January-February 2006 1

FORUM

Forensic Entomology and Main Challenges in Brazil

LEONARDO GOMES AND CLÁUDIO J. VON ZUBEN
Depto. Zoologia, Univ. Estadual Paulista (UNESP), Av. 24A, 1515, Bairro Bela Vista, 13506-900, Rio Claro, SP

Neotropical Entomology 35(1):001-011 (2006)

Entomologia Forense e os Principais Desafios no Brasil

RESUMO - Exceto o primeiro relato na China no século XIII, as primeiras observações de insetos e
outros artrópodes como indicadores forenses foram documentadas na Alemanha e França durante a
exumação em massa em 1880 por Reinhard, reconhecido como co-fundador dessa disciplina. Após a
publicação francesa do livro popular de Megnin com os aspectos aplicados da entomologia forense, o
conceito rapidamente se espalhou pelo Canadá e Estados Unidos. Agora, pesquisadores reconhecem
que faltam observações sistemáticas que permitam o uso de insetos como indicadores do intervalo
pós-morte. Após a I Guerra Mundial, poucos casos de entomologia forense entraram para a literatura.
Entre 1960 a 1980, Leclercq e Nuorteva foram os principais responsáveis por manter o método na
Europa Central, estudando casos isolados. Desde então, pesquisas básicas nos Estados Unidos, Rússia
e Canadá abriram caminho para a rotina do uso da Entomologia nas investigações forenses. Na
região Neotropical, a identificação dos insetos associados com cadáveres humanos são escassos e têm
recebido pouca atenção no Brasil. Este artigo dá uma visão dos desenvolvimentos históricos nesse
campo, os recentes estudos e os principais problemas e desafios na América do Sul e, principalmente,
no Brasil.

PALAVRAS-CHAVE: Cadáver, crime, entomologia forense, medicina legal

ABSTRACT - Apart from an early case report from China (13th century), the first observations on
insects and other arthropods as forensic indicators were documented in Germany and France during
mass exhumations in the 1880s by Reinhard, who is considered a co-founder of the discipline. After
the French publication of Mégnin’s popular book on the applied aspects of forensic entomology, the
concept quickly spread to Canada and United States. At that time, researchers recognized that the
lack of systematic observations of insects of forensic importance jeopardized their use as indicators of
postmortem interval. General advances in insect taxonomy and ecology helped to fill this gap over
the following decades. After World Wars, few forensic entomology cases were reported in the scientific
literature. From 1960s to the 1980s, Leclercq and Nuorteva were primarily responsible for maintaining
the method in Central Europe, reporting isolated cases. Since then, basic research in the USA, Russia
and Canada opened the way to the routine use of Entomology in forensic investigations. Identifications
of insects associated with human cadavers are relatively few in the literature of the Neotropical
region and have received little attention in Brazil. This article brings an overview of historic
developments in this field, the recent studies and the main problems and challenges in South America
and mainly in Brazil.

KEY WORDS: Cadaver, crime, forensic entomology, legal medicine

With about one million described species, insects
comprise the largest metazoan class (Price 1997) and are
found in virtually all habitats. Vertebrate corpses consist
on excellent food sources for a more or less specialized insect
community (Anderson & Cervenka 2002) and
approximately 400 insect species were found to colonize a
pig cadaver during its several decayment stages (Payne
1965).
In addition to their ecological importance in
decomposition, such insects may represent an important tool
in criminal investigations (Erzinclioglu 1983, Catts & Goff
1992), allowing the estimation of time at which a dead body
was colonized (Greenberg 1991). In particular, blowflies
(Calliphoridae) may serve as a biological clock in measuring
the time of death for two or more weeks. Such an
entomologically-based estimate may be far more precise than
2 Gomes & Zuben - Forensic Entomology and Main Challenges in Brazil
the medical examiner’s, which is limited to about a day or
two after death (Greenberg & Kunich 2002). Therefore
forensic entomology, defined as the use of insects and other
arthropods, such as mites, in medicocriminal investigations
(Hall 2001), is becoming an important field in legal
medicine. In Brazil, this field is recent and this article
describes the historical development of forensic entomology,
the methods, current fields and future trends in world and
mainly in Brazil.
History
The first documented forensic entomology case is
reported by the Chinese lawyer and death investigator Sung
Tz’u in the medico-legal text book “Hsi yüan chi lu” (one
possible translation: “The washing away of wrongs”)
(McKnight 1981, Benecke 2001). This author describes the
case of a stabbing near a rice field. The day after the murder,
the investigator told all workers to lay down their working
tools (sickles) on the floor. Invisible traces of blood drew
blow flies to a single sickle. So confronted, the tool’s owner
confessed to his crime and “knocked his head on the floor”.
During medieval times, the correlation between maggots
on a cadaver and the oviposition of adult flies was not
recognized. However, the realistic and detailed illustration
of corpses containing maggots was not unusual (Benecke
2001). The idea of the spontaneous development of life
emerging from pure matter prevailed.
At the beginning of the 19th century it was registered
that flies are attracted by corpses at a very early stage of
decomposition. Mende (1829) compiled a list of
necrophagous insects, including flies, beetles and other taxa
and provided more precise account, but did not link flies to
the time of death. Kamal (1958) described the opportunities
and problems associated with using insects for the estimation
of the postmortem interval (PMI), many of which are still
relevant today.
During the mass exhumation in France and Germany in
the 18th and 19th centuries, medico-legal doctors observed
that buried bodies are inhabited by arthropods of many kinds.
In 1831, the French physician Orfila observed a large number
of exhumations (Orfila & Lesueur 1831) and understood
that maggots play an important role in the decomposition
of corpses.
The first modern forensic entomology report to include
an estimation of PMI was provided by the French physician
Bergeret in 1855. In his journal article, next to a long
description of the criminalistic impact in the trial, he also
describes the court proceedings: “Within three years, four
different families lived in the flat. The first of them left in
December 1848, and the person examined started to live
there at the end of 1844. I was brought to the house of Mme.
Saillard (i.e. landlady) in rue du Citoyen, 4, to examine the
corpse of a child. The questions we had to deal now were:
(1) was the child born regularly/at the right time; (2) was it
alive when it was born; (3) how long did it live; (4) how did
it die?; (5) what was the time interval between birth and
death?” Questions 1-4 were answered with classical forensic
pathology methods. Question 5 was commented in the
following way: “To answer this question, legal medicine
must check with another science, the Natural Sciences.”
Bergeret provided a brief overview on the life cycle of
insects in general. He mistakenly assumed, however, that
metamorphosis would generally require a full year.
