SciELO - Scientific Electronic Library Online

 
vol.53 issue2New species of Anthrenoides Ducke (Hymenoptera, Apidae, Andreninae) from ArgentinaComparative external morphology of the three species of the Telchin licus (Drury) complex (Lepidoptera, Castniidae) with a synonymy author indexsubject indexarticles search
Home Pagealphabetic serial listing  

Revista Brasileira de Entomologia

Print version ISSN 0085-5626

Rev. Bras. entomol. vol.53 no.2 São Paulo June 2009

http://dx.doi.org/10.1590/S0085-56262009000200006 

SYSTEMATICS, MORPHOLOGY AND BIOGEOGRAPHY

 

Diagnosis and key of the main families and species of South American Coleoptera of forensic importance1

 

Diagnose e chave de identificação para as principais famílias e espécies de Coleoptera de importância forense da América do Sul

 

 

Lúcia M. AlmeidaI; Kleber M. MiseII

IDepartamento de Zoologia, Universidade Federal do Paraná, Caixa Postal 19020, 81581-980 Curitiba-PR, Brasil. Fellowship CNPq. lalmeida@ufpr.br
IIFellowship CAPES. klebermise@yahoo.com.br

 

 


ABSTRACT

The objective of this paper is to provide diagnosis and keys of the families and species, with illustrations of the main groups. A table of all related species recorded from South America is presented, including the substrate in which they were collected and their geographical distribution. The list comprises 221 species included in 15 families, of which 70% of the species are from Brazil. Scarabaeidae is the most diverse family with 121 species, followed by Staphylinidae with 68. Also we provide one database of Coleoptera species associated with carcasses in South America.

Keywords: Forensic entomology; necrophilous beetles; Neotropical region; taxonomy.


RESUMO

O objetivo deste trabalho é apresentar diagnoses e chaves de identificação das principais famílias e espécies de importância forense, com ilustrações dos principais grupos. É apresentada uma tabela de todas as espécies de ocorrência na América do Sul, incluindo o substrato nas quais foram coletadas e sua distribuição geográfica. A lista compreende 221 espécies incluídas em 15 famílias, das quais pelo menos 70% das espécies são distribuídas no Brasil. Scarabaeidae é a família com maior diversidade com 121 espécies, seguida por Staphylinidae com 68. Também é fornecida uma base de dados para as espécies de Coleoptera associadas a carcaças na América do Sul.

Palavras-chave: Besouros necrófilos; entomologia forense; neotropical; taxonomia.


 

 

Coleoptera is the second largest order of forensic interest, with several necrophagous representatives, most being predators but their feeding habit may change between larval stages and adulthood. The species of Coleoptera increase in number both of individuals and species during advanced stages of decomposition in open environment and are absent or less represented indoors (Goff 1991). Beetles are encountered in great numbers during the faunal succession process, moreover their biological traits may be used to estimate the post mortem interval.

According to Smith (1986) the families of Coleoptera of forensic interest are: Carabidae, Hydrophilidae, Silphidae, Leiodidae, Staphylinidae, Histeridae, Cleridae, Anthicidae, Dermestidae, Nitidulidae, Rhizophagidae, Ptinidae, Tenebrionidae, Scarabaeidae, Geotrupidae and Trogidae.

In Brazil one of the most comprehensive studies of this fauna was done by Luederwaldt (1911), who found about 62 species related to carcasses. Subsequently papers were mainly focused on Diptera, but some of them reported Coleoptera species.

Some authors aimed on specific families contributing with taxonomic or survey studies of necrophilous beetles. Pessôa & Lane (1941) did studies on Scarabaeinae fauna of legal medicine interest, found in São Paulo and neighborhood (southeast of Brazil) comprising 113 species of 26 genera.

The fauna of Staphylinidae was studied by Jimenez-Sanchez et al. (2000) on Nanchititla, Mexico. The 50 species in their study were collected with traps using squid as bait.

Scampini et al. (2002) found six species of Carabidae from Buenos Aires (Santa Catalina), Argentina, using pig carcasses and pit-fall traps during three months.

