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Beetles (Insecta, Coleoptera) associated with pig carcasses exposed in a Caatinga area, Northeastern Brazil

Coleópteros (Insecta, Coleoptera) associados a carcaças de suínos em Caatinga, Nordeste do Brasil

Abstracts

The species richness, abundance and seasonality of Coleoptera fauna associated with pig carcasses exposed in a Caatinga area were examined. Tray, pitfall and modified Shannon traps were settled together to collect these insects during two seasons (dry and rainy). 4,851 beetles were collected, belonging to 19 families and 88 species. Staphylinidae (2,184) and Histeridae (1,264) were the most abundant families and accounted for 71.1% of the specimens collected. Scarabaeidae (15) showed the highest species richness. The most abundant species were Atheta iheringi Bernhauer, 1908 (Staphylinidae) (1,685), Euspilotus sp. (Histeridae) (461), Stelidota geminata (Say, 1825) (Nitidulidae) (394), Xerosaprinus diptychus(Marseul, 1855) (Histeridae) (331) and Dermestes maculatus De Geer, 1774 (Dermestidae). Amongst these species, X. diptychus showed to be strongly influenced by seasonality, since 96.1% of the specimens were collected during the dry season.

animal decomposition; biodiversity; forensic entomology; necrophagous insects; seasonality


A riqueza, abundância e sazonalidade da coleopterofauna associada a carcaças de suínos expostas em uma área de Caatinga foram examinadas. Para captura desses insetos foram utilizadas armadilhas do tipo bandeja, pitfall e Shannon modificada, durante duas estações (seca e chuvosa). 4.851 coleópteros foram coletados, pertencentes a 19 famílias e 88 espécies. Staphylinidae (2.184) e Histeridae (1.264) foram as famílias mais abundantes e somaram 71,1% dos espécimes coletados. Scarabaeidae (15) apresentou a maior riqueza. As espécies mais abundantes foram Atheta iheringi Bernhauer, 1908 (Staphylinidae) (1,685), Euspilotus sp. (Histeridae) (461), Stelidota geminata (Say, 1825) (Nitidulidae) (394), Xerosaprinus diptychus(Marseul, 1855) (Histeridae) (331) e Dermestes maculatus De Geer, 1774 (Dermestidae). Entre essas espécies, X. diptychus mostrou-se fortemente influenciada pela sazonalidade, uma vez que 96,1% dos espécimes foram coletados durante a estação seca.

decomposição animal; biodiversidade; entomologia forense; insetos necrófagos; sazonalidade


1.

Introduction

Insects are usually the first animals to locate a cadaver, attracted by the odours and gases released by it, and then start to use it as a resource for feeding, reproduction and larval development (Cornaby, 1974CORNABY, BW., 1974. Carrion reduction by animals in contrasting tropical habitats. Biotropica, vol. 6, no. 1, p. 51-63. http://dx.doi.org/10.2307/2989697.
http://dx.doi.org/10.2307/2989697...
). This entomofauna attracts also another kind of fauna, consisting of predatory insects (Smith, 1986). The beetles belong to both ecological categories. Few species are effectively necrophagous, while most of them are predatory, but both categories can provide useful information from the forensic point of view, especially concerning to the estimation of the postmortem interval (PMI) (Catts and Goff, 1992CATTS, EP. and GOFF, ML., 1992. Forensic entomology in criminal investigations. Annual Review of Entomology, vol. 37, no. 1, p. 253-272. http://dx.doi.org/10.1146/annurev.en.37.010192.001345. PMid:1539937
http://dx.doi.org/10.1146/annurev.en.37....
; Santos et al. 2013SANTOS, WE., ALVES, ACF., FARIAS, RCAP. and CREÃO-DUARTE, AJ., 2013. Ecological roles of Coleoptera associated with carcasses in Caatinga. EntomoBrasilis, vol. 6, no. 3, p. 248-250. http://dx.doi.org/10.12741/ebrasilis.v6i3.325.
http://dx.doi.org/10.12741/ebrasilis.v6i...
).

In Brazil, important contributions to diversity, ecology and taxonomy of the fauna associated with carcasses have been made in the last two decades, but the majority of the studies have focussed on Diptera and are concentrated in the Southeast and South regions (Pujol-Luz et al., 2008PUJOL-LUZ, JR., ARANTES, LC. and CONSTANTINO, R., 2008. Cem anos da entomologia forense no Brasil (1908-2008). Revista Brasileira de Entomologia, vol. 52, no. 4, p. 485-492. http://dx.doi.org/10.1590/S0085-56262008000400001.
http://dx.doi.org/10.1590/S0085-56262008...
). However, due to their succession pattern, Coleoptera are the most useful insects for the estimation of PMI when skeletonised corpses are found (Kulshrestha and Satpathy, 2001KULSHRESTHA, P. and SATPATHY, DK., 2001. Use of beetles in forensic entomology. Forensic Science International, vol. 120, no. 1-2, p. 15-17. http://dx.doi.org/10.1016/S0379-0738(01)00410-8.PMid:11457603
http://dx.doi.org/10.1016/S0379-0738(01)...
). In some cases, it is possible to estimate the PMI based on rates of development of some species (Arnaldos et al., 2004ARNALDOS, MI., SÁNCHEZ, F., ÁLVARES, P. and GARCÍA, MD., 2004. A forensic entomology case from the Southeastern Iberian Peninsula. Anil Aggrawal's Internet Journal of Forensic Medicine and Toxicology, vol. 5, p. 22-25.). Coleoptera can also be used to support the PMI estimate from Diptera data (Goff and Flynn, 1991GOFF, ML. and FLYNN, MM., 1991. Determination of postmortem interval by arthropod succession: a case study from the Hawaiian Islands. Journal of Forensic Sciences, vol. 36, no. 2, p. 607-614. PMid:2066736.).