Furthermore, he assumed that females generally lay eggs in
summer, and that the larvae would transform to pupae the
following spring and hatch in summer. Some details of
Bergeret’s calculation: “The eggs of larvae we found on the
corpse in March 1850 must have been deposited here in the
middle of 1849. Therefore, the corpse must have been
deposited before this time interval. Next to the many living
larvae there were numerous pupae present, and they must
come from eggs that have been laid earlier, i.e., 1848. Could
it be that the corpse was deposited even before that time
(i.e. 1848)? The fly that emerges from the pupae that we
found in the body cavities, is Musca carnaria L. that lays
its eggs before the body dries out. We found other pupae of
little butterflies of the night (i.e. moth, Benecke 2001) too,
that attack bodies that already dried out. If the body was
deposited, say in 1846 or 1847, we would not have found
those larvae (i.e. since they would have hatched). In
conclusion, two generations of insects were found on the
corpse, representing two years postmortem: on the fresh
corpse, the flesh fly deposited its eggs in 1848, on the dried
out corpse, the moth laid their eggs in 1849” (Benecke 2001).
In retrospect, one should understand that Bergeret did
not focus on forensic entomology in his report to the court
but used entomology as a forensic tool. Indeed, the
mummification of the cadaver appears to be his over-riding
issue of interest in this case. Bergeret quotes Orfila in the
matters of mummification and forensic entomology and also
clearly notes the lack of information concerning insect
succession on corpses in his days.
After referencing the work of Bergeret, Brouardel
describes the case of a newborn child that was autopsied by
him in 1878 (Brouardel 1879). The mummified body was
inhabited by several arthropods, including butterfly larvae
and mites, which led to a request for assistance from
Monsieur Perier, Professor at the Museum of Natural History
in Paris, and Army Veterinarian Pierre Mégnin. Perier
reported that the body was most likely dried out before it
was abandoned. The determination of mites was left to
Mégnin whereas Perier determined the butterfly larvae from
the genus Aglossa (Lepidoptera: Pyralidae). Perier concluded
that the baby may have been born and died the summer
before, i.e. around 6-7 months before corpse was autopsied.
Mégnin (1894) reported that the whole body was covered
with a brownish layer composed exclusively of mite skins
and mite feces, but not living mites. Inside the cranium, he
found large numbers of a single mite species. Initially, a
few larval mites must have been carried to the corpse by
other arthropods. Mégnin calculated that on the whole body,
2.4 million mites were present dead or alive. He also
calculated that after 15 days, the first generation with 10
females and five males had developed; after 30 days, 100
females and 50 males; after 45 days, 1000 females and 500
males. Finally, after 90 days, one million females and
500,000 males were present. Since this was the number of
January-February 2006 Neotropical Entomology 35(1)
individuals he estimated being on the corpse, he made a
conservative guess and reported that the corpse must have
been abandoned for at least five months (three months of
mite development, preceded by two months for desiccation)
but more likely 7-8 months.
In the following years, side issues such as grave fauna
(Reinhard 1882, Schmitz 1928), the skeletonizing of corpses
(Hauser 1926), or modification or corpses caused by insects
were explored, but data concerning the biology, ecology and
succession of necrophagous insects (e.g. Fuller 1934,
Bornemissza 1957) were not applied to determine
postmortem interval in Europe. Leclercq & Leclercq (1948),
Nuorteva (1959a, 1959b), Nuorteva et al. (1967) and
Leclercq (1983), were among the first to use forensic
entomology for the determination of the postmortem interval
in Europe.
This topic was revived by researchers such as Reiter &
Wolleneck (1982, 1983) and Reiter (1984) in studies on the
development of Calliphora vicina (Wiedemann) and by
several others (Marchenko 1980, Leclercq 1983, Rodriguez
& Bass 1983, Greenberg 1985, Lord et al. 1986, Goff et al.
1986, Introna et al. 1989). Now, at the beginning of the 21st
century, forensic entomology is recognized in many countries
as an important forensic tool (Goff 1991, Greenberg 1991,
Anderson 1995, Introna et al. 1998, Amendt et al. 2004).
Postmortem Changes of the Human Body
After death, human or animal bodies undergo many
changes caused by autolysis of tissue, which is promoted by
the internal chemical breakdown of cells and released enzymes
as well as by activity of bacteria and fungi, from the intestine
and the external environment (Amendt et al. 2004).
Table 1. Selection of insects of forensic importance.
Order Order/family
Coleoptera Cleridae
Dermestidae
Geotrupidae
Histeridae
Silphidae
Staphilinidae
Diptera Calliphoridae
Drosophilidae
Fanniidae
Muscidae
Phoridae
Piophilidae
Sarcophagidae
Stratiomyidae
Lepidoptera Tineidae
Hymenoptera Ichneumonidae
Pteromalidae
3
However, the precise rate of postmortem decay is affected
by a wide range of variables associated with the corpse itself
and the surrounding environment. Moreover, after body
temperature has equilibrated with the environment and
following the initial putrefaction, no reliable estimation of
the postmortem interval is possible. Therefore, insects found
on the body provide an important source of information.
Insects are attracted to a body immediately after death,
often within minutes (Erzinclioglu 1983, Smith 1986,
Anderson 2001). However, oviposition may not occur. Many
taxa which appear very early at a death scene are late
colonizers or even non-necrophagous species.
According to Smith (1986), four ecological categories
can be identified in a carrion community:
1. Necrophagous species, feeding and breeding? on the
carrion.
2. Predators and parasites of necrophagous species,
feeding on other insects or arthropods. This group also
comprises species which feed on the carrion at first, but
may become predaceous in later larval stages.
3. Omnivorous species such as wasps, ants and some
beetles feeding both on the corpse and its colonizers.
4. Other species, such as springtails and spiders, which
use the corpse as an extension of their environment.
Groups 1 and 2 comprise representatives of the orders
Diptera and Coleoptera (Table 1). The succession on corpses
can be divided into different waves over the various stages of
decay, although this has been debated (Schoenly & Reid 1987).
Nevertheless, as the attractiveness of a decaying body differs
among necrophylous insects, changes over time and corpse
colonization will occur in a predictable sequence (Fig. 1).
Blowflies are typically the first colonizers, attracted by
the odour produced during decomposition (Wall & Warnes
Important genera
Necrobia
Dermestes
Geotrupes
Hister
Necrodes, Silpha
Aleochara
Calliphora, Chrysomya, Cochliomyia, Lucilia
Drosophila
Fannia
Musca, Ophyra, Muscina
Conicera, Megaselia
Piophila
Liopygia, Sarcophaga
Hermetia
Tineola
Alysia
Nasonia, Muscidifurax
4 Gomes & Zuben - Forensic Entomology and Main Challenges in Brazil
Figure 1. Succession of adult arthropods on human cadavers in east Tenesse; adapted from Rodriguez & Bass (1983) and Hall
(2001).
1994, Anderson 2001), even over large distances (Braack
1981, Erzinclioglu 1996). Oviposition first occurs at the
orifices or wounds of the corpse. The size of the carcass
affects its attractiveness to flies of the families Calliphoridae
and Sarcophagidae (Nuorteva 1959b, Davies 1990,
Erzinclioglu 1996, Povolný & Verves 1997). Other
necrophagous species oviposit preferentially on smaller
animals such as rodents or even snails. Several factors restrict
the colonization of a corpse, such as its burial (Mann et al.
