Forensic Science International 226 (2013) 41–45

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Forensic Science International

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

Necrophagous beetles associated with carcasses in a semi-arid environment

in Northeastern Brazil: Implications for forensic entomology
Ana C.G. Mayer, Simão D. Vasconcelos *
Necrophagous Insects Research Group, Department of Zoology, Universidade Federal de Pernambuco, Av. Prof. Moraes Rego, Recife, Pernambuco, 50.670-420, Brazil

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

Article history: Data on the ecology and bionomics of necrophagous beetles are scarce in tropical countries despite their
Received 6 July 2012 relevance in forensic investigations. We performed a survey on the diversity and temporal pattern of
Received in revised form 27 October 2012 colonization of beetles on pig carcasses in a fragment of dry forest in northeastern Brazil. We collected
Accepted 30 November 2012
1550 adults of diverse feeding habits from 12 families, of which 96% had necrophagous and/or copro-
Available online 9 February 2013
necrophagous habits and belonged to four families: Dermestidae, Scarabaeidae, Cleridae and Trogidae.
Three species, Dermestes maculatus, Necrobia rufipes and Omorgus suberosus are reported for the first time
Keywords:
with an expanded geographical distribution that includes the semi-arid region in Brazil. Adult beetles
Forensic entomology
Coleoptera
were collected as early as 24 h after death. One endemic species, Deltochilum verruciferum, stood out in
Deltochilum verruciferum terms of numerical dominance and temporal occurrence during different stages of decomposition. Its
Necrobia rufipes intimate association with carrion emphasizes their potential role in forensic entomology in the region.
Caatinga ß 2012 Elsevier Ireland Ltd. All rights reserved.
Homicide

1. Introduction entomology, especially in countries located in the tropics, some

Coleoptera (beetles) comprises the most diverse of all living
groups, from which several families exhibit intimate association
with carrion as a source of food, nesting sites and an extension of
their habitat [1,2]. Species of diverse feeding habits have been
recorded on carcasses and cadavers, including not only necropha-
gous, but also coprophagous, predator, saprophagous and omni-
vore species [3,4]. Members of the families Cleridae, Silphidae,
Scarabaeidae and Dermestidae have been demonstrated to show
necrophagous and copro-necrophagous habits and have potential
use in forensic entomology [5]. In recent years, information on the
diversity, geographical distribution and life cycle of necrophagous
beetles has been incorporated into forensic investigations, ranging
from the estimation of post-mortem interval [5,6] to the detection
of insect-mediated changes at the scene of death [7] and to the
study of the rates of cadaver decomposition [8].
Despite this, knowledge on the ecology of forensically impor-
tant beetles is remarkably low when compared to dipteran species;
blow flies and flesh flies, for example. The scarcity of information
on bionomics and biogeography is still a major obstacle to the use
of coleopterans in a reliable, quantitative basis in forensic
* Corresponding author. Tel.: +55 81 21268353; fax: +55 81 21268353.
E-mail addresses: simaovasconcelos@yahoo.com.br, simao@ufpe.br
(S.D. Vasconcelos).
of which are exposed to high rates of lethal crimes.
Violent crimes in Brazil are amongst the highest in the world:
from 1998 to 2008, the absolute number of homicides increased
101.5% in the northeastern region of Brazil [9]. Although violent
deaths had been traditionally associated with large, urban
environments, small rural municipalities have suffered from
increasing homicide rates, partially related to drug production
and trafficking. This is particularly true for cities located in areas
exposed to sudden economic development in the semi-arid region
in Northeastern Brazil, where the use of biological information to
provide evidence in homicide investigations is still incipient.
In this context, we aimed to identify beetle species of forensic
relevance in a semi-arid region of the Brazilian countryside.
Specifically we addressed the following questions: (i) How
diverse is the coleopteran community in a dry forest biome
subjected to harsh conditions of temperature and water regime?
(ii) Are there temporal differences in carcass colonization by
different species? (iii) Are there endemic species that could be
used as indicators of site of death? Specifically, we tested two
hypotheses: (1) Decomposition rates in environments subjected
to extreme temperatures may favor faster colonization by
necrophagous beetles during the early stages of decomposition;
(2) the unique environmental characteristics of the Brazilian dry
forest would support a specific coleopteran fauna, which could
be employed to validate certain beetle species as indicators of
place of death.