Souza & Linhares (1997) studied Diptera and Coleoptera collected in pig carcass and reported 13 species in five families, of which Staphylinidae had greater number of species, only two being identified to the species level.

A study conducted by Mise et al. (2007) focusing on Coleoptera took place in Curitiba for one year using pig carcass with Shannon modified, pit-fall traps and active sampling. There were found 4,360 beetles belonging to 112 species of 26 families, 12 of them were considered of forensic potential.

In South America and in Brazil, the majority of studies focuses mainly on Diptera, one of the reasons is due to difficulties in identifying the species of Coleoptera. There are no published records of Coleoptera of forensic importance in South America in a checklist and this data will be useful to start a database of the fauna of Coleoptera associated with different kinds of carcasses.

Due to the importance of the beetles to forensic sciences and the few studies in the Coleoptera fauna of South America, the purpose of this paper is to provide data about the families and species, with brief descriptions, identification keys and illustrations for the main groups. Also when available the alimentary habits of the species had been added.

 

MATERIAL AND METHODS

Taxa were arranged in alphabetical order at family, genera and species level, when possible, followed by the substratum, geographical distribution and references. White (1983), Booth et al.(1990), Kingsolver (1991), Borror et al.(1992), Costa (2000), Newton et al. (2005) and Costa et al. (2006) were used for the families descriptions, their ecological data and number of genera and species.

The list of the important groups of Coleoptera show different species found on different animal carcasses across the Neotropical region. Many species were reported only once because there is a huge difference among regions. This could be due to the endemism of certain species, outlining the importance of regional surveys, which could indicate the species of Coleoptera of potential forensic importance.

The keys from the most general to the most specific, deal with beetles associated with decomposing animal carcasses. Accidental families, genera and species are not included because the lack of specimens or literature.

 

RESULTS

The list of the main Coleoptera species of forensic importance has 221 species included in 15 families; however this is a conservative estimate, considering that most of the specimens cited in papers are not identified at species level (Table I). At least 70% of the species occurs in Brazil, being Scarabaeidae the most diverse family with 122 species, followed by Staphylinidae with 69. Also we provide a diagnosis and keys of the families and species, with illustrations to enable the identification of the main groups.

Key of South American Coleoptera of forensic importance

1.   Notopleural suture distinct; metatrochanter very large; first visible abdominal sternite divided by metacoxae (Fig. 1) ........ (Suborder Adephaga) Carabidae
    Without notopleural suture; first visible abdominal sternite entire (Fig. 2) ...... ....................................................................... (Suborder Polyphaga) 2
     
2.   Antennae lamellated (Fig. 3) .............................................................. 3
    Antennae not lamellated ................................................................... 4
     
3.   Clypeus large covering labrum in dorsal view; mandible usually hidden from above; abdomen with 5 or 6 visible segments ....................... Scarabaeidae
    Head bent down almost hypognathous; labrum distinct and bent down; mandible proeminent not covered by clypeus; elytra often with tubercles (Fig. 4); abdomen with 5 visible segments .................................. Trogidae
     
4.   Head prognathous; elytra usually very short and truncate, exposing more than three abdominal tergites (Fig. 5); antennae filiform or moniliform, occasionally with weak club (Fig. 6); tarsal formula variable, 3-3-3 to 5-5-5; abdomen usually capable of being flexed .............................. Staphylinidae
    Without the above combination of characters ....................................... 5
     
5.   Antennae with a compact club; elytra often short and truncate exposing abdominal tergites or entire ............................................................... 6
    Without the above combination of characters ....................................... 7
     
6.   Procoxae transverse with exposed trochantin (Fig. 7); tarsal formula often 5-5-5 with 4th reduced ......................................................... Nitidulidae
    Procoxae transverse without exposed trochantin; elytra often short and truncate exposing pygidium and propygidium (Fig. 8); tibiae flattened with spines or teeth, tarsal formula 5-5-5, rarely 5-5-4 ...................... Histeridae
     