1990) and most Diptera are not able to colonize bodies buried
deeper than 30 cm (Introna & Campobasso 2000,
Campobasso et al. 2001); however exceptionally, groups
such as the Phoridae may be found in buried coffins (Schmitz
1928, Stafford 1971). Burial, therefore, will influence the
time required for insects to reach the carcass as well as the
species composition of the necrophagous fauna (Payne et
al. 1968, Campobasso et al. 2001). Such delay may not only
occur in buried corpses, but in those that are covered or
wrapped (Goff 1991) or in cadavers found at indoor scenarios
(Gomes et al. 2005).
The role of freshwater and marine fauna in forensic
investigations has received very little attention (Payne &
King 1972, Goff et al. 1986). Knowledge about the role of
aquatic arthropods during decomposition is still scanty
(Anderson 2001). Compared to terrestrial habitats,
decomposition in an aquatic environment is completely
different. It occurs at a rate roughly half that of
decomposition on land, mainly due to the prevention of
insect activity and lower temperatures (Knight 1991). Merrit
& Wallace (2001) have distinguished six decompositional
stages ranging from submerged fresh, floating decay to
sunken remains. Aquatic insects of forensic importance
belong to the Ephemeroptera, Trichoptera and Diptera; the
latter are mainly represented by Chironomidae (midges) and
Simuliidae (black flies). However, these insects, unlike their
terrestrial counterparts, are not obligatory saprophages, but
instead use the submerged carrion as a food source and a
breeding site. The use of these insects for estimating the
time of death is therefore more difficult and depends on the
season and on other conditions of the aquatic systems. No
successional insect model exists which describes the different
waves of colonization of a corpse in aquatic habitats (Merrit
& Wallace 2001).
Studies on animal carcasses have demonstrated that
species composition and insect succession on a cadaver vary
with respect to the geographical region and season
(Bornemissza 1957, Arnaldos et al. 2001, Carvalho &
Linhares 2001, Grassberger & Frank 2004). Data collected
for a particular region or area should be used with caution
when determining time of death in another region. Even
local characteristics of the death scene, like the ecology of
the area or the degree of sun exposure, can alter the pattern
of insect colonization (Smith 1986, Erzinclioglu 1996).
Some insect species are found in both urban and rural areas,
while others are very specific to a certain habitat (Catts &
Haskell 1990). Since species commonly considered as rural
species have also been collected in urban regions, care must
be used in determining whether remains have been moved
based on entomological evidence alone (Anderson 2001,
Grassberger & Frank 2003a).
Estimating Time Since Death
When human remains are found days, weeks, or even
longer after death, body temperature, and conditions such
as rigor mortis or livor mortis are no longer appropriate for
estimating time since death. In such cases, insects may
provide important indications of the postmortem interval
(PMI). The ages of insect immature stages found on a dead
body can provide evidence for the estimation of a minimum
PMI ranging from one day up to more than one month,
depending on the insects species involved and the climatic
conditions at the death scene. However, this period will not
always match the exact PMI (Oliveira-Costa 2003).
Exact species identification of insect samples is the first
essential step in estimating the age of the larvae found. Insect
January-February 2006 Neotropical Entomology 35(1)
larvae differ in growth rates and biology. Larvae of Lucilia
sericata (Wiedemann), for instance, grow faster at 25oC than
larvae of Calliphora vicina (Fabricius) (Amendt et al. 2000);
blowflies (Calliphoridae) usually deposit eggs on a corpse
while flesh flies (Sarcophagidae) are larviparous. These
examples demonstrate that the same stage of development
of larvae found on a corpse do not necessarily indicate the
same age or the same time of colonization. For estimating
the minimum PMI, the age of the immature larval stages
must be determined. Various procedures for estimating their
age exist, but they are all based on the fact that the rate of
development depends on the ambient temperature (Amendt
et al. 2004).
Numerous papers describe ADH/ADD values and
thresholds for forensically important insects, mainly
blowflies and flesh flies (e.g. Kamal 1958, Anderson 2000,
Kaneshrajad & Turner 2004). The usefulness of these
methods depends on the thermal history of the immature
stages and, to be used, they therefore require the evaluation
of the temperature data at the death scene before the body
was found. Hence, multiple temperature measurements have
to be taken at the crime scene, at the body as well in the
maggot masses on the cadaver. These data should be
compared with data obtained from the nearest weather
station. If these data states are reasonably similar, the
weather station data can simply be used to extrapolate
temperatures at the crime scene (Greenberg & Kunich 2002).
Alternatively, if more substantive differences exist, the
weather station temperature data have to be corrected for
the average difference using mathematical methods such as
linear regression, prior to extrapolation of the expected
temperatures at the crime scene. However, when using
estimates of ambient temperature to calculate development
rates, it should be noted that these data are not necessarily
representative of the temperatures experienced by insects
in the corpse. Large numbers of maggots may create a so-
called “maggot mass effect” which, according to their
metabolic and feeding rate, can generate a temperature
substantially higher than those from the ambient (Wells &
LaMotte 1995). Moreover, development data obtained in
different geographical regions may not be comparable and
however, in PMI evaluations, great caution must be applied
when using data collected by researchers form other
countries. Changes in range and precipitation, which may
lead species to change their time of hatching, length of life
cycle, photoperiod and diapause, must all be taken in
consideration (Greenberg 1991, Grassberger & Reiter 2001,
Turchetto & Vanin 2004).
Knowing the chronology of insects colonizing carrion
in a certain area, analysis of the fauna on a carcass is useful
to estimate the time elapsed since the death (Goff & Flynn
1991, Anderson 2001). A simple succession model can be
used when estimating both the age of a larva and the time
interval between death and the insect’s arrival on the body
(Wells et al. 2001). Succession data have been used to
calculate a PMI up to 52 days (Schoenly et al. 1996) and, if
there are adequate data, may be applied to a much longer
time interval.
It is important to consider factors that might alter the
5
time of oviposition, such as covering corpses with branches
or tight wrapping with blankets, carpets or plastic bags, and
indoor placement, because these factors may delay initial
oviposition (Higley & Haskell 2001). Seasonal influences,
such as cold and rainy weather, may inhibit or even prevent
fly activity and delay oviposition (Erzinclioglu 1996).
Blowflies oviposition activity usually peaks in the early
afternoon (Nuorteva 1959a; Baumgartner & Greenberg
1984, 1985; Greenberg 1990). These insects are not active
at night and generally do not lay eggs during nighttime
(Greenberg 1985). A postmortem interval estimation based
on that assumption has to consider the possibility that a
corpse which was found about noon and was infested by
recently hatched maggots, could have been deposited there
in the late evening of the previous day. Hence, fly eggs
detected on a corpse during the night would lead to the
conclusion that death occurred during previous day or earlier
(Nuorteva 1977). On the other hand, Tessmer et al. (1995)
reported that blowflies fail to lay eggs at night both in urban
(with lighting) and rural dark habitats. Hence nocturnal
oviposition is a possibility and should be taken into
consideration. Diapause is still a challenge for the forensic
entomology (Ames & Turner 2003).