0379-0738/$ – see front matter ß 2012 Elsevier Ireland Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.forsciint.2012.11.019
42 A.C.G. Mayer, S.D. Vasconcelos / Forensic Science International 226 (2013) 41–45
Fig. 1. Location of the dry forest (caatinga) biome, with emphasis on the experimental
area in Pernambuco State, northeastern Brazil.
2. Materials and methods
2.1. Area of study and experimental design
The experiment was performed in a conserved fragment of caatinga, a type of
seasonally dry forest endemic to Brazil (Fig. 1), which covers an estimated area of
(6–9)  105 km2 and it is characterized by semi-arid climate, high potential
evapotranspiration throughout the year (1500–2000 mm/year), and low and erratic
rainfall (300–1000 mm/year) [10]. Severe droughts lasting 4–5 years are also
common. The study took place in the municipality of Serra Talhada (078590 S;
388170 W, altitude 429 m), State of Pernambuco, Northeastern Brazil, in a private
farm (ca. 700 ha) maintained for conservation purposes. The climate is defined as
hot semiarid (BSh – according to the Köppen climate classification). The rainy
season occurs from November to April; the mean annual rainfall is 431 mm and
daily temperatures usually reach 40 8C [11].
Vegetation is characterized by the presence of shrub native species typical to dry
biomes, with the predominance of Cactaceae and sparsely distributed trees (e.g.,
Fabaceae, Anacardiaceae). Local vertebrate fauna comprises mostly small and
medium sized species of lizards, anurans, snakes, wild dogs, marmosets, armadillos
and birds.
Field trials took place between March and May 2010. As a model we used a male
pig (Sus scrofa L.) of ca. 15 kg, killed by a concussion on the occipital region, in order
to mimic violent death. The animal was placed inside a metal cage
(80 cm  60 cm  50 cm) to prevent access of large scavengers. The top of the
cage was removable to allow sampling. The cage was placed in a way that the
extremities of the body would be in contact with the soil while the middle part of
the carcass was placed on a ditch in which a tray (30 cm  70 cm  10 cm)
containing sawdust was placed under the cage to collect beetles.
All procedures were approved by the Committee of Ethics in Animal
Experimentation at the Universidade Federal de Pernambuco. Three replicates of
the experiment were performed simultaneously using carcasses of similar weight,
color and age, separated from each other by a distance of 1 km.
2.2. Insect collection and identification
Two sampling methods were used: in the active collection, the whole carcass
was thoroughly inspected for 20 min during which adult beetles were collected
using soft forceps, with minimum disturbance. The passive method consisted of
removal of beetles caught in the tray. Sampling by both methods was performed at
the same time of the day (9.00–11:00) by the same operator to minimize variation
in sampling efficiency.
Collections began 24 h after death and were repeated at 24 h intervals until the
eleventh day, after which collection took place at 48 h intervals, until the complete
skeletonization of the carcass. Each carcass was observed and photographed daily in
order to characterize the stages of decomposition. All adult beetles were collected
irrespective of their feeding habits and were stored in 70% alcohol in vials until
identification in the laboratory, using taxonomic keys [1,12–16].
2.3. Data analysis
The following variables were analyzed: species richness, abundance, feeding
habit, and the percentage frequency of occurrence ([No. of samples containing a
given species/total number of samples]  100), all of which were related to the
collecting method and stage of decomposition. Chi-square tests were performed to
compare abundance of necrophagous and non-necrophagous species using a 5%
confidence level. Shannon’s diversity indices were calculated for each collecting
method and each stage of decomposition. A correspondence analysis was
performed to assess interactions between the most abundant species and the
decomposition stage using the Program MVSP 3 (Multivariate Statistical Package).
3. Results
3.1. Stages of decomposition and environmental conditions
During the experiment, mean air temperature was 30.6 8C and
relative humidity was 60.0%; accumulated precipitation was
12.7 mm throughout the whole experiment. Decomposition in
all replicates occurred in five distinct stages: fresh (1-day
duration); bloated (2-day duration); active decay (4 days);
advanced decay (5 days) and finally, the dry stage which lasted
12 days. There was no difference in the duration of each phase
among the three replicates (P > 0.05; d.f. = 2).
3.2. Overall pattern of diversity and feeding habits
In total, 1550 adult beetles from 24 species belonging to nine
families were collected (Table 1). There was no significant
difference among the number of families and specimens among
the three replicates (P > 0.05; d.f. = 2). Most species were
registered in low abundance, with the exception of Deltochilum
verruciferum (762 adults, considering the sum of the three
replicates), Necrobia rufipes (269), Dermestes maculatus (241) and
Omorgus suberosus (108). There was no significant difference in the
number of species collected by active search when compared to
tray captures (x2 = 1.58; P > 0.05; d.f. = 1). However, the number
of individuals collected in the tray was significantly higher than
that from active collection (x2 = 248.0; P < 0.05; d.f. = 1), due to
the overwhelming presence of D. verruciferum (over 700 individu-
als collected in trays).
Beetle species with varying feeding habits were associated with
the decomposing carcasses, including herbivore and omnivore
species (Table 1). No difference in the number of feeding categories
was observed between the two collecting methods. In contrast,
there was a dominance of necrophagous and copro-necrophagous
individuals for both active (x2 = 392.1; d.f. = 1; P < 0.0001) and
tray collecting methods (x2 = 879.8; d.f. = 1; P < 0.0001) (Fig. 2).
Shannon’s index (mean  s.d.) of diversity did not differ between
the two collecting methods (1.34  0.05 for active collection;
1.2  0.13 for tray) (P = 0.616; d.f. = 1). A closer similarity in the
pattern of occurrence – both in terms of abundance and temporal
colonization – was observed between D. maculatus and N. rufipes.
3.3. Beetle diversity and decomposition stages
The number of species on the carcasses varied according to the
stage of decomposition, from three (fresh stage) to 16 (dry)
(Table 2). Both necrophagous and non-necrophagous species were
registered throughout the entire decomposition. Fewer individuals
were collected at the first day post-death, although in the fresh
stage the relative proportion of necrophagous species was higher
than at the other phases. The relative frequency of necrophagous
individuals did not vary significantly among different decomposi-
tion stages (P > 0.05; d.f. = 4) (Table 2). Shannon’s diversity indices
were lower during the initial stages of decomposition (0.41 at fresh
stage) and increased to 1.69 at the dry stage.
The temporal colonization differed between species: D.
verruciferum and O. suberosus appeared within the first 24 h post
death and continued to exploit the carcass until skeletonization
(Table 3). In contrast, N. rufipes and D. maculatus only appeared at
the beginning of the active decay stage and their abundance did not
varied markedly during the entire decomposition. No marked
differences in the abundance of each necrophagous or copro-
necrophagous species according to the stage of decomposition
 A.C.G. Mayer, S.D. Vasconcelos / Forensic Science International 226 (2013) 41–45 43