7.   Body usually oval in shape; sometimes parallel sided; antennae short with a four segmented club, basal segment shiny, the apical three segments pubescent; maxillary palpi long often longer than antennae; procoxae conical; tibiae often spinose with two usually large spurs; tarsi usually 5-5-5; abdomen with 5 visible sternites ........................................ Hydrophilidae
    Without the above combination of characters ....................................... 8
     
8.   Tarsal formula 5-5-4 ....................................................... Tenebrionidae
    Tarsal formula 5-5-5 ........................................................................ 9
     
9.   Head flattened, inserted in broad imargination of pronotum; antennae 10-11 segmented usually capitate with a loose club but may be compact, club 3-5 segmented; abdomen with 5 visible sternites the sixth often partially visible ......................................................................................... Leiodidae
   

Body castaneus covered with decumbent hairs; antennae with a loose club, the eighth segment reduced and the eleventh pointed; approximately 2,33 mm (Fig. 19) ..................................... Dissochaetus murray Reitter, 1884

    Without the above combination of characters ...................................... 10
     
10.   Head hypognathous (Fig. 9) ............................................................. 11
    Head not as above; antennae sub-clubbed or clubbed with 10 or 11 segments (Fig. 10), segments 9-11 with dense pubescence; trochantin exposed (Fig. 7); elytra often with longitudinal striae, color black, testaceous, or black with orange or yellow markings, sometimes with pronotum tomentose, parcial or totally colored of yellow or reddish .............. ........................................................................................... Silphidae
     
11.  

Metacoxae often excavated for reception of metafemora; frons often with median ocellus; antennae short usually clubbed often received into grooves on underside of prothorax; five visible sternites ...................... Dermestidae

    Body elongated, covered with bristly hairs; frons oblique and eyes large; antennae not received into grooves on underside of prothorax; pronotum narrower than elytra, often nearly cylindrical; procoxae usually conical; tarsal formula 5-5-5 or sometimes pseudotetramerous ............................ Cleridae

Key to species of Cleridae

1.  

Head, pronotum and elytra base reddish brown and the rest of the elytra metallic blue; approximately 4,77 mm (Fig. 20) ......................................... ....................................................... Necrobia ruficollis (Fabricius, 1775)

    Body metallic blue (Fig. 21); approximately 5,17 mm ................................. .......................................................... Necrobia rufipes (De Geer, 1775)

Key to species of Dermestidae

1.  

Elytra apex serrate with a small terminal spine (Figs. 22 e 23); approximately 9,16 mm ...................................... Dermestes maculatus (De Geer, 1774)

    Elytra apex entire lacking serration and spines ...................................... 2
     
2.   Abdominal venter without pattern; lateral sulcus of abdominal sternum I closely parallel to lateral margin; approximately 8,5 mm ............................ ................................................... Dermestes peruvianus Laporte, 1840
    Abdominal venter patterned (Fig. 11); lateral sulcus of abdominal sternum I not closely parallel to lateral margin; approximately 8 mm ......................... ...........................................................Dermestes ater (De Geer, 1774)

Key to species of Histeridae

1.  

Small sized beetles (<0,3mm); antennal insertions usually exposed; elytra at most with vague impressions; scutellum hidden; hind tarsi with four segments .............................................................................. Aeletes

    Without the above combination of characters; large sized beetles (>0,3mm) .................................................................................................. 2
     
2.  

Prosternum with an antennal cavity for reception of antennae ............... 3

    Prosternum not as above ................................................................ 4
     
3.   Prosternal process with a preapical fovea (Fig. 12) ................... Euspilotus
    Lateral lobe of the abdominal 8th tergite with rounded apex; pygidial line rounded at apex (sometimes weakly sinuated or interrupted); size approximately 7,66 mm (Fig. 24) ......... Euspilotus nigrita (Blanchard, 1842)
    Prosternal process without a preapical fovea .............................. Saprinus
     
4.   Antennal club with two oblique sutures, V-shaped (Figs. 13 e 25) ............. ....................................................................................... Omalodes
    Pronotum with two fovea on the lateral sides; size approximately 11 mm (Fig. 25) .................................... Omalodes bifoveolatus (Marseul, 1853)
    Antennal club not as above ............................................................. 4
     