Regional and seasonal studies are essential to identify specific cadaveric fauna, since each biome and season has its specific fauna because of particular environmental conditions (Mise et al., 2010MISE, KM., SOUZA, ASB., CAMPOS, CM., KEPPLER, RLF. and ALMEIDA, LM., 2010. Coleoptera associated with pig carcass exposed in a forest reserve, Manaus, Amazonas, Brazil. Biota Neotropica, vol. 10, no. 1, p. 321-324. http://dx.doi.org/10.1590/S1676-06032010000100027.
http://dx.doi.org/10.1590/S1676-06032010...
; Rosa et al., 2011ROSA, TA., BABATA, MLY., SOUZA, CM., SOUSA, D., MELLO-PATIU, CA., VAZ-DE-MELLO, FZ. and MENDES, J., 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Revista Brasileira de Entomologia, vol. 55, no. 3, p. 424-434. http://dx.doi.org/10.1590/S0085-56262011005000045.
http://dx.doi.org/10.1590/S0085-56262011...
). The main objective was to analyse the diversity of the Coleoptera fauna associated with pig (Sus scrofa Linnaeus, 1758) carcasses, during the dry and wet seasons in Caatinga, a dry forest endemic to Brazil, whose insect fauna is poorly known (Vasconcellos et al., 2010VASCONCELLOS, A., ANDREAZZE, R., ALMEIDA, AM., ARAUJO, HFP., OLIVEIRA, ES. and OLIVEIRA, U., 2010. Seasonality of insects in a semi-arid Caatinga of Northeastern Brazil. Revista Brasileira de Entomologia, vol. 54, no. 3, p. 471-476. http://dx.doi.org/10.1590/S0085-56262010000300019.
http://dx.doi.org/10.1590/S0085-56262010...
; Creão-Duarte et al. 2013; Alves et al., 2014ALVES, ACF., SANTOS, WE., FARIAS, RCAP. and CREÃO-DUARTE, AJ., 2014. Blowflies (Diptera, Calliphoridae) associated with pig carcasses in a Caatinga area, Northeastern Brazil. Neotropical Entomology, vol. 43, no. 2, p. 122-126. http://dx.doi.org/10.1007/s13744-013-0195-4.
http://dx.doi.org/10.1007/s13744-013-019...
). Brief comments on the main families of forensic importance are also included, highlighting the richness, abundance, behaviour and ecological aspects of some species.

2.

Material and Methods

The study was carried out during October 2010 (dry season) and February 2011 (rainy season) at the Private Reserve for the Environmental Inheritance Fazenda Almas (RPPN Fazenda Almas), in São José dos Codeiros, state of Paraíba, Brazil (7° 28′ 19″ S, 36° 53′ 40″ W). The study area is located at an altitude of 650 m a.s.l. and 300 km away from the coast. The vegetation varies from open to dense arboreal Caatinga, with a strong deciduous characteristic during the dry season. The soil is sandy and arid and the topography is irregular, with inselbergs and rocky outcrops (Vasconcellos et al., 2010VASCONCELLOS, A., ANDREAZZE, R., ALMEIDA, AM., ARAUJO, HFP., OLIVEIRA, ES. and OLIVEIRA, U., 2010. Seasonality of insects in a semi-arid Caatinga of Northeastern Brazil. Revista Brasileira de Entomologia, vol. 54, no. 3, p. 471-476. http://dx.doi.org/10.1590/S0085-56262010000300019.
http://dx.doi.org/10.1590/S0085-56262010...
).

Two male pig carcasses, weighing around 15 kg each, were used in both seasons. The animals were killed with a single gunshot to the head. Each carcass was exposed in an iron cage to prevent interference of vertebrate scavengers. Underneath the cage there was a trap tray buried at ground level containing sawdust in order to collect actively the Coleoptera that take shelter under the carcasses. Four pitfall traps were settled around the cage, 1 m away from it, to passive capture. Each cage was also covered by a modified Shannon trap with a collector tube containing 70% alcohol at the apex (Alves et al., 2014ALVES, ACF., SANTOS, WE., FARIAS, RCAP. and CREÃO-DUARTE, AJ., 2014. Blowflies (Diptera, Calliphoridae) associated with pig carcasses in a Caatinga area, Northeastern Brazil. Neotropical Entomology, vol. 43, no. 2, p. 122-126. http://dx.doi.org/10.1007/s13744-013-0195-4.
http://dx.doi.org/10.1007/s13744-013-019...
). Coleoptera were collected daily in and from the traps.

The identification of the Coleoptera was done by experts and specific literature (Mazur, 2001MAZUR, S., 2001. Review of the Histeridae (Coleoptera) of México. Dugesiana, vol. 8, no. 2, p. 17-66.; Navarrete-Heredia et al., 2002NAVARRETE-HEREDIA, JL., NEWTON, AF., THAYER, MK., ASHE, JS. and HANDLER, DS., 2002. Guía ilustrada para los géneros de Staphylinidae (Coleoptera) de México. México: Universidad de Guadalajara/CONABIO. 401 p.; Almeida and Mise, 2009ALMEIDA, LM. and MISE, KM., 2009. Diagnosis and key of the main families and species of South American Coleoptera of forensic importance. Revista Brasileira de Entomologia, vol. 53, no. 2, p. 227-244. http://dx.doi.org/10.1590/S0085-56262009000200006.
http://dx.doi.org/10.1590/S0085-56262009...
; Vaz-de-Mello et al., 2011VAZ-DE-MELLO, FZ., EDMONDS, WD., OCAMPO, FC. and SCHOOLMEESTERS, P., 2011. A multilingual key to the genera and subgenera of the subfamily Scarabaeinae of the New World (Coleoptera: Scarabaeidae). Zootaxa, vol. 2854, p. 1-73.; Aballay et al., 2013ABALLAY, FH., ARRIAGADA, G., FLORES, GE. and CENTENO, ND., 2013. An illustrated key to and diagnoses of the species of Histeridae (Coleoptera) associated with decaying carcasses in Argentina. ZooKeys, vol. 261, no. 261, p. 61-84. http://dx.doi.org/10.3897/zookeys.261.4226. PMid:23653510
http://dx.doi.org/10.3897/zookeys.261.42...
). All material used was incorporated into the Coleção Entomológica do Departamento de Sistemática e Ecologia at Universidade Federal da Paraíba (DSEC/UFPB).