Competition may affect development and growth of the
larvae. According to Smith and Wall (1997a, b), larvae of
L. sericata in carcasses experience significant levels of
competition and that the intensity of this competition may
be sufficient to reduce the number of adult L. sericata able
to emerge successfully.
Reiter (1984), Smith (1986) and Erzinclioglu (1990)
pointed out another factor that could complicate the
estimation of the postmortem interval: in some female flies,
eggs may be retained in the oviduct, having been fertilized
as they pass through the spermathecal ducts in advance of
the act of oviposition (Wells & King 2001).
Parasitoids are also used to calculate de PMI (Amendt
et al. 2000, Grassberger & Frank 2004). The calculation of
developmental times for the host and the parasitoid allowed
the estimation of a greater minimum postmortem interval
than the estimated development time. However, when
considering the potential influence, especially of larval
parasitoids, it is important to remember that this specialized
group might also create significant problems for forensic
entomology (Erzinclioglu1990).
DNA Analysis and Entomotoxicology in Forensic
Entomology
Although identification keys are available, only a few
experts are able to identify the larvae of forensically relevant
insects and differentiation of the larval stages using
morphological criteria is still not possible. Time-consuming
rearing of the larvae to adults for identification may delay
the criminal investigation or cause significant problems
when rearing fails. Under these circumstances, species
identification based on genetic examination is an option.
PCR amplification of suitable regions of the genome,
sequence analysis of the amplicons obtained, and alignment
of the data with reference sequences is the usual and
6 Gomes & Zuben - Forensic Entomology and Main Challenges in Brazil
recommended methods (Sperling et al. 1994, Stevens & Wall
1996, Wells & Sperling 1999).
Larvae which feed on corpses may sequester drugs and
toxicants which had been ingested by the deceased person.
Analysis of carrion-feeding insects, to detect toxic substances
and to investigate the effects on insect development, is known
as entomotoxicology (Goff & Lord 2001). Bodies in a state
of advanced decomposition or that are skeletonized may be
difficult to examine for toxicologically relevant substances
due to the lack of appropriate sources such as tissue, blood
or urine. Instead, analysis of insects encountered may enable
toxicological assessment of the cause of death (Nolte et al.
1992; Goff & Lord 1994, 2001; Introna et al. 2001;
Campobasso et al. 2001).
The detection of mercury in the larvae of various species
of blowflies reared on tissues containing known concentrations
of this metal was described by Nuorteva and Nuorteva (1982).
Ingested drugs or toxicants may influence the development
of the necrophagous insects (O’Brien & Turner 2004). Goff
et al. (1991) studied the effect of cocaine and heroin (on the
rate of development in Sarcophagidae and demonstrated that
maggots of Boetterisca peregrina (Fabricius) develop more
rapidly if reared on the liver or spleen of rabbits which had
been killed by a lethal dose of cocaine or heroin.
These examples demonstrate that insects found on
corpses can be used in toxicological analyses, but also
illustrate the risk of calculating an incorrect postmortem
interval because of a modified rate of development of the
immature stages in the presence of drugs and toxicants.
Further research should focus on the bioaccumulation and
metabolism of drugs in necrophagous insects and their effects
on the rate of development.
Forensic Entomology in South America
Identifications of insects associated with human cadavers
are scarce in the literature of the Neotropical region. Dunn
(1916) reported Hermetia illuscens (L.), a stratiomyid in
Panama; Pessôa and Lane (1941) presented previous records
and reviewed necrophilous Scarabaeidae from museums in
Brazil; Jirón et al. (1982) and Jirón & Solano (1988) gave
information on Calliphoridae, Sarcophagidae, Muscidae and
Stratiomyidae of Costa Rica. Recently, some experimental
studies of forensic entomology have been carried out in
different regions of Colombia (Wolff & Uribe 2000). In Cali,
Olaya (1999) determined the arthropod succession and the
rates of decomposition in dogs, but apparently no specific
identifications of insects found on human bodies have been
published.
In Argentina, some descriptive reports on cadaveric insects
have been published (Mariluis 1983, Mariluis et al. 1989),
but up to now, no attempt was made to determine faunal
associations related to stages of decomposition. Then, Centeno
et al. (2002) compared decay stages and cadaveric fauna on
sheltered pig carcasses, for each season of the year.
Forensic Entomology in Brazil
Until several years ago, Forensic Entomology had received
little attention in Brazil despite the considerable value of this
procedure that has been demonstrated in several countries.
Few papers were published between 1908 and 1940 (Pessôa
& Lane 1941, Carrera 1991), mostly directly influenced by
the pioneer work of Mégnin (1894). Between 1940 and 1991,
not a single paper on this subject was published in Brazil.
However, since then, research groups have appeared in states
of São Paulo, Rio de Janeiro, Paraná and Distrito Federal
(Souza 1994, Von Zuben et al 1996, Moura et al. 1997) which
have dealt with some of related topics.
In the end of 20th century, several studies dealing with
Forensic Entomology become to appear. For example,
Carvalho & Linhares (2001) identified arthropods
associated with pig carrion and human corpses in Campinas,
State of São Paulo. Andrade et al. (2005) investigated
Calliphoridae in human corpses in the State of Rio Grande
do Norte. Oliveira-Costa (2002, 2003), Oliveira-Costa
(2001a,b) and Oliveira-Costa & Mello-Patiu (2004, 2005)
carried out several studies with human cadavers in crime
scenes to estimate the PMI.
Other important studies were carried out by Gomes &
Von Zuben (2003a,b; 2004a,b,c; 2005a,b) and Gomes et al.
(2002, 2003, 2005) studying postfeeding larval dispersal of
blowflies (Calliphoridae). This sort of study may have
important implications for medical-criminal investigations
that use forensic entomology (Amendt et al. 2004) and can
be underestimated if older dispersing larvae or those that
disperse farther, faster and deeper are not taken into account
(Von Zuben et al. 1996; 2002, 2003, 2005).
In 2002, a project was initiated in Brasília, DF, the
“Centro Nacional de Entomologia Forense” (National Center
for Forensic Entomology), linked to the Laboratory of
Diptera and Forensic Entomology of the Department of
Zoology of Universidade de Brasília. This project is financed
by the Departamento de Justiça, Secretaria Nacional de
Segurança Pública and CNPq/MCT and aims at studying
the arthropod succession in region of Brasília and has
contacted the other laboratories of Forensic Entomology in
Brazil, but unfortunately this centre also has no permission
to follow the real cases of crimes.