5.   Antennal club with two annuli (Fig. 14); frontal suture, if present, distant from antennal base (Fig. 26) ..................................................... Hister
    Antennal club with only a straight, transverse subapical annulus of short setae; frontal striae reaching the antennal base, if present (Fig. 27) ......... .......................................................................................... Phelister

 


 

Key to species of Scarabaeidae

1.   Mid coxae with external lateral border parallel to the body axis and located in the lateral of the metasternum in the limit of the lateral border of the body (Fig. 28) ......................................................................... Eurysternus
    Without the above combination of characters ...................................... 2
     
2.   Elytral apex with carinae or distinct tubercles between the interstriae ........ ...................................................................................... Deltochilum
    Clypeus with four teeth; elytrae with numerous striae; metallic colored; size approximately 26 mm (Fig. 29) ............... Deltochilum icarus (Olivier, 1789)
    Without the above combination of characters ...................................... 3
     
3.  

Body length less than 12 mm; elytra and pronotum hairy; first tarsal segment of posterior leg almost as longer as the others together (Fig. 30) .......................... .............................................................................................Onthophagus

    Without the above combination of characters ............................................. 4
     
4.   Pigidium flattened, with dull shine ..................................................... Canthon
   

Pronotum shining yellow colored; elytrae brownish-yellow and opacous; size approximately 11 mm (Fig. 31) ...................... Canthon triangularis (Drury, 1770)

    Without the above combination of characters .............................................. 5
     
5.  

Body oval elongated; colour usually black, sometimes green or blue; abdominal sternites fused and very short (Fig. 32) .......................................... Ontherus

    Without the above combination of characters .............................................. 6
     
6.   Antennal club with segments wide and flattened; anterior margin of clypeus with three emarginations, with two proeminent teeth and two lateral round angles; first segment of mid and hind tarsus elongated, longer than wide ............................ ......................................................................................... Coprophanaeus
    Metallic bluish colored with integument hardly puncturate; size approximately 37 mm (Fig. 33) ..................................... Coprophaneus lancifer (Linnaeus, 1767)
    Metallic bluish colored with integument smooth; size approximately 19 mm (Fig. 34) .........................................................Coprophaneus saphirinus (Sturm, 1826)
     
7.  

Antennal club with segments elongated and thin; anterior margin of clypeus without emargination, with two short teeth .................................. Dichotomius

    Protibiae with a slender spine in the apice; size approximately 22 mm (Fig. 35) ..... ............................................................... Dichotomius boreus (Olivier, 1789)

Key to species of Silphidae

1.  

With a tooth in the humeral region of the elytra; size approximately 17 mm (Fig. 36) ....................................... Oxelytrum discicolle (Brullé, 1840)

    Without a tooth or only with a small protuberance in the humeral region of the elytra; size approximately 16 mm ................................................ 2
     
2.   Pronotum with a maculae quadrangular-shaped, occupying most of the pronotal disk (Fig. 37) ................. Oxelytrum erythrurum (Blanchard, 1840)
    Pronotum with uniform color or with the maculae rounded, occupying the center of the pronotal disk (Fig. 38) ... Oxelytrum cayennense (Sturm, 1826)

Key to species of Staphylinidae

1.   Antennae inserted before anterior margin of eyes ................................ 2
    Antennae inserted between eyes (Fig. 15) ........................... Aleocharinae
    Maxilar palpi with five segments, the apical minute (pseudosegment) (Figs. 15 e 39) ............................................................................... Aleochara
    Maxilar palpi with four segments, without pseudosegment (Fig. 40) ............. ............................................................................................ Atheta
     
2.   Abdomen with seven visible sternites ..................................... Oxytelinae
    Scutellum with a diamond shaped impression; abdominal tergum II with curved basal lateral ridge ...................................................... Oxytelus
    Scutellum with a tri- or bilobed impression; abdominal tergum II without curved basal lateral ridge (Fig. 41) .......................................... Anotylus
     
2'.   Abdomen with six visible sternites ...................................... Staphylininae
a.  

