the role of ants (hymenoptera: formicidae) in forensic entomology

THE ROLE OF ANTS (HYMENOPTERA: FORMICIDAE)
IN FORENSIC ENTOMOLOGY
Giovanni Ramón1, David A Donoso2
Instituto Nacional de Investigación en Salud Pública, Iquique N14-285 y Yaguachi, Quito, Ecuador.1
Museo de Colecciones Biológicas MUTPL, Departamento de Ciencias Naturales,
Universidad Técnica Particular de Loja; Loja, Ecuador.2
[email protected]
ABSTRACT
Ants (Hymenoptera: Formicidae) are an important component of insect biodiversity, and their study may
help forensic entomologists to solve criminal cases. We review published literature and ecological principles
that may guide forensic entomologists in the usage of ants as evidence with forensic value. Because ants are
top predators in litter environments, their activities can affect decomposition processes and influence on
the estimation of the post-mortem interval. We also revise artifacts caused by ants on decomposing bodies
(e.g. biting, stinging and modification of blood patterns).
KEYWORDS: BIODIVERSITY, biting, post-mortem interval, stinging
RESUMEN
Las hormigas (Hymenoptera: Formicidae) son un componente importante de la biodiversidad de insectos
y su estudio puede ayudar a los entomólogos forenses a resolver casos criminales. Nosotros revisamos
la literatura y principios ecológicos que pueden guiar a los entomólogos forenses a usar hormigas como
evidencia con valor forense. Las hormigas son depredadores en la hojarasca, por lo tanto sus actividades
pueden afectar los procesos de descomposición e influenciar la estimación del intervalo post mortem.
También, nosotros revisamos los artefactos causados por hormigas en cuerpos en descomposición (p.ej.
mordeduras, picaduras y modificación de los patrones en manchas de sangre).
PALABRAS CLAVES: BIODIVERSIDAD, intervalo post mortem, mordeduras, picaduras.
INTRODUCTION
As a group, insects constitute the most successful
living organisms in the history of the planet. Insects
represent more than 50% of all the eukaryotic
species estimated to exist in the world (Grimaldi
and Engel, 2005), they live in all land masses of
the planet and perform a myriad of ecological
functions. Forensic entomology studies the role of
insects, and other arthropods, in legal issues (Hall
and Huntington, 2010), and thus seeks to apply a
vast amount of scientific knowledge about insects,
their distribution and their development to solve
crime. Ultimately, forensic entomology aids legal
and justice systems and provides relief to families
and societies.
Ants are an important and ubiquitous component of
the insect fauna on all terrestrial habitats, except the
poles, with about 140 million years of evolutionary
history (Moreau and Bell, 2013). There are currently
more than 13,000 ant species recognized worldwide
(Bolton, 2014). They play crucial ecological roles as
predators, soil engineers, seed dispersers, plant
symbionts, nutrient cyclers and more (Del Toro et
al., 2012). In tropical forests ants represent, together
with termites, a major proportion (i.e. 30%) of the
animal biomass (Fittkau and Klinge, 1973). At the
ground level they can reach densities of up to 23
nests per square meter (Kaspari, 1996a).
Most ants are beneficial to human beings. Some
species are used as food (DeFoliart, 1999), others
produce compounds with pharmaceutical and
biomedical applications (Reddy et al., 2011; Brown
et al., 2004), they are important in many cultures
around the world, and they provide ecological
services as seed dispersers, pollinators and
biological control agents (Lengyel et al., 2010; Van
Mele, 2008; Philpott and Armbrecht, 2006). Some
species, however, are detrimental for humans
when they establish nearby human settlings. Fire
ants, for example, frequently colonize electrical
equipment, attack livestock, cause infrastructure
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REMCB 36 pp. 19-26. 2015
damage in roads and buildings, and with their
painful stings are a nuisance for human activities
(Ascunce et al., 2011). Of special concern are
ants that invade hospitals, as they represent a
nuisance for patients and staff and are capable of
mechanically transporting bacteria (Rodovalho et
al., 2007; Moreira et al., 2005).