Table 1
Diversity and abundance of adult Coleoptera collected on decomposing pig carcasses in a semi-arid environment in Northeastern Brazil according to the collecting method
and main feeding habit; total numbers represent the summation of three replicates.

Family Species Active collection Tray Main feeding habita

Scarabaeidae Ateuchus carbonarius (Harold, 1868) 24 73 CN

Canthidium manni Arrow, 1913 2 C
Canthon sp. 3 C
Coprophanaeus cyanescens (d’Ousoufieff, 1924) 2 C
Deltochilum verruciferum Felsche, 1911 29 733 CN
Dichotomius nisus (Olivier, 1789) 14 C
Dichotomius geminatus (Arrow, 1913) 1 C
Trichillum externepunctatum Borre, 1886 4 4 C

Histeridae Atholus sp. 2 P

Euspilotus (Hesperosaprinus) sp. 4 P
Euspilotus sp. 1 P
Hololepta sp. 1 P
Omalodes foveola Erichson, 1834 1 P
Omalodes planifrons Marseul, 1853 1 P
Xerosaprinus sp. 7 7 P

Carabidae Galerita sp. 12 P

Arthrostictus speciosus (Drury, 1829) 1 P
Clivina sp. 1 P

Curculionidae Scolytinae sp. 2 O

Bostrichidae Bostrichidae sp. 2 O
Cleridae Necrobia rufipes (De Geer, 1775) 222 47 N
Dermestidae Dermestes maculatus (De Geer, 1774) 147 94 N
Lymexylidae Lymexylidae sp. 1 O
Trogidae Omorgus suberosus Fabricius, 1775 24 84 N