The first visible segment of abdominal terga with impressed curved line ........ .................................................................................... Xanthopygus

    Head and pronotum green bluish colored; elytrae and basal segments of abdomen yellow brownish; size approximately 14 mm (Fig. 44) .................... ..................................................... Xanthopygus bicolor (LaPorte, 1835)
a'.   Without above combination of characters ............................................. b
b.   With sclerotized plate in front of prosternum (Figs. 16) .............................. ............................................................................................ Eulissus
    Body green bluish, size approximately 21 mm (Fig. 45) .............................. ................................................. Eulissus chalybaeus (Mannerheim, 1830)
b'.   Without sclerotized plate in front of prosternum .................................... c
c.   Pronotum with translucent post-coxal process ........................ Platydracus
    Head, pronotum and elytrae metallic bluish; abdomen black, with the three last segments yellow colored; size approximately 16 mm (Fig. 46) ................ ............................................... Platydracus ochropygus (Nordman, 1837)
c'.   Without above combination of characters ........................................... d
d.   Apical segment of maxilar palpi twice the size of the penultimate segment (Figs. 17, 42) ................................................................... Belonuchus
d'.   Apical segment of maxilar palpi more or less of the same size of the penultimate segment (Figs. 18, 43) ......................................... Philonthus

Key to species of Trogidae

1.  

Clypeus forming a 90º angle with frons (Figs. 47, 48) ............... Polynoncus

    Clypeus not forming a 90º angle with frons (Figs. 49, 50) ............ Omorgus

 



 

Characteristics of families and species

Carabidae. Ground beetles. A family with about 1,500 genera and 30,000 species, with 336 genera and 6,258 species in the Neotropical region. Usually found under stones, logs, leaves, bark and debris. The larvae and adults hide during the day and feed at night, preferring to walk rather than fly. Most of them are predators of other insects and may be used in biological control. Some species feed on dead or dying insects and others on living insects, such as maggots, beetle larvae, etc.

Cleridae. Checkered beetles. A family with about 150 genera and 4,000 species, with 61 genera and 886 species in the Neotropical region. They attack insects specially wood-boring beetles. The adults are very active specially during the day. They are often found on flowers, foliage and tree trunks. Necrobia ruficollis (Fabricius, 1775) and Necrobia rufipes (DeGeer, 1775) occur in carrion and products of animal origin.

Necrobia rufipes is predator of dipteran and coleopteran larvae. It is also associated with stored meats, such as dry fish, skin, dead animal bones, some oily seeds and stored products, mainly those with high protein indices, being also found in animal rations (Gredilha et al. 2005; Ashman 1963).

Dermestidae. Odd beetles. A family with about 45 genera and 850 species, with 20 genera and 248 species in the Neotropical region. Also known as skin beetles, they are primarily scavengers that feed on dried skin and other soft remains of animals such as fur, feathers, wool and leather. They also feed on carpets, silk, dried meats and dead insects. Some are pests of storage products such as grains, seeds, corks or cereal products.

Dermestes maculatus De Geer is a notorious pest of dried fish and fish meal, is known to damage wooden frames as well as polystyrene and glass fibre wadding in premises when the last instar larva is about to pupate (Turner 1986; Wildey & Wayman 1979). The insect pests of dried animal products also attack living insects Dermestes spp. on silkworm pupae and adults (Kumar et al. 1988; Veer et al. 1996). Use of infested woolen materials can cause allergic reactions like urticarial and papulovesicular lesions in man (Ahmed et al. 1981).

Histeridae. Clown beetles. A family with about 200 genera and 3,000 species, with 139 genera and 1,047 species in the Neotropical region. They are mainly predators of soft body insects larvae and eggs, particularly those of Cyclorraphan Diptera. Most occur in carrion, dung, decomposing plant materials, such as fungi, and tree wounds. Some live under loose bark or in galleries of wood-boring insects, where they prey on other organisms. The greatly flattened species live under bark of dead or dying trees. Cylindrical species occur in tunnels of bark beetles and other wood-boring insects. Most species are neither flattened nor cylindrical and are abundant in the early stages of decay of carcasses.