3.

Results and Discussion

A total of 4,851 adults belonging to 19 families and 88 species was collected (Table 1). Staphylinidae was the most abundant family with 2,184 individuals, followed by Histeridae (1,264), Nitidulidae (395), Dermestidae (333), Scarabaeidae (300), Cleridae (120), Carabidae (75) and Trogidae (75), amounting to 97.8% of the specimens.

Table 1.
Absolute (n) and relative (%) abundance, and species richness (S) of Coleoptera families associated to pig carcasses during dry and rainy seasons in a Caatinga area, state of Paraíba, Brazil.

Under similar conditions different results were obtained by Carvalho et al. (2000)CARVALHO, LM., THYSSEN, PJ., LINHARES, AX. and PALHARES, FAB., 2000. A checklist of arthropods associated with pig carrion and human corpses in southeastern Brazil. Memorias do Instituto Oswaldo Cruz, vol. 95, no. 1, p. 135-138. http://dx.doi.org/10.1590/S0074-02762000000100023. PMid:10656720
http://dx.doi.org/10.1590/S0074-02762000...
, in Campinas, state of São Paulo, whose decreasing order of abundance was Scarabaeidae, Histeridae, Silphidae and Staphylinidae. Mise et al. (2007)MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
, in Curitiba, State of Paraná, found Staphylinidae as the most abundant family, as in the present study, followed by Silphidae, Histeridae and Cleridae. Rosa et al. (2011)ROSA, TA., BABATA, MLY., SOUZA, CM., SOUSA, D., MELLO-PATIU, CA., VAZ-DE-MELLO, FZ. and MENDES, J., 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Revista Brasileira de Entomologia, vol. 55, no. 3, p. 424-434. http://dx.doi.org/10.1590/S0085-56262011005000045.
http://dx.doi.org/10.1590/S0085-56262011...
, in areas of Cerrado, in Uberlândia, State of Minas Gerais, collected mainly Dermestidae, Histeridae, Cleridae and Staphylinidae. These divergent results highlight the importance of local studies for forensic practice in Brazil.

High species richness of beetles associated to carcasses has been registered in the Neotropical region: 62 and 113 species were mentioned respectively by Luederwaldt (1911)LUEDERWALDT, G., 1911. Os insectos necrophagos paulistas. Revista do Museu Paulista, vol. 8, p. 414-433. and Pessôa and Lane (1941)PESSÔA, SB. and LANE, F., 1941. Coleópteros de interesse médico-legal. Arquivos de Zoologia do Estado de São Paulo, vol. 2, p. 389-504. in São Paulo and neighbourhood; 112 species were collected by Mise et al. (2007)MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
in Curitiba, state of Paraná; and 41 species were found by Mise et al. (2010)MISE, KM., SOUZA, ASB., CAMPOS, CM., KEPPLER, RLF. and ALMEIDA, LM., 2010. Coleoptera associated with pig carcass exposed in a forest reserve, Manaus, Amazonas, Brazil. Biota Neotropica, vol. 10, no. 1, p. 321-324. http://dx.doi.org/10.1590/S1676-06032010000100027.
http://dx.doi.org/10.1590/S1676-06032010...
in Manaus, State of Amazonas. These data demonstrate the importance of Coleoptera when sampling strategies do not neglect this taxonomic group.

Smith (1986) considered as Coleoptera families with forensic importance: Carabidae, Hydrophilidae, Silphidae, Leiodidae, Staphylinidae, Histeridae, Cleridae, Anthicidae, Dermestidae, Nitidulidae, Rhizophagidae, Ptinidae, Tenebrionidae, Scarabaeidae, Geotrupidae and Trogidae. Ten of these families were found in the present study and represented 98.7% of the specimens collected (Table 2).

Table 2.
Absolute (n) and relative (%) abundance of Coleoptera species of forensic importance associated to pig carcasses during dry and rainy seasons in a Caatinga area, State of Paraíba, Brazil.

Amongst the collected species, Pterostichus sp., Eremosaprinus sp., Ateuchus carbonarius(Harold, 1868), Canthidium manni Arrow, 1913, Coprophanaeus pertyi (Olsoufieff, 1924), Deltochilum verruciferum Felsche, 1911, Dichotomius geminatus (Arrow, 1913), Acylophorus sp., Philonthus figulus Erichson, 1840 and Tribolium sp. are reported for the first time associated to carcasses in the Neotropical region.