Thyssen et al. (2005) assessed the usefulness of
polymerase chain reaction-restriction fragment length
polymorphism (PCR-RFLP) in the differentiation of two
species of forensic entomology importance. Two specific
regions of mitochondrial DNA, the Cytochrome oxidase
subunit I (COI) and the control region (CR), were amplified
by PCR and digested using restriction endonucleases. The
cleavage patterns generated by the endonucleases DraI and
SspI were suitable for differentiating the two Hemilucilia
species. This method can be helpful for the forensic
entomologist in estimating the PMI because it provides a
fast identification, also making possible the use of the insect
at any life stage, including immature specimens, regardless
of the conditions of preservation (dead or live specimens).
The Challenges of Forensic Entomology in Brazil
In spite of some efforts in divulge the works of Forensic
Entomology among scientists, there are some problems that
must be solved to improve and raise this area in Brazil.
An admissible death investigation requires the combined
January-February 2006 Neotropical Entomology 35(1)
efforts and cooperation of experts in different disciplines,
crime scene technicians, death investigators, forensic
pathologists, anthropologists, entomologists, other medical
and non-medical professionals. All these front-line experts
play a crucial role in every death investigation process.
Unfortunately, not all these disciplines are always
represented. They may become involved during the course
of the investigation depending on the procedures adopted
by medico-legal jurisdiction, on the nature of the case and
the need for verification of essential evidence (Campobasso
& Introna 2001, Von Zuben 2002).
When insects are found on the body, samples must be
collected in conjunction with other biological evidence. Even
if the important role of insects in human decay is well known,
unfortunately the medical training and experience of forensic
pathologists usually do not fully prepare them to extract all
of the available information from arthropods. For many
years, maggots crawling on dead bodies were considered
just another disgusting element of decay; something to be
rinsed away as soon as the body was placed on the table for
autopsy (Campobasso & Introna 2001).
Specific collecting techniques and standard procedures
in different environments at the death scene, at the autopsy
and in the laboratory have been developed to enhance the
recovery of insect specimens. Respecting the different
disciplines and competences, in our opinion, the complex
ecology of the decay process and the role played by insects
suggest that the collection and interpretation of entomological
evidence associated with dead bodies should be done
preferably by both the forensic entomologist and pathologist
correlating circumstantial information and findings noted at
the crime scene and at the autopsy (Von Zuben 2002).
It is no longer admissible that law enforcement officers,
death investigators or pathologists could be skeptical or could
not appreciate the value of insects as forensic indicators
(Campobasso & Introna 2001). Actually, the entomologist is
still not often included in the investigation from the beginning
and he is rarely called upon to visit the death scene.
As forensic scientists, it is our responsibility to encourage
all local jurisdictions and police officers to spend some
resources including on routine crime-scene investigation
teams or adopting in death investigations many different
professionals (as entomologist familiar with cadavers or
pathologist familiar with insects) or, at least, training crime-
scene technicians to recognize and properly preserve
entomological evidence.
In conclusion, we strongly encourage the interaction
between Brazilian forensic scientists involved in criminal
investigations and in particular between pathologists and
entomologists in death cases where insects are associated
with the remains.
Final Remarks
Despite 150 years of use, forensic entomology is still a
young discipline. The number of scientists working in this
field and the degree of international cooperation among the
various groups are increasing. In Europe there are at least
10 countries with groups of researchers concentrating on
7
entomology (http://www.eafe.org). There are international
attempts as standardization and quality assessment of the
techniques and methodologies. It can be hoped that in future
forensic entomology will play an even more important role
in the elucidation of acts of crime.
In spite of the problems in countries from South America,
the precise estimation of PMI is the most important goal of
forensic entomology by refining the techniques used.
Developmental and succession data, consideration of a
greater number of geographical regions and a range of death
scene scenarios are essential.
One of the most important challenges is to combine
experimental data and practical case work. Due to the wide
variations in biotic and abiotic factors which occur at death
scenes, an improvement of the existing understanding can
only be established trough an increased number of detailed
and quantified observations. Forensic entomologists are
always presented with the task of reconstructing the death
scene conditions as closely as possible. A model for the
calculation and handling of the data is crucial for the
credibility of this discipline (Amendt et al. 2000).
Acknowledgments
Thanks are due to Iracema Monteiro da Silva and
Guilherme Gomes for collecting the data and to Regina Lúcia
Sugayama for the opportunity. Leonardo Gomes has been
supported by fellowships from FAPESP (03/00540-3) and
Cláudio José Von Zuben by research fellowships from CNPq.
References
Amendt, J., R. Krettek, C. Niess, R. Zehner & H. Bratzke.
2000. Forensic entomology in Germany. Forensic Sci.
Int. 113: 309-314.
Amendt, J., R. Krettek & R. Zehner. 2004. Forensic
entomology. Naturwissenschaften 91: 51-65.
Ames, C. & B. Turner. 2003. Low temperature episodes in
development of blowflies: Implications for postmortem
interval estimation. Med. Vet. Entomol.17: 178-186.
Anderson, G.S. 1995. The use of insects in death investigations:
An analysis of cases in British Columbia over a five year
period. Can. Soc. Forensic. J. 28: 277-292.
Anderson, G.S. 2000. Minimum and maximum development
rates of some forensically important Calliphoridae
(Diptera). J. Forensic Sci. 45: 824-832.
Anderson, G.S. 2001. Succession on carrion and its
relationship to determining time of death, p.143-175.
In J.H. Byrd & J.L. Castner (eds.), Forensic entomology:
The utility of arthropods in legal investigations. CRC,
Boca Raton, 418p.
Anderson, G.S. & V.J. Cervenka. 2002. Insects associated
with the body: Their use and analyses,p.73-200. In W.D.
Haglund & M.H. Sorg (eds.), Advances in forensic
taphonomy: Method, theory and archaeological
perspectives. CRC, Boca Raton, 507p.
8 Gomes & Zuben - Forensic Entomology and Main Challenges in Brazil
Andrade, H.T.A., A.A. Varela-Freire, M.J.A. Batista &
Medeiros, J.F. 2005. Calliphoridae (Diptera) coletados
em cadáveres humanos no Rio Grande do Norte.
Neotrop. Entomol. 34: 855-856.
Arnaldos, I., E. Romera, M.D. Garcia & A. Luna. 2001. An
initial study on the succession of sarcosaprophagous
Diptera (Insecta) on carrion in the southeastern Iberian
peninsula. Int. J. Legal Med. 114: 156-162.
Baumgartner, D.L. & B. Greenberg. 1984. The genus
Chrysomya (Diptera: Calliphoridae) in the New World.
J. Med. Entomol. 21: 105-113.
Baumgartner, D.L. & B. Greenberg. 1985. Distribuition and
medical ecology of the blowflies (Díptera: Calliphoridae)
of Peru. Ann. Entomol. Soc. Am 78: 565-587.
Benecke, M. 2001. A brief history of forensic entomology
Forensic. Sci. Int. 120: 2-114.
Bornemissza, G.F. 1957. An analysis of arthropod succession
in carrion and the effect of its decomposition on the
soil fauna. Aust. J. Zool. 5: 1-12.