Hydrophilidae. Water scavenger beetles. A family with about 140 genera and 2,025 species, with 42 genera and 3,064 species in the Neotropical region. Larvae and adults of many are aquatic or semiaquatic, but some live in fresh mammal dung, humus-rich soil, or decaying leaves. Adults are mainly scavengers but the larvae are usually predacious. The terrestrial species occur in dung.

Leiodidae. Round fungus beetles. A family with about 334 genera and 4,240 species, with 38 genera and 279 species in the Neotropical region. Most species occur in carrion but some are found in fungi, some feed on slime molds, and others occur in ant nests.

Nitidulidae. Sap beetles. A family with about 160 genera and 3,000 species, with 79 genera and 770 species in the Neotropical region. They have a varied biology. Many are phytophagous, including pollen feeders, seeds, tree sap, others feed on dead or decaying plants, rotten fruits, leaf litter and a few are carrion feeders. Some are predators of scale insects.

Scarabaeidae. Scarab beetles. A family with about 2,000 genera and 25,000 species, with 362 genera and 4,706 species in the Neotropical region. Adults and larvae may be detritivorous, saprophagous, herbivorous, necrophagous or coprophagous. Larvae of most injurious species live in soil, feeding on roots, others may feed in rotten wood, dry carrion or skins.

Silphidae. Carrion beetles. A family with about 14 genera and 175 species, with 9 genera and 82 species in the Neotropical region. Most common on carrion but sometimes found on decaying vegetation or living plants. They feed on maggots and also are associated with vertebrate carcasses.

Staphylinidae. Rove beetles. A family with about 659 genera and 48,000 species, with 652 genera and 8,124 species in the Neotropical region. Adults are found in a wide range of habitats, under stones and other objects on ground, along shores of streams and lakes. Some live along ocean shores, on carrion, in manure, on fungi, on flowers in ant and termite nests, under bark, in soil or soil litter and in caves. They are among the most proficient fliers of all beetles. Many species run with the tip of abdomen raised. Larvae usually occur in the same habitat as adults. Most species are saprophagous feeding on dead or decaying plant or animal materials, such as carrion, dung, dead logs, etc. Most species are predacious and a few are parasites of other insects.

Tenebrionidae. Darkling beetles. A family with about 1,700 genera and 18,000 species, with 478 genera and 4,624 species in the Neotropical region. Adults and larvae live in a variety of terrestrial habitats. Some lives in rooting wood, on plant materials, under logs and stones, in termite and ant nests, in houses, in fungi and in debris. They feed on material of plant origin including decaying plant litter, dead wood, fungal fruiting bodies. Some feed on dead animal material and a few are predators. A few species are pests of stored products and the root-feeding larvae can also be agricultural pests specially of young plants and during dry conditions.

Trogidae. Hide beetles. A family with about 5 genera and 300 species, with 3 genera and 48 species in the Neotropical region. Adults and larvae are among the last inhabitants of fish, amphibians, reptiles, birds and mammals carcasses. The species of the genus Trox occur in nests of birds and burrows of mammals, especially those with masses of fur or feathers. When disturbed these beetles draw in their legs and lie motionless resembling dirt and rubbish and are often overlooked.

Acknowledgments. The authors thank to doctors José Roberto Pujol-Luz, Universidade de Brasília; José Albertino Rafael and Ruth Leila Ferreira Keppler, Instituto Nacional de Pesquisas da Amazônia; Marta Wolff, Instituto de Biología de la Universidad de Antioquia; Janyra Oliveira-Costa, Universidade Castelo Branco and Rodrigo Krüger, Universidade Federal de Viçosa, for loan of the material. To Nicolas Dégallier; Carla de Lima Bicho (Histeridae); Pedro Gnaspini Netto (Leiodidae); Edilson Caron (Staphylinidae) and Paschoal Coelho Grossi (Scarabaeidae) for aid in identifying the material.