The most abundant beetles collected, with number of individuals greater than 1% of the total, were represented by 17 species: Atheta iheringi Bernhauer, 1908 (1,685), Euspilotussp. (461), S. geminata (394), Xerosaprinus diptychus (Marseul, 1855) (331), Dermestes maculatus De Geer, 1774 (330), Philonthus sp. 2 (294), Euspilotus azureus (Sahlberg, 1823) (188), A. carbonarius (129), Necrobia rufipes (De Geer, 1775) (120), Phelister sp. (114), Aleochara bonariensis Lynch, 1884 (102), Hister punctifer Paykull, 1811 (82), Omorgus suberosus Fabricius, 1775 (75), Omalodes foveola Erichson, 1834 (67), Loxandrus sp. (65), Belonuchus sp. 2 (48) and C. manni (47) (Table 2). These species totaled together 93.1% of all beetles collected. Future specific studies, mainly about succession pattern, life cicle and diet, are required for a better understanding of the relationship of these species with carcasses and thus establish its forensic importance.

Rove beetles (Staphylinidae) are common on carcasses because they are predators of larvae of other insects, with a preference for fly larvae (Navarrete-Heredia et al., 2002NAVARRETE-HEREDIA, JL., NEWTON, AF., THAYER, MK., ASHE, JS. and HANDLER, DS., 2002. Guía ilustrada para los géneros de Staphylinidae (Coleoptera) de México. México: Universidad de Guadalajara/CONABIO. 401 p.; Byrd and Castner, 2010BYRD, JH. and CASTNER, JL., 2010. Insects of forensic importance. In BYRD, JH. and CASTNER, JL. (Eds.). Forensic entomology: the utility of arthropods in legal investigations. Boca Raton: CRC Press. p. 39-126.). In addition, larvae of some species of Aleochara are ectoparasites of fly pupae (Smith 1986). Staphylinidae was the most abundant family in the study, with 2,184 specimens, mainly represented by A. iheringi(1,685) and species of Philonthus (326). Most of the Staphylinidade were collected in the rainy season (86.6%), which also showed higher species richness (13) (Table 2).

Mise et al. (2007)MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
highlighted Staphylinidae as the most abundant family, as in this study, with 29 species and 2,450 individuals captured. In many studies conducted in the Neotropical region, due to lack of specialists, the identification of these beetles is done only until genus. However, there seems to be a similarity of the composition of the Staphylinidae fauna among different regions in Brazil, mainly represented by Aleochara and Philonthus (Souza and Linhares, 1997SOUZA, AM. and LINHARES, AX., 1997. Diptera and Coleoptera of potential forensic importance in southeastern Brazil: relative abundance and seasonality. Medical and Veterinary Entomology, vol. 11, no. 1, p. 8-12. http://dx.doi.org/10.1111/j.1365-2915.1997.tb00284.x. PMid:9061672
http://dx.doi.org/10.1111/j.1365-2915.19...
; Mise et al., 2007MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
; Mise et al., 2010MISE, KM., SOUZA, ASB., CAMPOS, CM., KEPPLER, RLF. and ALMEIDA, LM., 2010. Coleoptera associated with pig carcass exposed in a forest reserve, Manaus, Amazonas, Brazil. Biota Neotropica, vol. 10, no. 1, p. 321-324. http://dx.doi.org/10.1590/S1676-06032010000100027.
http://dx.doi.org/10.1590/S1676-06032010...
; Rosa et al., 2011ROSA, TA., BABATA, MLY., SOUZA, CM., SOUSA, D., MELLO-PATIU, CA., VAZ-DE-MELLO, FZ. and MENDES, J., 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Revista Brasileira de Entomologia, vol. 55, no. 3, p. 424-434. http://dx.doi.org/10.1590/S0085-56262011005000045.
http://dx.doi.org/10.1590/S0085-56262011...
; Silva and Santos, 2012SILVA, RC. and SANTOS, WE., 2012. Fauna de Coleoptera associada a carcaças de coelhos expostas em uma área urbana no Sul do Brasil. EntomoBrasilis, vol. 5, no. 3, p. 185-189. http://dx.doi.org/10.12741/ebrasilis.v5.i3.245.
http://dx.doi.org/10.12741/ebrasilis.v5....
). In other countries, several different genera were recorded occurring on carcasses (Wolff et al., 2001WOLFF, M., URIBE, A., ORTIZ, A. and DUQUE, P., 2001. A preliminary study of forensic entomology in Medellín, Colombia. Forensic Science International, vol. 120, no. 1-2, p. 53-59. http://dx.doi.org/10.1016/S0379-0738(01)00422-4. PMid:11457610
http://dx.doi.org/10.1016/S0379-0738(01)...
; Battán Horenstein and Linhares, 2011BATTÁN HORENSTEIN, M. and LINHARES, AX., 2011. Seasonal composition and temporal succession of necrophagous and predator beetles on pig carrion in central Argentina. Medical and Veterinary Entomology, vol. 25, no. 4, p. 395-401. http://dx.doi.org/10.1111/j.1365-2915.2011.00969.x. PMid:21762401
http://dx.doi.org/10.1111/j.1365-2915.20...
; Aballay et al., 2012ABALLAY, FH., MURUA, AF., ACOSTA, JC. and CENTENO, ND., 2012. Succession of carrion fauna in the arid region of San Juan Province, Argentina and its forensic relevance. Neotropical Entomology, vol. 41, no. 1, p. 27-31. http://dx.doi.org/10.1007/s13744-011-0005-9. PMid:23950006
http://dx.doi.org/10.1007/s13744-011-000...
).

The great abundance of the family Histeridae is mainly due to the fact that their species are predators of larvae of flies, food resource widely available on carcasses (Smith, 1986; Byrd and Castner, 2010BYRD, JH. and CASTNER, JL., 2010. Insects of forensic importance. In BYRD, JH. and CASTNER, JL. (Eds.). Forensic entomology: the utility of arthropods in legal investigations. Boca Raton: CRC Press. p. 39-126.). Histeride was the second most abundant family in this study, represented mainly by species of Euspilotus (649), collected mostly during the rainy season (92.4%), and X. diptychus (331), in the dry season (96.1%) (Table 2).