One overlooked utility of ants is their use on
forensic investigations. Traditionally, ants have
been considered minor actors on forensic studies,
well after flies and beetles (Byrd and Castner, 2010).
Ants, however, are important in forensic studies in
several ways. Here, we search in the literature for
the role of ants on forensic entomology, specifically
on medicocriminal entomology. First, we review
decomposition processes and how ants modify
them. We found that ant activities over corpses
can influence the estimation of the post-mortem
interval (PMI). Second, we explore the role of ants
as causing agents of human injuries and death.
Third, we list the artifacts ants may cause over
corpses. Several of these artifacts can confuse
forensic specialists and lead them to erroneous
conclusions. Fourth, we review cases where ants
can help determine the place of death. Across the
paper, we offer our personal observations and
suggestions on the ways forward and recognize the
underappreciated role of this fascinating group of
insects in forensic science.
Ants and decomposition processes
At macroscale, decomposition of organic material
can be explained as a function of temperature and
precipitation (Chapin et al., 2011). However, at
local scale differences in decomposition of organic
matter are due to substrate quality (e.g. soil),
material quality (e.g. plant material decomposes
much faster than animal material), and the
decomposer community (e.g. bacteria, fungi, and
the microorganism of the litter layer) (Chapin et al.,
2011). The role of litter predators (such as ants) on
the decomposition of plant material has been little
studied. Generally, ants play an important role
as predators of other arthropods in the leaf litter
layer (Wilson, 2005; Kaspari 1996b), and so it has
been shown that they may limit the decomposition
rates by predating on them and also by altering the
composition of the microbial community (Del Toro
et al., 2012; Wardle et al., 2011).
The importance of ants on the decomposing process
of animal and human carcasses has received less
attention. It is commonly recognized that ants are
attracted to arthropods that colonize corpses, not
the corpses themselves (Campobasso et al., 2009).
20
This hypothesis is mostly based in observations
from temperate regions where ants dwelling in the
corpses are attracted to fly eggs, larvae and pupae,
but not the flesh itself (Campobasso et al., 2009;
Barros et al., 2008). With some exceptions (ValdesPerezgasga et al., 2010), in tropical ecosystems most
reports only mention ants as casual visitors to
corpses that feed on arthropods and provide little
or none information of forensic relevance at all (e.g.
Bermúdez and Pachar, 2010).
Ants (Cephalotes atratus Linnaeus 1758, Crematogaster
sp.) however, were found to cut and carry skin and
muscle tissue of an experimental fresh pig carcass
located in Yasuní National Park in Orellana Province,
Ecuador, (Fig. 1) (Emilia Moreno, pers. comm.).
This behavior have been observed elsewhere (e.g.
Prado e Castro et al., 2014; and references therein)
Moreover, among the Ecuadorian Cofan ethnicity
(and possibly widespread in the Amazon basin)
ants of the genus Dolichoderus are usually used to
clean animal bones from their flesh (David Donoso,
pers. obs.). Regardless, ant activity can potentially
decrease (flesh removal) or increase (fly egg, larvae
and pupae removal) the estimation of the postmortem interval (Early and Goff, 1986)
Interactions of ants with other arthropods around
decomposing bodies
While they are capable of scavenging on the flesh,
the role of ants as predators over other arthropods
(and their eggs and larvae) has the most substantial
impact on forensic studies. In some instances the
predation rate of ants on flies, beetles and their eggs
might be so important that it can delay the onset of
initial colonization up to 3 days (Byrd and Castner,
2010). Barros et al. (2008) recorded Pheidole and
Camponotus ants (two widely distributed genera
in Ecuador and elsewhere) removing fly eggs,
larvae and pupae from pig carcasses. In Ecuador
(Union del Toachi, Pichincha), it has also been
observed, repeatedly, that army ants (Formicidae:
Dorylinae) raid on decomposing pigs, removing
both cadaveric pig remains and fly larvae (Emilia
Moreno, pers. comm.).
In some instances, ants are even capable of
establishing their colony on decomposing bodies.