 

REFERENCES

Ahmed, A. R.; R. Moy; A. R. Barr & Z. Prize. 1981. Carpet beetle dermatitis. Journal of the American Academy of Dermatology 5: 428-432.         [ Links ]

Ashman, F. 1963. Factors affecting the abundance of the copra beetle Necrobia rufipes (De Geer) (Coleoptera, Cleridae). Bulletin of Entomological Research 53: 671-680.         [ Links ]

Booth, R. G.; M. L. Cox & R. B. Madge. 1990. 3. Coleoptera IIE. Guides to insects of importance to man. Cambridge, Cambridge University Press, vi+384 pp.         [ Links ]

Borror, D. J.; C. A. Triplehorn & N. F. Johnson. 1992. An Introduction to the study of insects (6ª ed.). Fort Worth, Saunders College Publishing, Harcourt Brace College Publishers, xiv+875 pp.         [ Links ]

Carvalho, L. M. L.; P. J. Thyssen; A. X. Linhares & F. A. B. Palhares. 2000. A checklist of Arthropods associated with pig carrion and human corpses in southeastern Brazil. Memórias do Instituto Oswaldo Cruz 95: 135-138.         [ Links ]

Carvalho, L. M. L.; P. J. Thyssen; M. L. Goff & A. X. Linhares. 2004. Observations on the succession patterns of necrophagous insects on a pig carcass in an urban area of southeastern Brazil. Aggrawal's Internet Journal of Forensic Medicine and Toxicology 5: 33-39.         [ Links ]

Centeno, N.; M. Maldonado & A. Oliva. 2002. Seasonal patterns of arthropods occurring on sheltered and unsheltered pig carcasses in Buenos Aires province (Argentina). Forensic Science International 126: 63-70.         [ Links ]

Costa, C.; S. Ide & C. E. Simonka. 2006. Insetos imaturos. Metamorfose e identificação. Ribeirão Preto, Holos Editora, 249 p.         [ Links ]

Costa, C. 2000. Estado de conocimiento de los Coleoptera neotropicales, pp. 99-114. In: F., Martín-Piera, J. J. Morrone, & A. Melic (eds). PrIBES. Proyecto Iberoamericano de Biogeografía y Entomología Sistemática. Zaragoza. Vol. 1. Villa de Leyva, Colômbia, Sociedad Entomológica Aragonesa, 326p.         [ Links ]

Cruz, T. M. & S. D. Vasconcelos. 2006. Entomofauna de solo associada à decomposição de carcaça suína em um fragmento de Mata Atlântica de Pernambuco, Brasil. Biociências 14: 193-201.         [ Links ]

Goff, M. L. 1991. Comparison of insect species associated with decomposing remains recovered inside dwellings and outdoors on the island of Oahu, Hawaii. Journal of Forensic Sciences 36: 748-753.         [ Links ]

Gredilha, R.; P. R. Saavedra; L. G. Guerim; A. F. Lima & N. M. Serra-Freire. 2005. Ocorrência de Oryzaephilus surinamensis Linnaeus, 1758 (Coleoptera: Cucujidae) e Necrobia rufipes DeGeer, 1775 (Coleoptera: Cleridae) infestando rações de animais domésticos. Entomologia Y Vectores, 12: 95-103.         [ Links ]

Iannacone, J. 2003. Artropofauna de importancia forense en un cadáver de cerdo en el Callao, Perú. Revista Brasileira de Zoologia 20: 85-90.         [ Links ]

Jimenez-Sanchez, E.; J. L. Navarrete-Heredia & J. R. Padilla-Ramirez. 2000. Estafilinidos (Coleoptera: Staphylinidae) necrofilos de la sierra de Nanchititla, Estado de Mexico, Mexico. Folia de Entomologia Mexicana 108: 53-78.         [ Links ]

Jirón, L. F. & V. M. Cartín. 1981. Insect succession in the decomposition of a mammal in Costa Rica. Journal of New York Entomological Society 89: 158-165.         [ Links ]