Aballay et al. (2012)ABALLAY, FH., MURUA, AF., ACOSTA, JC. and CENTENO, ND., 2012. Succession of carrion fauna in the arid region of San Juan Province, Argentina and its forensic relevance. Neotropical Entomology, vol. 41, no. 1, p. 27-31. http://dx.doi.org/10.1007/s13744-011-0005-9. PMid:23950006
http://dx.doi.org/10.1007/s13744-011-000...
found X. diptychus and five species of Euspilotuson pig carcasses in a different kind of arid environment, in San Juan, Argentina. The other genera and species of Histeride collected are commonly found on carcasses throughout the Neotropical region (Souza and Linhares, 1997SOUZA, AM. and LINHARES, AX., 1997. Diptera and Coleoptera of potential forensic importance in southeastern Brazil: relative abundance and seasonality. Medical and Veterinary Entomology, vol. 11, no. 1, p. 8-12. http://dx.doi.org/10.1111/j.1365-2915.1997.tb00284.x. PMid:9061672
http://dx.doi.org/10.1111/j.1365-2915.19...
; Carvalho et al., 2000CARVALHO, LM., THYSSEN, PJ., LINHARES, AX. and PALHARES, FAB., 2000. A checklist of arthropods associated with pig carrion and human corpses in southeastern Brazil. Memorias do Instituto Oswaldo Cruz, vol. 95, no. 1, p. 135-138. http://dx.doi.org/10.1590/S0074-02762000000100023. PMid:10656720
http://dx.doi.org/10.1590/S0074-02762000...
; Wolff et al., 2001WOLFF, M., URIBE, A., ORTIZ, A. and DUQUE, P., 2001. A preliminary study of forensic entomology in Medellín, Colombia. Forensic Science International, vol. 120, no. 1-2, p. 53-59. http://dx.doi.org/10.1016/S0379-0738(01)00422-4. PMid:11457610
http://dx.doi.org/10.1016/S0379-0738(01)...
; Mise et al., 2007MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
, 2010MISE, KM., SOUZA, ASB., CAMPOS, CM., KEPPLER, RLF. and ALMEIDA, LM., 2010. Coleoptera associated with pig carcass exposed in a forest reserve, Manaus, Amazonas, Brazil. Biota Neotropica, vol. 10, no. 1, p. 321-324. http://dx.doi.org/10.1590/S1676-06032010000100027.
http://dx.doi.org/10.1590/S1676-06032010...
; Battán Horenstein and Linhares, 2011BATTÁN HORENSTEIN, M. and LINHARES, AX., 2011. Seasonal composition and temporal succession of necrophagous and predator beetles on pig carrion in central Argentina. Medical and Veterinary Entomology, vol. 25, no. 4, p. 395-401. http://dx.doi.org/10.1111/j.1365-2915.2011.00969.x. PMid:21762401
http://dx.doi.org/10.1111/j.1365-2915.20...
; Silva and Santos, 2012SILVA, RC. and SANTOS, WE., 2012. Fauna de Coleoptera associada a carcaças de coelhos expostas em uma área urbana no Sul do Brasil. EntomoBrasilis, vol. 5, no. 3, p. 185-189. http://dx.doi.org/10.12741/ebrasilis.v5.i3.245.
http://dx.doi.org/10.12741/ebrasilis.v5....
; Aballay et al., 2013ABALLAY, FH., ARRIAGADA, G., FLORES, GE. and CENTENO, ND., 2013. An illustrated key to and diagnoses of the species of Histeridae (Coleoptera) associated with decaying carcasses in Argentina. ZooKeys, vol. 261, no. 261, p. 61-84. http://dx.doi.org/10.3897/zookeys.261.4226. PMid:23653510
http://dx.doi.org/10.3897/zookeys.261.42...
).

Larvae and adults of Nitidulidae are generally saprophagous or xylophagous, feeding on decaying or dead wood. However, some species are exclusively phytophagous or mycophagous and others occur on carcasses or are predators (Smith, 1986; Marinoni et al., 2001MARINONI, RC., GANHO, NG., MONNÉ, ML. and MERMUDES, JRM., 2001. Hábitos alimentares em Coleoptera (Insecta). Ribeirão Preto: Holos. 63 p.). Nitidulidae was the third family in order of abundance and was almost totally represented by S. geminata with 394 individuals collected, mostly during the dry season (79.2%) (Table 2).

According to Ewing and Cline (2005)EWING, CP. and CLINE, AR., 2005. Key to adventive sap beetles (Coleoptera: Nitidulidae) in Hawaii, with notes on records and habits. The Coleopterists Bulletin, vol. 59, no. 2, p. 167-183. http://dx.doi.org/10.1649/736.
http://dx.doi.org/10.1649/736...
, S. geminata is commonly found in leaf litter and in decaying fruits and wood. The species probably plays a facultative necrophagous role in the carcasses, although it has not been observed feeding directly from it. Future studies about the biology of the species need to be done to elucidate this issue. Payne and King (1970)PAYNE, JA. and KING, EW., 1970. Coleoptera associated with pig carrion. Entomologist's Monthly Magazine, vol. 105, p. 224-232., in South Carolina, USA, found nine species of Nitidulidae during advanced stages of decomposition of pigs together with Dermestidae. However, records of the family on carcasses are rare in the Neotropical region (Wolff et al., 2001WOLFF, M., URIBE, A., ORTIZ, A. and DUQUE, P., 2001. A preliminary study of forensic entomology in Medellín, Colombia. Forensic Science International, vol. 120, no. 1-2, p. 53-59. http://dx.doi.org/10.1016/S0379-0738(01)00422-4. PMid:11457610
http://dx.doi.org/10.1016/S0379-0738(01)...
; Mise et al., 2007MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
; Battán Horenstein and Linhares, 2011BATTÁN HORENSTEIN, M. and LINHARES, AX., 2011. Seasonal composition and temporal succession of necrophagous and predator beetles on pig carrion in central Argentina. Medical and Veterinary Entomology, vol. 25, no. 4, p. 395-401. http://dx.doi.org/10.1111/j.1365-2915.2011.00969.x. PMid:21762401
http://dx.doi.org/10.1111/j.1365-2915.20...
; Silva and Santos, 2012SILVA, RC. and SANTOS, WE., 2012. Fauna de Coleoptera associada a carcaças de coelhos expostas em uma área urbana no Sul do Brasil. EntomoBrasilis, vol. 5, no. 3, p. 185-189. http://dx.doi.org/10.12741/ebrasilis.v5.i3.245.
http://dx.doi.org/10.12741/ebrasilis.v5....
).