As social insects, ant colonies as a whole pass
through developmental stages in which the number
and proportion of castes vary and generally, after
a colony has reached a certain size, it begins to
produce sexual castes (Peeters and Molet, 2010).
When the development time of a given ant species
is known it is be possible to estimate PMI based on
the time needed to establish a colony or produce
The role of ants (Hymenoptera: Formicidae) in forensic entomology
Giovanni Ramón1, David A Donoso2
certain castes. This was the case of some human
remains discovered inside a tool box; based on
the presence of an Anoplolepis gracilipes colony, the
PMI was estimated to be 14 to 18 months (Goff
and Win, 1997).
Ants as assassins: frequency of injuries
and fatalities caused by ants
The continuing expansion in the range of invasive
species, especially fire ants (Solenopsis spp.)
(Morrison et al., 2005; Rodriguez-Acosta and ReyesLugo, 2002; Vinson, 1997), which can inflict painful
stings (More et al., 2008; Prahlow and Barnard,
1998), aggravates the problem. At the highly antdiverse Neotropical region with many potentially
“dangerous” species of Ponerinae and Dorylinae
ant subfamilies whose stings can produce all
the range from mild reactions to severe allergic
reactions and even death, the increase in encounters
between people and stinging ants could represent a
serious public health concern.
As many other species in the order Hymenoptera
like wasps, bumblebees, velvet ants and hornets,
ants possess a sting formed by the modified
terminal abdominal segments. With it, ants are
capable of injecting venom typically produced in
the attached poison glands (Hölldobler and Wilson,
1990). While some ants use the stinging apparatus
to send chemical messages to nestmates or to
produce trail pheromones (Hölldobler and Wilson,
1990), most of them use it to hunt for prey or as a
defense mechanism. It is the latter in which we will
focus the discussion of this section.
While humans are not a suitable prey for ants
they still can become victims of ant stings when
they come cross their paths. Although in most
tropical and subtropical countries, ant stings are
only a minor concern (McGain and Winkel, 2002),
in some, encounters with ants are becoming more
common (Kemp et al., 2000; deShazo et al., 1984).
As noted by Rodriguez-Acosta and Reyes-Lugo
(2002), the reaction from ant stings can range from
moderate to extensive local allergic reactions,
systemic hypersensitivity, anaphylactic shock and,
in extreme cases, even death (Hardwick et al., 1992).
In the United States alone, 44 fatalities from fire
ant (Solenopsis invicta and Solenopsis richteri) stings
have been reported in the last 50 years (Bury et al.,
2012). Most fatalities however are attributed to
anaphylaxis rather than to the direct effect of the
venom (Bury et al., 2012).
Artifacts caused by ants on decomposing bodies:
biting, stinging, burying and modification of
blood patterns
Ant stings are painful. Some ants, especially
poneroids (from the Greek word ponera, for pain),
which have primitive characteristics, can inflict some
of the most painful insect stings. In the Neotropical
region, for example, the bullet ant Paraponera clavata
sting has been categorized as “traumatically painful”
(Starr, 1985). Similarly, in Australia Myrmecia spp.
ants are also respected for their painful stings. It is
thought that painful stings evolved as a mechanism
to deter vertebrate aggressors (Schmidt, 1990). This
stinging behavior can potentially inflict marks in
the decomposing body that can provide valuable
evidence for the investigation.
Another case of ant sting with forensic value is
the fire ants (S.invicta, S. richteri and S. geminata).
Introduced in the 20th century from South America
to the United States, fire ants have been expanding
their range at fast pace. Today they are reported
throughout most US southern states (Ascunce et al.,
2011) and also in Australia, New Zealand, Taiwan,
Hong Kong, Macao, and China (Wetterer, 2011).
It has been estimated that between 30 and 60% of
people living in infested areas are stung every year;
from these in the US it has been reported that 0.6 to
16% presented anaphylactic reactions (Caldwell et
al., 1999; Adams and Lofgren, 1981). Fire ants get
their names from the painful stings they are capable
of inflicting. Scientific and medical literature is full
of reports of fire ant stings and their consequences.