Kingsolver, J. M. 1991. Dermestid Beetles (Dermestidae, Coleoptera), p. 115-135. In: J.R. Gorham (ed.) Insect and Mite Pests in Food. Vol. 1. Washington, D.C., U.S. Government Printing Office, vii+310 p.         [ Links ]

Kumar, P.; C. A. Jayaprakas; B. D. Singh & K. Sengupta. 1988. Studies on the biology of Dermestes ater (Coleoptera: Dermestidae) - a pest of silkworm pupae and adults. Current Science 57: 1253.         [ Links ]

Luederwaldt, G. 1911. Os insectos necrophagos paulistas. Revista do Museu Paulista 8: 414-433.         [ Links ]

Márquez-Luna, J. 2001. Especies necrofilas de Staphylinidae (Insecta: Coleoptera) del municipio de Tlayacapan, Morelos, Mexico. Folia Entomologica Mexicana 40: 93-131.         [ Links ]

Mise, K. M.; L. M. Almeida & M. O.Moura. 2007. Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia 51: 358-368.         [ Links ]

Moura, M. O.; C. J. B. de Carvalho & E. L. A. Monteiro-Filho. 1997. A preliminary analysis of insects of medico-legal importance in Curitiba, state of Paraná. Memórias do Instituto Oswaldo Cruz 92: 269-274.         [ Links ]

Monteiro-Filho, E. L. A. & J. L. Penereiro. 1987. Estudo de decomposição e sucessão sobre uma carcaça animal numa área do estado de São Paulo, Brasil. Revista Brasileira de Biologia 47: 289-295.         [ Links ]

Newton, A. F.; C. Gutiérrez Chacón & D. S. Chandler. 2005. Checklist of the Staphylinidae (Coleoptera) of Colombia. Biota Colombiana 6: 1-72.         [ Links ]

Pessôa, S. B. & F. Lane. 1941. Coleópteros Necrófagos de Interêsse Médico-Legal. Ensaio monográfico sobre a família Scarabaeidae de S. Paulo e regiões visinhas. Revista do Museu Paulista 25: 389-504.         [ Links ]

Scampini, E.; A. Cichino & N. Centeno. 2002. Especies de Carabidae (Coleoptera) asociadas a cadavers de cerdo (Sus scrofa L.) en Santa Catalina (Buenos Aires, Argentina). Revista de la Sociedad Entomologica Argentina 61: 85-88.         [ Links ]

Smith, K. G. V. 1986. A manual of forensic entomology. Ithaca, NY, Cornell University Press, 205 p.         [ Links ]

Souza, A. M. & A. X. Linhares. 1997. Diptera and Coleoptera of potential forensic importance in southeastern Brazil: relative abundance and seasonality. Medical and Veterinary Entomology 11: 8-12.         [ Links ]

Turner Jr., E.C. 1986. Structural and litter Pests. Poultry Science 65: 644-648.         [ Links ]

Veer, V.; B. K. Negi & K. M. Rao. 1996. Dermestid beetles and some other insect pests associated with stored silkworm cocoons in India, including a world list of dermestid species found attacking this commodity. Journal of Stored Products Research 32: 69-89.         [ Links ]

Velásquez, Y. 2007. A checklist of arthropods associated with rat carrion in a montane locality of northern Venezuela. Forensic Science International 174: 68-70.         [ Links ]

White, R. E. 1983. A field guide to the beetles of North America. The Peterson field guide series 29. Boston, Houghton Mifflin Company, 368 p.         [ Links ]

Wildey, K. B. & C. Wayman. 1979. The hide beetle (Dermestes maculatus) as a deep pit pest in poultry houses. ADAS Quarterly Review 34: 187-193.         [ Links ]

Wolff, M.; A. Uribe; A. Ortiz & P. Duque. 2001. A preliminary study of forensic entomology in Medellín, Colombia. Forensic Science International 120: 53-59.         [ Links ]

 

 

Received 31/10/2008; accepted 20/03/2009

 

 

1 Contribution number 1768 of the Department of Zoology, Universidade Federal do Paraná, Brazil.