Braack, L. 1981. Visitation patterns of principal species of
the insect complex at carcasses in the Kruger National
Park. Koedoe 24: 33-49.
Brouardel, P. 1879. De la détermination de l’epoque de la
naissance et de la mort d’um nouveau-né, faite a l’aide
de la présence des acares et des chenilles d’aglosses
dans um cadavre momifié. Ann. Hyg. Pub. Méd. Lég.
2: 153-158.
Campobasso, C.P. & F. Introna. 2001. The forensic
entomologist in the context of the forensic pathologist’s
role. Forensic Sci. Int. 120: 132-139.
Campobasso, C.P., G. Di Vella & F. Introna. 2001. Factors
affecting decomposition and Diptera colonization.
Forensic Sci. Int. 120: 18-27.
Carrera, M. 1991. Insetos de interesse médico e veterinário.
Editora da Universidade Federal do Paraná, Curitiba
228p.
Carvalho, L.M.L. & A.X. Linhares. 2001. Seasonality of
insect succession and pig carcass decomposition in a
natural forest area in southeastern Brazil. J. Forensic
Sci. 46: 604-608.
Catts, E.P. & M.L. Goff. 1992. Forensic entomology in
criminal investigations. Annu. Rev. Entomol. 37: 253-
272.
Catts, E.P. & N.H. Haskell. 1990. Entomology and death: A
procedural guide. Joyce’s Print Shop, Clemson, USA,
182p.
Centeno, N., M. Maldonado. & A. Oliva. 2002. Seasonal
patterns of arthropods occuring on sheltered and
unsheltered pig carcasses in Buenos Aires Province
(Argentina). Forensic Sci. Int. 126: 63-70.
Davies, L. 1990. Species composition and larval habitats of
blowfly (Calliphoridae) populations in upland areas in
England and Wales. Med. Vet. Entomol. 4: 61-88.
Dunn, L.H. 1916. Hermetia illuscens breeding in a human
cadaver (Dipt.) Entomol. News 27: 59-61.
Erzinclioglu, Y.Z. 1983. The application of entomology to
forensic medicine. Med. Sci. Law 23: 57-63.
Erzinclioglu, Y.Z. 1990. On the interpretation of maggot
evidence in forensic cases. Med. Sci. Law 30: 65-66.
Erzinclioglu, Y.Z. 1996. Blowflies. Richmond Publishing,
Slough, 71p.
Fuller, M.E. 1934. The insect inhabitants of carrion: A study
in animal ecology. Council for Scientific and Industrial
Research Bulletin 82.
Goff, M.L. 1991. Comparison of insect species associated
with decomposing remains recovered inside dwellings
and outdoors on the island of Oahu, Hawaii. J. Forensic
Sci. 3: 748-753.
Goff, M.L. & M.M. Flynn. 1991. Determination of
postmortem interval by arthropod succession: A case
study from the Hawaiian Island. J. Forensic Sci. 36:
607-614.
Goff, M.L. & W.A. Brown, K.A. Hewadikaram & A.I.
Omori. 1991. Effects of heroin in decomposing tissues
on the development rate of Boettcherisca peregrina
(Díptera: Sarcophagidae) and implications of this effect
on estimation of postmortem intervals using arthropod
development patterns. J. Forensic Sci. 36: 537-542.
Goff, M.L. & W.D. Lord. 1994. Entomotoxicology: A new
area for forensic investigation. Am. J. Forensic Med.
Pathol. 15: 51-57.
Goff, M.L. & W.D. Lord. 2001. Entomotoxicology: Insects
as toxicological indicators and the impact of drugs and
toxins on insect development, p.331-340. In J.H. Byrd
& J.L. Castner (eds.), Forensic entomology: The utility
of arthropods in legal investigations. CRC, Boca Raton,
418p.
Goff, M.L., C.B. Odom & M. Early. 1986. Estimation of
postmortem interval by entomological techniques: a case
study from Oahu, Hawaii. Bull. Soc. Vector Ecol. 11:
242-246.
Gomes, L., C.J. Von Zuben & J.S. Govone. 2002.
Comportamento da dispersão larval radial pós-alimentar
em moscas-varejeiras do gênero Chrysomya (Diptera:
Calliphoridae): Busca por novas fontes de alimento.
Entomol. Vect. 9: 115-132.
Gomes, L. & C.J. Von Zuben. 2003a. Distribuição larval
radial pós-alimentar em Chrysomya albiceps (Wied.)
(Diptera: Calliphoridae): Profundidade, distância e peso
de enterramento para pupação. Entomol. Vect. 10: 211-
222.
Gomes, L. & C.J. Von Zuben. 2003b. Dispersão larval pós-
alimentar em Chrysomya megacephala (F.) (Diptera:
Calliphoridae): Profundidade, distância e peso de
enterramento para pupariação. Biosc. J. 18: 67-76.
January-February 2006 Neotropical Entomology 35(1)
Gomes, L. & C.J. Von Zuben. 2004a. Dispersão larval radial
pós-alimentar em Lucilia cuprina (Diptera,
Calliphoridae): profundidade, peso e distância de
enterramento para pupação. Iheringia. Série Zoologia
94: 135-138.
Gomes, L. & C.J. Von Zuben. 2004b. Efeito da temperatura
na profundidade de enterramento de larvas de
Chrysomya megacephala (Wied.) (Diptera:
Calliphoridae). Entomol. Vect. 11: 551-557.
Gomes, L. & C.J. Von Zuben. 2004c. Insetos ajudando a
desvendar crimes: Entomologia forense. Ciência Hoje
208: 28-31.
Grassberger, M. & C. Frank. 2004. Initial study of arthropod
succession on pig carrion in a central European urban
habitat. J. Med. Entomol. 41: 511-523.
Grassberger, M. & C. Reiter. 2001. Effect of temperature
on Lucilia sericata (Diptera: Calliphoridae)
development with special reference to the isomegalen-
and isomorphen-diagram. Forensic Sci. Int.120: 32-36.
Greenberg, B. 1985. Forensic entomology: Case studies.
Bull. Entomol. Soc. Am. 31: 25-28.
Greenberg, B. 1990. Nocturnal oviposition behaviour of blow
flies (Diptera: Calliphoridae). J. Méd. Entomol. 27: 807-
810.
Greenberg, B. 1991. Flies as forensic indicators. J. Med.
Entomol. 28: 565-577.
Greenberg, B. & J.C. Kunich. 2002. Entomology and the
law: Flies as forensic indicators. Cambridge University
Press, Cambridge, 306p.
Hall, R.D. 2001. Perceptions and status of forensic
entomology, p.1-15. In J.H. Byrd & J.L. Castner (eds.),
Forensic entomology: The utility of arthropods in legal
investigations. CRC, Boca Raton, 418p.
Hauser, G. 1926. Ein Beitrag zum Madenfraß an
menschlichen Leichen. Dtsch. Z. Gesamte. Gerichtl.