Many larder beetles (Dermestidae) species, especially those of the genus Dermestes, are of considerable forensic importance. In cases in which the mummification of the body takes place, even indoors, these beetles can accelerate the skeletonisation process (Voigt, 1965VOIGT, J., 1965. Specific postmortem changes produced by larder beetles. Journal of Forensic Medicine, vol. 12, p. 76-80. PMid:14328682.; Schroeder et al., 2002SCHROEDER, H., KLOTZBACH, H., OESTERHELWEG, L. and PÜSCHEL, K., 2002. Larder beetles (Coleoptera, Dermestidae) as an accelerating factor for decomposition of a human corpse. Forensic Science International, vol. 127, no. 3, p. 231-236. http://dx.doi.org/10.1016/S0379-0738(02)00131-7. PMid:12175954
http://dx.doi.org/10.1016/S0379-0738(02)...
) and, even so, adults and larvae can be found associated with the remains years after death (Byrd and Castner, 2010BYRD, JH. and CASTNER, JL., 2010. Insects of forensic importance. In BYRD, JH. and CASTNER, JL. (Eds.). Forensic entomology: the utility of arthropods in legal investigations. Boca Raton: CRC Press. p. 39-126.). In the present study, three individuals of Dermestes haemorrhoidalis Küster, 1852 and 330 individuals of D. maculatus were collected. The last one is the most forensically important Coleoptera species in the Neotropical region and was more abundant in the dry season (69.4%) (Table 2).

Several authors have reported the presence of adults and larvae of D. maculatus on carcasses and corpses in Brazil (Souza and Linhares, 1997SOUZA, AM. and LINHARES, AX., 1997. Diptera and Coleoptera of potential forensic importance in southeastern Brazil: relative abundance and seasonality. Medical and Veterinary Entomology, vol. 11, no. 1, p. 8-12. http://dx.doi.org/10.1111/j.1365-2915.1997.tb00284.x. PMid:9061672
http://dx.doi.org/10.1111/j.1365-2915.19...
; Carvalho et al., 2000CARVALHO, LM., THYSSEN, PJ., LINHARES, AX. and PALHARES, FAB., 2000. A checklist of arthropods associated with pig carrion and human corpses in southeastern Brazil. Memorias do Instituto Oswaldo Cruz, vol. 95, no. 1, p. 135-138. http://dx.doi.org/10.1590/S0074-02762000000100023. PMid:10656720
http://dx.doi.org/10.1590/S0074-02762000...
; Carvalho et al., 2004CARVALHO, LML., THYSSEN, PJ., GOFF, ML. and LINHARES, AX., 2004. Observations on the succession patterns of necrophagous insects on a pig carcass in an urban area of Southeastern Brazil. Anil Aggrawal's Internet Journal of Forensic Medicine and Toxicology, vol. 5, p. 33-39.; Mise et al., 2007MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
; Rosa et al., 2011ROSA, TA., BABATA, MLY., SOUZA, CM., SOUSA, D., MELLO-PATIU, CA., VAZ-DE-MELLO, FZ. and MENDES, J., 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Revista Brasileira de Entomologia, vol. 55, no. 3, p. 424-434. http://dx.doi.org/10.1590/S0085-56262011005000045.
http://dx.doi.org/10.1590/S0085-56262011...
) and the rest of the Neotropical region (Iannacone, 2003IANNACONE, J., 2003. Artropofauna de importancia forense en un cadaver de cerdo en el Callao, Peru. Revista Brasileira de Zoologia, vol. 20, no. 1, p. 85-90. http://dx.doi.org/10.1590/S0101-81752003000100010.
http://dx.doi.org/10.1590/S0101-81752003...
; Velásquez, 2008VELÁSQUEZ, Y., 2008. A checklist of arthropods associated with rat carrion in a montane locality of northern Venezuela. Forensic Science International, vol. 174, no. 1, p. 68-70. http://dx.doi.org/10.1016/j.forsciint.2007.02.020. PMid:17386987
http://dx.doi.org/10.1016/j.forsciint.20...
; Valdes-Perezgasga et al., 2010VALDES-PEREZGASGA, MT., 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, vol. 55, no. 4, p. 1098-1101. http://dx.doi.org/10.1111/j.1556-4029.2010.01381.x. PMid:20384924
http://dx.doi.org/10.1111/j.1556-4029.20...
; Battán Horenstein and Linhares, 2011BATTÁN HORENSTEIN, M. and LINHARES, AX., 2011. Seasonal composition and temporal succession of necrophagous and predator beetles on pig carrion in central Argentina. Medical and Veterinary Entomology, vol. 25, no. 4, p. 395-401. http://dx.doi.org/10.1111/j.1365-2915.2011.00969.x. PMid:21762401
http://dx.doi.org/10.1111/j.1365-2915.20...
; Aballay et al., 2012ABALLAY, FH., MURUA, AF., ACOSTA, JC. and CENTENO, ND., 2012. Succession of carrion fauna in the arid region of San Juan Province, Argentina and its forensic relevance. Neotropical Entomology, vol. 41, no. 1, p. 27-31. http://dx.doi.org/10.1007/s13744-011-0005-9. PMid:23950006
http://dx.doi.org/10.1007/s13744-011-000...
). Mise et al. (2007)MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
, in an Araucaria forest fragment, found only 22 specimens of D. maculatus, while Rosa et al. (2011)ROSA, TA., BABATA, MLY., SOUZA, CM., SOUSA, D., MELLO-PATIU, CA., VAZ-DE-MELLO, FZ. and MENDES, J., 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Revista Brasileira de Entomologia, vol. 55, no. 3, p. 424-434. http://dx.doi.org/10.1590/S0085-56262011005000045.
http://dx.doi.org/10.1590/S0085-56262011...
, in an area of Cerrado, collected 4,874 individuals, the majority in the dry season (94.6%), the same as in the present study. Despite differences between sampling methods, it seems to be clear the divergence in the abundance status of D. maculatus among different habitats, with a preference for drier regions.