Biting can also exert significant damage to corpses.
Barros et al. (2008) recorded the presence of Pheidole
and Camponotus ants inflicting significant damage to
the nose and ears of pig carcasses (e.g. taphonomic
marks). Similarly, in a study on the decomposition
of rat corpses in Brazil, small injuries made by ants
were detected at every decomposition stage and
over different parts of the body (Moura et al., 1997).
These bites can be confounded with pre-mortem
caused injuries. A throughout review of the main
artifacts caused by ants on decomposing bodies can
be found in Campobasso et al. (2009). Without careful
consideration of these ant-inflicted marks, they can
misguide investigators about the cause of death.
Another way in which ants can influence the
decomposition process is by burying the body, or
parts of the body. Fire ants are also a good example
as they have been found capable of this. In Texas
for example, a wound on the abdominal area of
a partially buried human body was found filled
with soil by S. invicta, preventing other insects
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from reaching it. Only nine days later were other
insects capable of colonizing the corpse to continue
the usual decomposition process; as noted by the
authors, the estimation of the PMI based only in the
fly activity would have been dramatically skewed
(Lindgren et al., 2011). Similarly, in a study by
Moura et al. (1997) in Brazil, the head and anal area
of decomposing rats were found buried by ants,
preventing the access by flies.
Ants can make difficult the interpretation of events
on a crime scene by altering the bloodstain patterns
or by moving blood to new locations. Investigators
use the size and shape of the bloodstains to try to
understand what happened at the crime scene.
Some insects like flies, however, by walking over
a corpse or by feeding on blood can alter the blood
pattern. Moreover, some of the patterns they leave
can be confused with those left by a specific gun or
event. Although the possibility exists, we are not
aware of any specific report.
Determination of place of death
Insect species found inside a decomposing body
that do not correspond to the insect fauna normally
found at the area can indicate that a body has been
moved from a different place, and given the case that
the geographic distribution of the insect is known, it
can even signal the site from which the body was
relocated (Catts and Goff, 1992). In their study, Chen
et al. (2014) found that the distribution of ants was
irregular, so the ants found in monkey remains were
useful in determining the place of death.
In the Neotropics, composition of ant communities
of various sites has been well documented.
For instance, in Ecuador, the ant community
composition of a pre-montane forest (Ramón
et al., 2013; Donoso and Ramón, 2009), of an
Amazonian forest (Mertl et al., 2010; Ryder Wilkie
et al., 2010; 2009; 2007) and of mountain forests in
the southern provinces (Delsinne et al., 2013; 2012;
Jacquemin et al., 2012) have been well documented
in the last years. This knowledge could aid forensic
investigators to resolve criminal cases if, for
example, ants are found in a corpse that has been
moved to determine the site of origin. Forensic
investigations would benefit from more studies
of local ant communities in natural and urban
environments, and of their seasonal changes in
composition, in Ecuador in particular and in the
Neotropical region in general.
22
CONCLUSIONS
Our main conclusion after reviewing the literature
and from our field experience, is that ants cannot be
overlooked anymore as merely casual or secondary
visitors at crime scenes. We have seen that ants can
cause errors in the estimation of the PMI in three
ways: by acting as predators of other arthropods
that feed on carrion, by altering the composition of
the microbial community and by directly removing
chunks of the corpse. The latter one has been
overlooked in the Neotropical region and needs to
be researched more deeply in the future.
While traditionally flies and beetles have been
used to establish the PMI, we found that ants that
establish their colonies within decomposing bodies
can also be used for that purpose. By determining
the number of individuals on each caste, or by
observing the production of sexual individuals, a
fairly precise estimate of the PMI can be obtained
with a previous knowledge of the species life cycle.
In this regard, forensic entomology would benefit
from the study of the life cycle of the most common
local species at each country.
While the stings of most species are capable of
causing an acute pain at the most, some people
can present allergic reactions that can even cause
death. In view of the increasing expansion of cities
towards rural areas and with the increase in the
geographical range of invasive species like fire
ants, encounters between ants and humans are
expected to become more frequent. This could
start to become a health problem that needs to be
researched in order to reduce its impact.