Med. 7: 179-192.
Higley, L.G. & N.H. Haskell. 2001. Insect development and
forensic entomology, p.287-302. In J.H. Byrd & J.L.
Castner (eds.), Forensic entomology: The utility of
arthropods in legal investigations. CRC, Boca Raton, 418p.
Introna, F., B.M. Altamura, A. Dell’Erba & V. Dattoli. 1989.
Time since death definition by experimental
reproduction of Lucilia sericata cycles in growth
cabinet. J. Forensic Sci. 34: 478-480.
Introna, F. & C.P. Campobasso. 2000. Forensic dipterology,
p.793-846. In Papp, L. & B. Darvas (eds.), Contributions
to a manual of palearctic diptera. 1. General and applied
dipterology. Science Herald, Budapest, 978p.
Introna, F., C.P. Campobasso & A. Difazio. 1998. Three
case studies in forensic entomology from Southern Italy.
J. Forensic Sci. 43: 210-214.
Introna, F., C.P. Campobasso & M.L. Goff. 2001.
9
Entomotoxicology. Forensic Sci. Int. 120: 42-47.
Jirón, L.F. & M.E. Solano. 1988. Notes on the eclectical
feeding preferences of the black soldier fly Hermetia
illuscens (Diptera: Stratiomyidae) in Costa Rica.
Brenesia 30: 101-103.
Jirón, L.F., L.G. Vargas & E. Vargas-Alvarado. 1982. Four
muscoid flies (Sarcophagidae and Muscidae) associated
with human cadavers in Costa Rica. Brenesia 21: 1-5.
Kamal, A.S. 1958. Comparative study of thirteen species of
sarcosaprophagous Calliphoridae and Sarcophagidae
(Diptera). I. Bionomics. Ann. Entomol. Soc. Am. 51:
261-270.
Kaneshrajah, G. & B. Turner. 2004. Calliphora vicina larvae
grow at different rates on different body tissues. Int. J.
Legal Med. 118: 242-244.
Knight, B. 1991. Forensic pathology. Edward Arnold,
London, 636p.
Leclercq, M. 1983. Entomologie et medicine legale; datation
de la mort, observation indite. Rev. Med. Liege 38: 735-
738.
Leclercq, J. & M. Leclercq. 1948. Donnees bionomiques
pour Calliphora erythrocephala (Meigen) et cas
d’application la medecine legale. Bull. Soc. Entomol.
Fr. 53: 101-103.
Lord, W.D., E.P. Catts, D.A. Scarboro & D.B. Hadfield.
1986. The green blow fly, Lucilia illustris (Meigen), as
an indicator of human postmortem interval: A case of
homicide from Fort Lewis, Washington. Bull. Soc.
Vector Ecol. 11: 271-275.
Mann, R.W., W. M. Bass & L. Meadows. 1990. Time since
death and decomposition of the human body: Variables
and observations in case and experimental field studies.
J. Forensic Sci. 35:103-111.
Marchenko, M.J. 1980. Classifying of cadaveric
entomofauna. Biology of flies: The forensic medical role.
Sud-Med Ekspert 23: 17-20.
Mariluis, J.C. 1983. Presencia del genero Chrysomya
Robineau-Desvoidy, 1830 en la region Neotropical.
(Calliphoridae, Chrysomyiinae, Chrysomyiini) Rev. Soc.
Entomol. Argentina 42: 141-142.
Mariluis, J.C., M.C. Lagar & E.J. Bellegarde. 1989.
Disemination de enteroparasitos por Calliphoridae (Insecta,
Diptera) Mem. Inst. Oswaldo Cruz 84: 349-351.
McKnight, B.E. 1981. The washing away of wrongs:
Forensic medicine in thirteenth-century China.
University of Michigan, Ann Arbor, 181p.
Mégnin, P. 1894. La faune de cadaveres. Application de
l’entomologie a la médicine légale, Encyclopdie
scientifique des Aides-Mémoire, Masson, Paris
Gauthier-Villars, Paris, 214p.
Mende, L.J.K. 1829. Ausführliches Handbuch der
gerichtlichen Medizin für Gesetzgeber, Rechtsgelehrte,
10 Gomes & Zuben - Forensic Entomology and Main Challenges in Brazil
Aerzte und Wundaerzte, Teil 5.
Merrit, R.W. & J.R.Wallace. 2001. The role of aquatic insects
in forensic investigations, p.177-222. In Byrd, J.R. &
J.L. Castner (eds.), Forensic entomology: The utility of
arthropods in legal investigations. CRC, Boca Raton,
418p.
Moura, M.O., C.J.B de Carvalho & E.L.A. Monteiro-Filho.
1997. A preliminary analysis of insects of medico-legal
importance in Curitiba, State of Paraná. Mem. Inst.
Oswaldo-Cruz 92: 269-274.
Nolte, K.B., R.D. Pinder & W.D. Lord.1992. Insect larvae
used to detect cocaine poisoning in a decomposed body.
J. Forensic Sci. 37: 1179-1185.
Nuorteva, P. 1959a. Studies on the significance of flies in
the transmission of poliomyelitis. III. The composition
of the blow fly fauna and the activity of the flies in
relation to the weather during the epidemic season of
poliomyelitis in south Finland. Ann. Entomol. Fenn.
25: 137-162.
Nuorteva, P. 1959b. Studies on the significance of flies in
the transmission of poliomyelitis. IV. The composition
of the blow fly fauna in different part of Finland during
1958. Ann. Entomol. Fenn. 25: 137-162.
Nuorteva, P. 1977. Sarcosaprophagous insects as forensic
indicators, p.1072-1095. In G.C. Tedeshi, W.G. Eckert
& L.G. Tedeshi (eds.), Forensic medicine: A study in
trauma and environmental hazards, v. 2, Saunders,
Philadelphia, 1637p.
Nuorteva, P., M. Isokoski & K. Laiho. 1967. Studies on the
possibilities of using blowflies (Dipt.) as medicolegal
indicators in Finland. Ann. Entomol. Fenn. 33: 217-
225.
Nuorteva, P. & S.L. Nuorteva 1982. The fate of mercury in
sarcosaprophagous flies and in insects eating them.
Ambio 11: 34-37.
O’Brien, C., B. Turner. 2004. Impact of paracetamol on the
development of Calliphora vicina larval development.
Int. J. Legal Med. 118: 188-189.
Orfila, M.J.B. & C.A. Lesueur. 1831. Traité dês exhumations
juridiques. Paris.
Oliveira-Costa, J., C. Mello-Patiu & S. M. Lopes. 2001a.
Dipteros muscoides associados com cadáveres humanos
na cena da morte no estado do Rio de Janeiro. Bol.
Museu Nacional 464: 1-6.
Oliveira-Costa, J., C. Melo-Patiu & S. M. Lopes. 2001b.
Influência dos diferentes fatores na freqüência de
dípteros muscóides em cadáveres humanos no Rio de
Janeiro. Bol. Museu Nacional 470: 1-10.