Total of species 24 12 19
Total of individuals 1550 465 1085
Total of feeding categories 6 6
a
CN, copro-necrophagous; C, coprophagous; N, necrophagous; P, predator; O, others. Consulted bibliography [1,3,12–14,29,30,34–41].

were detected, with the exception of D. verruciferum which was
significantly more abundant during the bloated phase (x2 = 465.8;
d.f. = 4; P < 0.0001). O. suberosus was the least abundant of the
species classified as necrophagous (Table 3). The correspondence
analysis (Fig. 3) revealed the association of O. suberosus and D.
verruciferum with early and intermediate stages of decomposition
in the caatinga biome, while D. maculatus and N. rufipes were more
strongly related to advanced and dry stages.
4. Discussion
The stages of decomposition described in this experiment
appear to be similar to what has been previously described in arid
and semi-arid environments [17], although mummification was
Fig. 2. Species richness vs. abundance, according to the feeding habit, of
coleopterans captured by different collection methods in the dry forest
(caatinga) located in Northeastern Brazil.
not observed. Galloway et al. [17] used official reports on human
corpses found in semiarid regions to establish a pattern of
decomposition, and reinforced the similarity on the sequence
and duration of decomposition stages between pigs and humans.
The diversity of coleopteran species associated with decom-
posing animal carcasses in the Brazilian semi-arid region does not
seem to differ markedly from what has been observed in semi-
desert areas [18], tropical rain forests [19], temperate forests [20],
agroecosystems and urban areas [21,22]. Feeding habits varied
from predatory (Histeridae and Carabidae) to necrophagous
species (Dermestidae and Cleridae), as well as omnivores and
also species that, despite being associated with coprophagy, may
include decomposing animal matter in their diets, such as Ateuchus
carbonarius (Scarabaeidae). Predators, such as Galerita sp. (Car-
abidae), are attracted to the carcass by the high availability of prey,
particularly the large amount of blowfly larvae, and are common at
early stages of decomposition, as evidenced by the correspondence
analysis (Fig. 3).
Surprisingly, specimens of rove beetles (Staphylinidae) were
not collected despite the abundance of these insects in carcasses
and cadavers in diverse ecosystems, where they have been
reported to prey upon larvae of several species of necrophagous
Diptera [21,23–26]. Similarly, species of Silphidae were not found
in the experiment, in spite of the necrophagous habits previously
registered for beetles from this family and of their dominance in
field studies performed in Southern Brazil [26].
Clearly, necrophagous and copro-necrophagous species were
dominant in the assemblage, as they comprised nearly 96% of all
specimens collected. The high proportion of necrophagous and
copro-necrophagous specimens at early stages of decomposition
diverges, to some extent, from several records in the literature that
suggest that coleopterans do not occupy the cadaver until later in
the decomposition. However, further studies have revealed that
beetles can actively colonize a carcass in the first days after death
44 A.C.G. Mayer, S.D. Vasconcelos / Forensic Science International 226 (2013) 41–45

Fig. 3. Patterns of colonization of pig carcasses by Coleoptera species in a dry forest fragment (caatinga) in Northeastern Brazil, according to the stages of decomposition, as
produced by correspondence analysis.

Table 2
Diversity and occurrence of necrophagy among Coleoptera collected on pig carcasses at different decomposition stages in a semi-arid forest in Brazil. Values relate to
combined data from three replicates.

Fresh Bloated Active decay Advanced decay Dry/remains

Total number of species 3 9 13 11 16

% of copro-necrophagous/necrophagous species 67.7% 33.3% 53.8 50.0% 37.5%
Number of individuals 28 442 385 270 425
% of copro-necrophagous/necrophagous individuals 89.3% 96.4% 85.2% 87.4% 85.6%
Shannon’s index (mean  SD) 0.17  0.15 0.24  0.11 0.63  0.08 0.62  0.02 0.65  0.09