When ants are found on a crime scene, or a body is
found in a place where ants are abundant, evidence
should be interpreted with care. By biting, stinging,
burying the body or by altering the blood stain
patterns; ants can lead forensic investigators not
only to make mistakes on the estimation of the PMI
but also can be misguided when looking for the
possible cause of death.
Finally, when ants are found within a body that
has been moved, their identifications may help
investigators to determine the original place where
the body was located after identifying the ants
found to species level. This, however, requires a
previous knowledge not only of the ecology but
also of the biogeography of ant communities.
The role of ants (Hymenoptera: Formicidae) in forensic entomology
Giovanni Ramón1, David A Donoso2
With the ongoing interest from forensic
investigators and related institutions in Central
and South America in the last years (e.g. Pujol-Luz
et al., 2008; Arnaldos 2006; Centeno, 2000), there are
now more opportunities to increase the knowledge
on the insects of forensic importance and the
role they play in the decomposition process. As
previously mentioned, ants are important actors of
the decomposition process and forensic science in
the Neoptropical region would greatly benefit from
more studies on their biology.
ACKNOWLEDGMENTS
This work was financed, in part, by UTPL Grant #
PROY_CCNN_0022 to DAD. The authors would
like to than Álvaro Barragán and Emilia Moreno
for inviting us to write this manuscript. The
authors thank Thibaut Delsinne and Aura Paucar
for the valuable comments to earlier drafts of the
manuscript. DAD thanks UTPL and Aventura
Lojana for the constant support. We further thank
the FGE of Ecuador for the support given to the
development of forensic sciences in Ecuador.
BIBLIOGRAPHICAL REFERENCES
Adams C and Lofgren C. 1981. Red imported fire
ants (Hymenoptera: Formicidae): frequency of
sting attacks on residents of Sumter County,
Georgia. Journal of Medical Entomology, 18: 378–
382.
Arnaldos MI, Prado C, Castro E, Presa JJ, Luna A,
López Gallego E and García García MD. 2006.
Importancia de los estudios regionales de la
fauna sarcosaprófaga. Aplicación de la práctica
forense. Ciencia forense, 8: 63-81.
Ascunce MS, Yang CC, Oakey J, Calcaterra L.,
Wu W-J, Shih C-J, Goudet JRM, Ross K G and
Shoemaker D. 2011. Global invasion history of
the fire ant Solenopsis invicta. Science, 331: 1066–
1068.
Barros AS, Dutra F and Ferreira R. 2008. Insects of
forensic importance from Rio Grande do Sul
state in southern Brazil. Revista Brasileira de
Entomologia, 52(4): 641–646.
Bermúdez S and Pachar JV. 2010. Artrópodos
asociados a cadáveres humanos en Ciudad
de Panamá, Panamá. Revista Colombiana de
Entomología, 36(1): 86-89.
Bolton B. 2014. An online catalog of the ants of
the world. Available from http://antcat.org.
(Consultada 18-Marzo-2015).
Brown S, Haas M, Black J, Parameswaran A, Woods G
and Heddle R. 2004. In vitro testing to diagnose
venom allergy and monitor immunotherapy: a
placebo-controlled, crossover trial. Clinical and
Experimental Allergy, 34: 792–800.
Bury D, Langlois N and Byard RW. 2012. AnimalRelated Fatalities- Part II: Characteristic
Autopsy Findings and Variable Causes of Death
Associated with Envenomation, Poisoning,
Anaphylaxis, Asphyxiation, and Sepsis. Journal
of Forensic Sciences, 57: 375–380.
Byrd JH y Castner JL. 2010. Forensic Entomology, The
Utility of Arthropods in Legal Investigations. 2da
Edición. CRC Press, USA.
Caldwell ST, Schuman SH and Simpson W. 1999.
Fire ants: a continuing community health
threat in South Carolina. Journal of the South
Carolina Medical Association, 1975 (95): 231–235.