With over 19,000 species described, Scarabaeidae vary greatly in biology, ecology and behaviour (Byrd and Castner, 2010BYRD, JH. and CASTNER, JL., 2010. Insects of forensic importance. In BYRD, JH. and CASTNER, JL. (Eds.). Forensic entomology: the utility of arthropods in legal investigations. Boca Raton: CRC Press. p. 39-126.). The members of this family are mainly detritivorous, with larvae feeding on dung or pieces of decomposing carcasses (Smith, 1986; Marinoni et al., 2001MARINONI, RC., GANHO, NG., MONNÉ, ML. and MERMUDES, JRM., 2001. Hábitos alimentares em Coleoptera (Insecta). Ribeirão Preto: Holos. 63 p.). Many species dig oviposition chambers and tunnels under or near the carcasses, where pieces of the carcass are deposited to feed the larvae, which can cause postmortem movement of the bodies (Ururahy-Rodrigues et al., 2008URURAHY-RODRIGUES, A., RAFAEL, JA., WANDERLEY, RF., MARQUES, H. and PUJOL-LUZ, JR., 2008. Coprophanaeus lancifer (Linnaeus, 1767) (Coleoptera, Scarabaeidae) activity moves a man-size pig carcass: relevant data for forensic taphonomy. Forensic Science International, vol. 182, no. 1-3, p. e19-e22. http://dx.doi.org/10.1016/j.forsciint.2008.09.009. PMid:18990518
http://dx.doi.org/10.1016/j.forsciint.20...
). In the present study, this family was the fifth most abundant, with 300 individuals, mostly present in the rainy season (97%). Fifteen species were collected, especially of the subfamily Scarabaeinae, mainly represented by A. carbonarius (43.0%) and C. manni (15.7%) (Table 2), which were very abundant in a study carried out by Hernández (2007)HERNÁNDEZ, MIM., 2007. Besouros escarabeíneos (Coleoptera: Scarabaeidae) da Caatinga paraibana, Brasil. Oecologia Brasiliensis, vol. 11, no. 03, p. 356-364. http://dx.doi.org/10.4257/oeco.2007.1103.06.
http://dx.doi.org/10.4257/oeco.2007.1103...
in the same study area.

In Brazil, the studies by Luederwaldt (1911)LUEDERWALDT, G., 1911. Os insectos necrophagos paulistas. Revista do Museu Paulista, vol. 8, p. 414-433. and Pessôa and Lane (1941)PESSÔA, SB. and LANE, F., 1941. Coleópteros de interesse médico-legal. Arquivos de Zoologia do Estado de São Paulo, vol. 2, p. 389-504. about Scarabaeidae associated with carcasses of several animals in São Paulo and surrounding cities are historical references. Together, these authors recorded 124 species, most of them belonging to the genera Canthidium, Canthon, Coprophanaeus, Deltochilum and Phanaeus.

Most of Cleridae are predators of larvae of flies and other beetles, but the genus Necrobia has necrophagous habit, with a preference for carcasses and corpses in dry stages of decomposition (Payne and King, 1970PAYNE, JA. and KING, EW., 1970. Coleoptera associated with pig carrion. Entomologist's Monthly Magazine, vol. 105, p. 224-232.; Arnaldos et al., 2004ARNALDOS, MI., SÁNCHEZ, F., ÁLVARES, P. and GARCÍA, MD., 2004. A forensic entomology case from the Southeastern Iberian Peninsula. Anil Aggrawal's Internet Journal of Forensic Medicine and Toxicology, vol. 5, p. 22-25.). This family was the sixth in order of abundance and represented only by N. rufipes, with 120 specimens collected and showing preference for the dry season (61.7%) (Table 2).

