in carrion, Medical and Veterinary Entomology 15
(2001) 433–437.
[34] J.B. Cragg, P. Cole, Laboratory studies on the chemo-sensory reactions of blowflies,
Annals of Applied Biology 44 (1956) 478–491.
[35] J. Ashworth, R. Wall, Responses of the sheep blowflies Lucilia sericata and Lucilia
cuprina to odour and the development of semiochemical baits, Medical & Veteri-
nary Entomology 8 (1994) 303–309.
[36] M. Morris, et al., Orientation stimulants from substances attractive to Lucilia
cuprina (Diptera, Calliphoridae), Australian Journal of Experimental Agriculture
38 (1998) 461–468.
[37] C.H. Eismann, M.J. Rice, The origin of sheep blowfly, Lucilia cuprina (Wiedemann)
(Diptera: Calliphoridae), attractants in media infested with larvae, Bulletin of
Entomological Research 77 (1987) 287–294.
[38] M.J.R. Hall, Trapping the flies that cause myiasis: their responses to host-stimuli,
Annals of Tropical Medicine & Parasitology 89 (1995) 333–357.
75
[39] L. Barton-Brown, The relationship between oviposition in the blowfly Lucilia
cuprina and the presence of water, Journal of Insect Physiology 8 (1962)
383–390.
[40] L. Barton-Brown, W.M. Rogoff, The ‘sheep factor’ and oviposition in Lucilia cuprina,
Australian Journal of Science 21 (1959) 189–190.
[41] L. Lopes de Carvalho, A. Linhares, Seasonality of insect succession and pig carcass
decomposition in a natural forest area in southeastern Brazil, Journal of Forensic
Sciences 46 (2001) 604–608.
[42] G.S. Anderson, Minimum and maximum development rates of some forensically
important Calliphoridae (Diptera), Journal of Forensic Sciences 45 (2000)
824–832.
[43] M. Marchenko, Medicolegal relevance of cadaver entomofauna for the deter-
mination of the time of death, Forensic Science International 120 (2001)
89–109.
[44] S. Matuszewski, et al., Insect succession and carrion decomposition in selected
forests of Central Europe. Part 1: pattern and rate of decomposition, Forensic
Science International 194 (2010) 85–93.
[45] I.R. Dadour, D.F. Cook, N. Wirth, Rate of development of Hydrotaea rostrata under
summer and winter (cyclic and constant) temperature regimes, Medical &
Veterinary Entomology 15 (2001) 177–182.
[46] M. Fuller, The insect inhabitants of carrion: a study in animal ecology, Bulletin of
the Australian Council for Scientific and Industrial Research 82 (1934) 1–62.
[47] J.L. Froggatt, An economic study of Nasonia brevicornis, a Hymenopterous parasite
of Muscid Diptera, Bulletin of Entomological Reseach 9 (1918) 257–262.
[48] C. Campobasso, F. Introna, The forensic entomologist in the context of the forensic
pathologist’s role, Forensic Science International 120 (2001) 132–139.
[49] W.M. Davies, R.P. Hobson, Sheep blowfly investigations, Annals of Applied Biology
22 (1935) 279–293.
[50] M.D. Lang, G.R. Allen, B.J. Horton, Blowfly succession from possum (Trichosurus
vulpecula) carrion in a sheep-farming zone, Medical and Veterinary Entomology
20 (2006) 445–452.