Campobasso CP, Marchetti D, Introna F and
Colonna MF. 2009. Postmortem artifacts
made by ants and the effect of ant activity on
decompositional rates. The American Journal of
Forensic Medicine and Pathology, 30(1): 84–87.
Catts E and Goff ML. 1992. Forensic entomology
in criminal investigations. Annual Review of
Entomology, 37(1): 253–272.
Centeno N. 2000. La entomología forense:
aplicaciones, fundamentos y algunos datos
sobre la Argentina. Boletín de la Sociedad
Entomológica Argentina, 16: 9-11.
Chapin FS, Matson PA and Vitousek P. 2011.
Principles of Terrestrial Ecosystem Ecology.
Springer, USA.
Chen CD, Nazni WA, Lee HL, Hashim R, Abdullah
NA, Ramli R, Lau KW, Heo CC, Goh TG, Izzul
AA and Sofian-Azirun M. 2014. A preliminary
report on ants (Hymenoptera: Formicidae)
recovered from forensic entomological studies
conducted in different ecological habitats in
Malaysia. Tropical Biomedicine, 31(2): 381–386.
DeFoliart GR. 1999. Insects as food: why the
western attitude is important. Annual Review of
Entomology, 44: 21–50.
23
REMCB 36 pp. 19-26. 2015
Delsinne T, Arias-Penna T and Leponce M. 2013.
Effect of rainfall exclusion on ant assemblages
in montane rainforests of Ecuador. Basic and
Applied Ecology, 14(4): 357-365.
Delsinne T, Sonet G, Nagy ZNT, Wauters N,
Jacquemin J and Leponce M. 2012. High
species turnover of the ant genus Solenopsis
(Hymenoptera:
Formicidae)
along
an
altitudinal gradient in the Ecuadorian Andes,
indicated by a combined DNA sequencing
and morphological approach. Invertebrate
Systematics, 26(6): 457-469.
Del Toro I, Ribbons RR and Pelini SL. 2012. The
little things that run the world revisited: a
review of ant-mediated ecosystem services
and disservices (Hymenoptera: Formicidae).
Myrmecological News, 17: 133–146.
deShazo RD, Griffing C, Kwan TH, Banks W and
Dvorak H. 1984. Dermal hypersensitivity
reactions to imported fire ants. Journal of Allergy
and Clinical Immunology, 74: 841–847.
Donoso DA and Ramón G. 2009. Composition
of a high diversity leaf litter ant community
(Hymenoptera:
Formicidae)
from
an
Ecuadorian pre-montane rainforest. Annales de
la Société Entomologique de France 45(4): 487-499.
Early M and Goff ML. 1986. Arthropod succession
patterns in exposed carrion on the island of
O’ahu, Hawaiian Islands, USA. Journal of
Medical Entomology, 23:520–531.
Fittkau EJ and Klinge H. 1973. On biomass and
trophic structure of the central Amazonian rain
forest ecosystem. Biotropica, 5: 2–14.
Goff M and Win B. 1997. Estimation of postmortem
interval based on colony development
time for Anoplolepis longipes (Hymenoptera:
Formicidae). Journal of Forensic Sciences, 42(6):
1176–1179.
Grimaldi D and Engel MS. 2005. Evolution of the
Insects. Cambridge University Press. UK.
Hall RD and Huntington TE. 2010. Introduction:
Perceptions and Status of Forensic Entomology.
In: Byrd JH and Castner JL (eds.) Forensic
Entomology, The Utility of Arthropods in Legal
Investigations. 2da Edición. CRC Press, USA.
24
Hardwick WE, Royall JA, Petitt BA and Tilden SJ.
1992. Near fatal fire ant envenomation of a
newborn. Pediatrics, 90: 622–624.
Hölldobler B and Wilson E. 1990. The Ants. Belknap
Press. USA.
Jacquemin J, Drouet T, Delsinne T, Roisin Y and
Leponce M. 2012. Soil properties only weakly
affect subterranean ant distribution at small
spatial scales. Applied Soil Ecology, 62: 163-169.