Oliveira-Costa, J. 2002. Entomologia forense - Os insetos a
serviço da lei. Cad. Direito Univ. Castelo Branco, Rio
de Janeiro, 1: 41-51.
Oliveira-Costa, J. 2003. Entomologia forense: Quando os
insetos são vestígios. Ed. Millennium, Campinas, 257p.
Oliveira-Costa, J. & C.A. Mello-Patiu. 2004. Estimation of
PMI in homicide investigation by the Rio de Janeiro
Police. Anil Aggrawal’s Internet J. Forensic Med.
Toxicol 5: 40-44.
Oliveira-Costa, J. & C. A. Mello-Patiu. 2005. In press.
Association analysis of carrion arthropods to estimate
post mortem interval. J. Forensic Sci (in press).
Payne J.A. 1965. A summer carrion study of the baby pig
Sus scrofa Linnaeus. Ecology 46: 592-602.
Payne, J.A. & E.W. King. 1972. Insect succesion and
decomposition of pig carcasses in water. J. Ga. Entomol.
Soc. 7: 153-162.
Payne J.A., E.W. King & G. Beinhart. 1968. Arthropod
succession and decomposition of buried pigs. Nature
219: 1180-1181.
Pessôa S.B. & F. Lane. 1941. Coleópteros necrófagos de
interésse médico-legal. Ensáio monográfico sôbre a
familia Scarabaeidae de São Paulo e regiões vizinhas.
Arq. Zool. Est. São Paulo, II. Rev. Mus. Paulista 25:
389-504.
Povolný D. & Y. Verves. 1997. The flesh-flies of central
Europe. Spixiana Suppl 24: 1-260.
Price, P.W. 1997. Insect ecology. Wiley, New York, 874p.
Reinhard, H. 1882. Beiträge zur Gräberfauna. Verh Kaiserl-
Königl Zool-Bot Ges Wien 31: 207-210.
Reiter, C. 1984. Zum Wachstumsverhalten der Maden der
blauen Schmeißfliege Calliphora vicina. Z. Rechtsmed
91: 295-308.
Reiter, C. & G. Wolleneck. 1982. Bemerkungen zur
Morphologie forensisch bedeutsamer Fliegenmaden. Z.
Rechtsmed 89: 197-206.
Reiter, C. & G. Wolleneck. 1983. Zur Artbestimmung der
Maden forensisch bedeutsamer Schmeißfliegen. Z
Rechtsmed 90: 309-316.
Rodriguez, W.C. & W.M. Bass. 1983. Insect activity and its
relationship to decay rates of human cadavers in East
Tennessee. J. Forensic Sci. 28: 423-432.
Schmitz, H. 1928. Phoriden in doodkisten. Natuurhist
Maandbl 17: 150-153.
Schoenly, K., M.L. Goff, J.D. Wells & W.D. Lord. 1996.
Quantifying statistical uncertainty in succession-based
entomological estimates of the postmortem interval in
death scene investigations: A simulation study. Am.
Entomol. 42: 106-112.
Schoenly, K. & W. Reid. 1987. Dynamics of heterotrophic
succession in carrion arthropod assemblages: Discrete
series or a continuum of change? Oecologia 73: 192-
202.
Smith, K.E. & R. Wall. 1997a. Asymmetric competition
between larvae of the blowflies Calliphora vicina and
Lucilia sericata in carrion. Ecol. Entomol. 22: 467-474.
January-February 2006 Neotropical Entomology 35(1)
Smith, K.E. & R. Wall. 1997b. The use of carrion as breeding
sites by the blowfly Lucilia sericata and other
Calliphoridae. Med. Vet. Entomol. 11: 38-44.
Smith, K.G.V. 1986. A manual of forensic entomology.
British Museum, London, 205p.
Souza, A.M. 1994. Sucessão entomológica na decomposição
de carcaça animal, com ênfase nas famílias
Calliphoridae e Sarcophagidae (Diptera). M.Sc.
Dissertation, Universidade Estadual de Campinas, SP,
XI + 96p.
Sperling, F.A., G.S. Anderson & D.A. Hickey. 1994. A DNA-
based approach to the identification of insect species
used for postmortem interval estimation. J. Forensic Sci.
39: 418-427.
Stafford, F. 1971. Insects of a medieval burial. Sci Anthropol
7: 6-10.
Stevens, J.R. & R. Wall. 1996. Species, sub-species and
hybrid populations of the blowflies Lucilia cuprina and
Lucilia sericata (Diptera: Calliphoridae). Proc. R. Soc.
London B 263: 1335-1341.
Tessmer, J.W., C.L. Meek & V.L. Wright. 1995. Circadian
patterns of oviposition by necrophilous flies (Diptera:
Calliphoridae). Southwest. Entomol. 24: 439-445.
Thyssen, P.J., A.C. Lessinger, A.M.L. Azeredo-Espin & A.X.
Linhares. 2005. The value of PCR-RFLP molecular
markers for the differentiation of immature stages of
two necrophagous flies (Diptera: Calliphoridae) of
potential forensic importance. Neotrop. Entomol. 34:
777-783.
11
Turchetto, M. & S. Vanin. 2004. Forensic entomology and
climatic change. Forensic Sci. Int. 146 (S1) S195- S199.
Von Zuben, C.J. 2002. The forensic entomologist’s role in
medico-criminal investigations. Holos Environment
(CD-ROM).
Von Zuben, C.J., R.C. Bassanezi, S.F. dos Reis, W.A.C. Godoy
& F.J. Von Zuben. 1996. Theoretical approaches to forensic
entomology: I. Mathematical model of postfeeding larval
dispersal. J. Appl. Entomol. 120: 379-382.
Wall, R & M.L. Warnes. 1994. Responses of the sheep
blowfly Lucilia sericata to carrion odour and carbon
dioxide. Entomol. Exp. Appl. 73: 239-246.
Wells, J.D. & F.A. Sperling. 1999. Molecular phylogeny of
Chrysomya albiceps and C. rufifacies (Diptera:
Calliphoridae). J. Med. Entomol. 36: 222-226.
Wells, J.D., F. Introna Jr., G. Di Vella, C.P. Campobasso, J.
Hayes & F.A. Sperling. 2001. Human and insect
mitochondrial DNA analysis from maggots. J. Forensic
Sci. 46: 685-687.
Wells, J.D. & L. King. 2001. Incidence of precocious egg
development in flies of forensic importance
(Calliphoridae). Pan-Pac Entomol 77: 235-239.
Wells, J.D. & L.R. LaMotte. 1995. Estimating maggot age
from weight using inverse prediction. J. Forensic Sci.
40: 585-590.
Wolff, M. & A. Uribe. 2000. Sucesión de insectos carroñeros
em cerdo blanco (Sus scrofa). Mem. Congr. Soc. Col.
Entomol. 27: 177-185.
Received 29/III/05.