Rosa et al. (2011)ROSA, TA., BABATA, MLY., SOUZA, CM., SOUSA, D., MELLO-PATIU, CA., VAZ-DE-MELLO, FZ. and MENDES, J., 2011. Arthropods associated with pig carrion in two vegetation profiles of Cerrado in the State of Minas Gerais, Brazil. Revista Brasileira de Entomologia, vol. 55, no. 3, p. 424-434. http://dx.doi.org/10.1590/S0085-56262011005000045.
http://dx.doi.org/10.1590/S0085-56262011...
collected that species in great abundance (2,321) in an area of Cerrado, most of them in the dry season as well (94.1%). Adults and larvae of N. rufipes are widely found on carcasses and corpses in Brazil (Luederwaldt, 1911LUEDERWALDT, G., 1911. Os insectos necrophagos paulistas. Revista do Museu Paulista, vol. 8, p. 414-433.; Souza and Linhares, 1997SOUZA, AM. and LINHARES, AX., 1997. Diptera and Coleoptera of potential forensic importance in southeastern Brazil: relative abundance and seasonality. Medical and Veterinary Entomology, vol. 11, no. 1, p. 8-12. http://dx.doi.org/10.1111/j.1365-2915.1997.tb00284.x. PMid:9061672
http://dx.doi.org/10.1111/j.1365-2915.19...
; Carvalho et al., 2000CARVALHO, LM., THYSSEN, PJ., LINHARES, AX. and PALHARES, FAB., 2000. A checklist of arthropods associated with pig carrion and human corpses in southeastern Brazil. Memorias do Instituto Oswaldo Cruz, vol. 95, no. 1, p. 135-138. http://dx.doi.org/10.1590/S0074-02762000000100023. PMid:10656720
http://dx.doi.org/10.1590/S0074-02762000...
, 2004CARVALHO, LML., THYSSEN, PJ., GOFF, ML. and LINHARES, AX., 2004. Observations on the succession patterns of necrophagous insects on a pig carcass in an urban area of Southeastern Brazil. Anil Aggrawal's Internet Journal of Forensic Medicine and Toxicology, vol. 5, p. 33-39.; Mise et al., 2007MISE, KM., ALMEIDA, LM. and MOURA, MO., 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia, vol. 51, no. 3, p. 358-368. http://dx.doi.org/10.1590/S0085-56262007000300014.
http://dx.doi.org/10.1590/S0085-56262007...
; Silva and Santos, 2012SILVA, RC. and SANTOS, WE., 2012. Fauna de Coleoptera associada a carcaças de coelhos expostas em uma área urbana no Sul do Brasil. EntomoBrasilis, vol. 5, no. 3, p. 185-189. http://dx.doi.org/10.12741/ebrasilis.v5.i3.245.
http://dx.doi.org/10.12741/ebrasilis.v5....
) and elsewhere in the Neotropical region (Wolff et al., 2001WOLFF, M., URIBE, A., ORTIZ, A. and DUQUE, P., 2001. A preliminary study of forensic entomology in Medellín, Colombia. Forensic Science International, vol. 120, no. 1-2, p. 53-59. http://dx.doi.org/10.1016/S0379-0738(01)00422-4. PMid:11457610
http://dx.doi.org/10.1016/S0379-0738(01)...
; Iannacone, 2003IANNACONE, J., 2003. Artropofauna de importancia forense en un cadaver de cerdo en el Callao, Peru. Revista Brasileira de Zoologia, vol. 20, no. 1, p. 85-90. http://dx.doi.org/10.1590/S0101-81752003000100010.
http://dx.doi.org/10.1590/S0101-81752003...
; Valdes-Perezgasga et al., 2010VALDES-PEREZGASGA, MT., 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, vol. 55, no. 4, p. 1098-1101. http://dx.doi.org/10.1111/j.1556-4029.2010.01381.x. PMid:20384924
http://dx.doi.org/10.1111/j.1556-4029.20...
; Battán Horenstein and Linhares, 2011BATTÁN HORENSTEIN, M. and LINHARES, AX., 2011. Seasonal composition and temporal succession of necrophagous and predator beetles on pig carrion in central Argentina. Medical and Veterinary Entomology, vol. 25, no. 4, p. 395-401. http://dx.doi.org/10.1111/j.1365-2915.2011.00969.x. PMid:21762401
http://dx.doi.org/10.1111/j.1365-2915.20...
; Aballay et al., 2012ABALLAY, FH., MURUA, AF., ACOSTA, JC. and CENTENO, ND., 2012. Succession of carrion fauna in the arid region of San Juan Province, Argentina and its forensic relevance. Neotropical Entomology, vol. 41, no. 1, p. 27-31. http://dx.doi.org/10.1007/s13744-011-0005-9. PMid:23950006
http://dx.doi.org/10.1007/s13744-011-000...
).

The diversity of Coleoptera fauna was different from other studies carried out in the Neotropical region, which may be a result not only from different environments, but also the capture method adopted. Ten species were recorded for the first time associated with carcasses in the Neotropical region. Populations of Staphylinidae and Histeridae, the most abundant families, can be considered as affected by seasonality, with only 13.4% and 34.2% of the specimens collected, respectively, in the dry season. Amongst the most collected species, X. diptychus showed to be strongly influenced by the dry season, since 96.1% of the specimens were collected during that period. A. iheringi, representing 34.7% of the total, was the most abundant species. Future studies are required to confirm the role of these beetles on carcasses and its possible forensic potential to Caatinga. In addition, the exceptional Coleoptera fauna found reinforces the need of regional studies about the composition of insects associated with carcasses throughout Neotropical region.

Acknowledgements

The authors are thankful to Dr. Edilson Caron (UFPR), Dr. Daniel P. Moura (UFPR) and M.Sc. Carolina N. Liberal (UFPB) for their help with the identifications; Dr. Carla L. Bicho (UEPB), Dr. Patricia J. Thyssen (UFPel) and Dr. Rodrigo C. A. P. Farias (UFPB) for critical comments on the manuscript. Thanks are also due to the “Conselho Nacional de Desenvolvimento Científico e Tecnológico” (CNPq) and the “Coordenação de Aperfeiçoamento de Pessoal de Nível Superior” (CAPES) for providing scholarships to A. C. F. Alves and W. E. Santos.

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Publication Dates

  • Publication in this collection
    Aug 2014

History

  • Received
    11 Mar 2013
  • Accepted
    15 May 2013
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