Kaspari M. 1996a. Litter ant patchiness at the
1-m 2 scale: disturbance dynamics in three
Neotropical forests. Oecologia, 107: 265–273.
Kapari M. 1996b. Worker size and seed size
selection by harvester ants in a Neotropical
forest. Oecologia, 105: 397–404.
Kemp SF, deShazo RD, Moffitt JE, Williams DF and
Buhner WA. 2000. Expanding habitat of the
imported fire ant (Solenopsis invicta): a public
health concern. Journal of Allergy and Clinical
Immunology, 105: 683–691.
Lengyel S, Gove AD, Latimer AM, Majer JD and
Dunn RR. 2010. Convergent evolution of
seed dispersal by ants, and phylogeny and
biogeography in flowering plants: a global
survey. Perspectives in Plant Ecology, Evolution
and Systematics, 12: 43–55.
Lindgren NK, Bucheli SR, Archambeault AD and
Bytheway JA. 2011. Exclusion of forensically
important flies due to burying behavior by
the red imported fire ant (Solenopsis invicta) in
southeast Texas. Forensic Science International,
204(1): e1–e3.
McGain F and Winkel KD. 2002. Ant sting mortality
in Australia. Toxicon, 40: 1095–1100.
Mertl A, Sorenson M and Traniello J. 2010.
Community-level interactions and functional
ecology of major workers in the hyperdiverse
ground-foraging
Pheidole
(Hymenoptera,
Formicidae) of Amazonian Ecuador. Insectes
Sociaux, 57: 441-452.
More DR, Kohlmeier RE and Hoffman DR. 2008.
Fatal anaphylaxis to indoor native fire ant
stings in an infant. The American Journal of
Forensic Medicine and Pathology, 29: 62–63.
The role of ants (Hymenoptera: Formicidae) in forensic entomology
Giovanni Ramón1, David A Donoso2
Moreau CS and Bell CD. 2013. Testing the museum
versus cradle tropical biological diversity
hypothesis: phylogeny, diversification, and
ancestral biogeographic range evolution of the
ants. Evolution, 67: 2240–2257.
Moreira D, Morais VND, Vieira-da-Motta O,
Campos-Farinha AEDC and Tonhasca A. 2005.
Ants as carriers of antibiotic-resistant bacteria
in hospitals. Neotropical Entomology, 34: 999–
1006.
Morrison LW, Porter SD, Daniels E and Korzukhin
MD. 2005. Potential global range expansion of
the invasive fire ant, Solenopsis invicta. Biological
Invasions, 6: 183–191.
Moura MO, Carvalho CJD and Monteiro-Filho
EL. 1997. A preliminary analysis of insects of
medico-legal importance in Curitiba, State of
Paraná. Memórias do Instituto Oswaldo Cruz, 92:
269–274.
Pape T, Wolff M and Amat EC. 2004. Los califóridos,
óstridos, rinofóridos y sarcofágidos (Diptera:
Calliphoridae, Oestridae, Rhinophoridae,
Sarcophagidae) de Colombia. Biota Colombiana,
5: 201–208.
Peeters C and Molet M. 2010. Colonial Reproduction
and Life Histories. In: Lach L, Parr CL and
Abbott KL (eds.) Ant Ecology. Oxford University
Press, USA.
Philpott SM and Armbrecht I. 2006. Biodiversity in
tropical agroforests and the ecological role of
ants and ant diversity in predatory function.
Ecological Entomology, 31: 369–377.
Ramón G, Barragán A and Donoso DA. 2013.
Can clay banks increase the local ant species
richness of a montane forest? Métodos en
Ecologia y Sistematica, 8(2): 37–53.
Reddy N, Xu H and Yang Y. 2011. Unique naturalprotein
hollow-nanofiber
membranes
produced by weaver ants for medical
applications. Biotechnology and Bioengineering,
108: 1726–1733.
Rodovalho CM, Santos AL, Marcolino MT, Bonetti
AM and Brandeburgo MA. 2007. Urban ants
and transportation of nosocomial bacteria.
Neotropical Entomology, 36: 454–458.