[3,27]. The similar frequency of necrophagous/copro-necropha-
gous specimens across the different stages of decomposition
suggests that in environments exposed to high temperatures – and
probably to faster decomposition – beetles tend to exploit this
spatially and temporally finite resource comparatively earlier than
in temperate zones. This supports the hypothesis that under local
conditions necrophagous and copro-necrophagous beetles can be
forensically important during the initial stages of cadaver
decomposition.
The higher efficiency of the passive collection (tray) reflects the
longer periods of catchment (24 h intervals) when compared to
20 min of active search. However, given the similar pattern of
richness and feeding habits in species caught by both methods, the
practice of manual collection of larvae and adults at homicide
crime scenes would probably be representative of the coleopteran
diversity. Our results suggest that in the dry forest of Northeastern
Brazil even a short collection period can be effective to sample a
significant proportion of the diversity of necrophagous coleopter-
ans present in the ecosystem. However, it is important to stress out
that the pattern of diversity is also a reflection of the efficiency of
each method, as copro-necrophagous beetles may remain on trays
for longer periods than species that feed exclusively on carcasses.
The unfavorable environmental conditions peculiar to the
caatinga biome, namely water stress, high temperature, low
humidity and frequent periods of severe drought, was expected
to be associated with a specialized assemblage of endemic
necrophagous beetles that would be adapted to the marked
seasonal variations in the availability of resources. This hypothesis
was rejected, since only one species, D. verruciferum, appeared to
be specific to the habitat [28–31]. The two other most abundant
species, D. maculatus and N. rufipes occur commonly on animal
carcasses in several countries [21,22,32,33]. Our results represent a
contribution to the knowledge of the geographical distribution and
habitat use of these two forensically important species, since this is
their first record in the Brazilian semi-arid region.
N. rufipes and D. maculatus are by far the most studied
coleopteran species for forensic purposes, and they both have
been used to help in the estimation of post-mortem interval in
other countries [5,6,27]. In northeastern Brazil, the forensic
relevance of these species could only be assessed provided that

Table 3
Occurrence and class of abundance of copro-necrophagous/necrophagous beetles associated with pig carcasses in a semi-arid environment in Northeastern Brazil, according
to the day post-mortem. Values relate to combined data from three replicates.

Family Species Stage of decomposition/days post-mortem

Fresh Bloated Active decay Advanced decay Dry/remains

01 02 03 04 05 06 07 08 09 10 12 14 16 18 20 22 24 26

Scarabaeidae Ateuchus carbonarius – –                

Deltochilum verruciferum                  

Cleridae Necrobia rufipes – – –               

Dermestidae Dermestes maculatus – – –               
Trogidae Omorgus suberosus          -   -     

(–) absent; () <10; () 11–50; () 51–100; () >100.
 A.C.G. Mayer, S.D. Vasconcelos / Forensic Science International 226 (2013) 41–45
data on their life cycle, such as the duration of each larval stage, is
obtained under local or similar conditions. For example, it is
common to register a time interval of several years between
rainfalls in large areas in the caatinga, a fact that should trigger
variations in the bionomics of local species. It is also known that
temperature affects the development of dermestids and clerids
[27] but the exact duration of each larval stage of these species
when exposed to different temperatures is still poorly understood.
D. verruciferum is endemic to the Brazilian caatinga and its
surroundings and its previous records were based mostly on pitfall
traps baited with human feces [29,34]. This is the first reference of
its colonization of animal carcasses, adding necrophagy to its
registered feeding habits. This behavior supports the forensic
relevance of this species as a suitable indicator of place of death.
5. Conclusions
This is, to our knowledge, the first inventory of necrophagous
beetle species in a semi-arid Neotropical environment. The
continued presence over time of cosmopolitan species, such as
N. rufipes and D. maculatus, reinforces their relevance to forensic
studies, although the lack of detailed information on their life
cycles under the extreme conditions found in the caatinga hinders
the practical use of these species in the estimation of the minimum
post-mortem interval. In contrast, the discovery of an endemic
copro-necrophagous species, D. verruciferum, may help forensic
scientists to associate its presence with a particular biome, since it
has not yet been registered as a necrophagous species elsewhere.
Acknowledgements
We thank Capes (Ministry of Education, Brazil) for the
scholarship to A.C.G.M., Homembom Magalhaes for access to the
experimental area, Roberta Salgado for invaluable help in the field,
Carla Lima Bicho (UEPB, Brazil) and Fernando Vaz-de-Mello (UFMT,
Brazil) for helping with species identification, Trevor Williams
(Inecol, Mexico) for critical reading of the manuscript and
Rivaneide Matias and Renata Akemi for logistical support. S.D.V.
has a productivity grant from CNPq (Ministry of Science and
Technology, Brazil).
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