Rodriguez-Acosta A and Reyes-Lugo M.
2002. Severe human urticaria produced
by ant (Odontomachus bauri, Emery 1892)
(Hymenoptera:
Formicidae)
venom.
International Journal of Dermatology, 41: 801–803.
Ryder Wilkie K, Mertl A and Traniello J. 2010.
Species diversity and distribution patterns of
the ants of Amazonian Ecuador. PLoS ONE,
5(10): 1–12.
Ryder Wilkie K, Mertl A and Traniello J.
2009. Diversity of ground-dwelling ants
(Hymenoptera: Formicidae) in primary and
secondary forests in Amazonian Ecuador.
Myrmecological News, 12: 139–147.
Ryder Wilkie K, Mertl A and Traniello J. 2007.
Biodiversity below ground: probing the
subterranean ant fauna of Amazonia.
Naturwissenschaften, 94(9): 725–731.
Prado e Castro C, García M-D, Palma C and
Martínez-Ibáñez M-D. 2014. First report on
sarcosaprophagous Formicidae from Portugal
(Insecta: Hymenoptera). Annales de la Société
entomologique de France (N.S.), 50: 51–58.
Schmidt J. 1990. Hymenopteran venoms:
striving toward the ultimate defense against
vertebrates. In: Evans DL and Schmidt JO (eds.)
Insect defenses: adaptive mechanisms and strategies
of prey and predators. State University of New
York Press, USA.
Prahlow JA and Barnard JJ. 1998. Fatal anaphylaxis
due to fire ant stings. The American journal of
forensic medicine and pathology, 19: 137–142.
Starr CK. 1985. A simple pain scale for field
comparison of hymenopteran stings. Journal of
Entomological Science, 20: 225–231.
Pujol-Luz JR, Arantes LC y Constantino R. 2008.
Cem anos da Entomologia Forense no Brasil
(1908-2008). Revista Brasileira de Entomologia,
52(4): 485–492.
Valdes-Perezgasga
M,
Sanchez-Ramos
FJ,
Garcia-Martinez O and Anderson GS. 2010.
Arthropods of forensic importance on pig
carrion in the Coahuilan semidesert, Mexico.
Journal of Forensic Sciences 55(4): 1098–1101.
25
REMCB 36 pp. 19-26. 2015
Van Mele P. 2008. A historical review of research on
the weaver ant Oecophylla in biological control.
Agricultural and Forest Entomology, 10: 13–22.
Wetterer JK. 2011. Worldwide spread of the tropical
fire ant, Solenopsis geminata (Hymenoptera:
Formicidae). Myrmecological News, 14: 21–35.
Vasconcelos SOD and Araujo M. 2012.
Necrophagous species of Diptera and
Coleoptera in northeastern Brazil: state of the art
and challenges for the Forensic Entomologist.
Revista Brasileira de Entomologia, 56: 7–14.
Wilson EO. 2005. Oribatid mite predation by small
ants of the genus Pheidole. Insectes Sociaux, 52:
263–265.
Vinson SB. 1997. Invasion of the red imported
fire ant (Hymenoptera: Formicidae): spread,
biology, and impact. American Entomologist, 43:
23–39.
Wolff M, Giraldo N and Perez MV. 2004 Estudio
de la entomofauna cadavérica encontrada en
humanos alrededor de Medellín, Colombia y
su aplicación en la determinación del intervalo
postmortem. Socolen, Memorias XXXI Congreso.
Bogotá, Colombia: 125–130.
Wardle DA, Hyodo F, Bardgett RD, Yeates GW and
Nilsson M-C. 2011. Long-term aboveground and
belowground consequences of red wood ant
exclusion in boreal forest. Ecology, 92: 645–656.
FIGURE 1. A) Cephalotes atratus workers tearing
apart the skin on the leg of a dead pig (bloating
stage) at Yasuní National Park, Ecuador, B)
detail of a C. atratus worker biting out the flesh
of the dead pig and C) some Crematogaster sp.
workers tearing apart the skin of the dead pig.
Photos courtesy of Ricardo Jaramillo.
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