ZooKeys 261: 61–84, doi: 10.3897/zookeys.261.4226
An illustrated key to and diagnoses of the species of Histeridae (Coleoptera) associated with decaying carcasses in Argentina
Fernando H. Aballay 1, Gerardo Arriagada 2, Gustavo E. Flores 1, Néstor D. Centeno 3
1 Laboratorio de Entomología, Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA, CCT CONICET Mendoza), Casilla de correo 507, 5500 Mendoza, Argentina
2 Sociedad Chilena de Entomologia
3 Laboratorio de Entomología Aplicada y Forense, Universidad Nacional de Quilmes, Roque Sáenz peña 180, B1876BXD, Bernal, Buenos Aires, Argentina

Corresponding author: Fernando H. Aballay (faballay@mendoza-conicet.gob.ar)

Academic editor: M. Catherino

received 31 October 2012 | accepted 21 December 2012 | Published 24 January 2013


(C) 2013 Fernando H. Aballay. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


For reference, use of the paginated PDF or printed version of this article is recommended.

Abstract

A key to 16 histerid species associated with decaying carcasses in Argentina is presented, including diagnoses and habitus photographs for these species. This article provides a table of all species associated with carcasses, detailing the substrate from which they were collected and geographical distribution by province. All 16 Histeridae species registered are grouped into three subfamilies: Saprininae (twelve species of Euspilotus Lewis and one species of Xerosaprinus Wenzel), Histerinae (one species of Hololepta Paykull and one species of Phelister Marseul) and Dendrophilinae (one species of Carcinops Marseul). Two species are new records for Argentina: Phelister rufinotus Marseuland Carcinops troglodytes (Paykull). A discussion is presented on the potential forensic importance of some species collected on human and pig carcasses.

Keywords

Key, Histeridae, Saprininae, forensic, carcasses, Argentina

Introduction

Coleoptera is one of the major orders of insects represented on carcasses and its forensic importance has been frequently documented (Benecke 1988, Kulshrestha and Satpathy 2001, Schroeder et al. 2002). Among the most important families mentioned in the literature are: Dermestidae, Cleridae, Histeridae, Staphylinidae, Nitidulidae, Scarabaeidae, Tenebrionidae, and Trogidae (Mise et al. 2007, Özdemir and Set 2009, Almeida and Mise 2009, Battán Horenstein and Linhares 2011). Members of these families are associated with carcasses due to different trophic roles of adults or their immature stages, which can feed on cadaveric tissues (necrophagous) or on other insects in the body, such as larvae of Diptera or other Coleoptera species (necrophilous).

Histeridae comprises 4252 species and 391 genera worldwide, grouped in 11 subfamilies (Mazur 2011), with 139 genera and 1047 species in the Neotropical region (Almeida and Mise 2009). They are mostly predators of soft-bodied insect larvae and eggs, particularly of cyclorraphan Diptera, whose larvae develop on carcasses and dung of large mammals. The odoriferous products of microbial degradation attract both flies and histerids via olfaction (Kovarik and Catherino 2001).

Due to the fact that the Diptera colonize the body from the beginning of the decomposition process (Goff 1993), they have been the group most used for estimating Post Mortem Interval (PMI) within short periods after death. However, they have little usefulness after several weeks or months, when the body is in advanced stages of decomposition. Although histerids are less abundant than flies in the scavenger community, they complete its life cycle in the body (Aballay pers. obs.). Thus, histerids may be helpful when a long time has elapsed since death. Adult histerids reach their highest abundance in intermediate stages of decomposition such as Active and Advanced Decay (Özdemir and Set 2009) and can cause a remarkable decrease in the number of immature stages of Diptera: Calliphoridae (Nuorteva 1970).

Histerid adults have been frequently mentioned in forensic studies on decomposing pig carcasses (Wolff et al. 2001, Centeno et al. 2002, Aballay et al. 2008, Özdemir and Sert 2009, Battán Horenstein and Linhares 2011, Battán Horenstein et al. 2012, Aballay 2012) and on human corpses (Arnaldos et al. 2005, Mariani et al. 2010, Aballay obs. pers.). The correct identification of insects and knowledge of their life history as well as the duration of each stage of development leads to accurately establishing the PMI (Turchetto and Vanin 2004). In South America, the usefulness of histerids as PMI indicators has not been established due to the absence of taxonomic keys that allow their determination, as well as minimal documentation of detailed life histories. In previous forensic studies in the continent, histerids were identified to family level (Mariani et al. 2010), most to generic level (Carvalho et al. 2000, Wolff et al. 2001, Mise et al. 2007, Segura et al. 2009, Battán Horenstein and Linhares 2011, Battán Horenstein et al. 2012) and a few to species level (Centeno et al. 2002, Oliva and Ravioli 2004, Aballay et al. 2008, Mise et al. 2010). A key to the main families of South American Coleoptera of forensic importance was recently published (Almeida and Mise 2009), which includes histerids mentioned for some South American countries. In this key, only six genera and two species were determined for Histeridae, and it does not include most of the species collected in Argentina in decomposition assays (Aballay et al. 2008). For these reasons, it is necessary to have a tool that allows determination of the necrophilous species of Histeridae.

The objective of this paper is to provide an illustrated key to the histerid species associated with decaying carcasses in Argentina to achieve their correct identification. Additionally, diagnoses for these species are presented.

Material and methods

A total of 7070 specimens were collected mostly during forensic studies on decomposing pig carcasses because it is the preferred animal model for forensic entomological studies (Goff 1993). These decomposition experiments were conducted in three Argentinean provinces with arid conditions: Mendoza, San Juan and Catamarca. Histerids were collected during the entire decomposition process on 16 pig carcasses. In Mendoza, the study was carried out at the campus of Instituto Argentino de Investigaciones de las Zonas Áridas, CCT CONICET-Mendoza (32°53'53.3"S, 68°52'26.2"W, 850 m altitude) collecting histerids on 12 pig carcasses, during the four seasons of the year. In San Juan, histerids were collected on two decomposing pig carcasses in summer at the campus of Facultad de Ciencias Exactas Físicas y Naturales, Universidad Nacional de San Juan (31°32'34.1"S, 68°34'38.2"W, 673 m altitude). In Catamarca, histerids were collected on two decomposing pig carcasses during spring in Antofagasta de la Sierra (26°01'32.3"S, 67°20'36.5"W, 3600 m altitude).

In addition specimens from decomposing pig carcasses were recorded in the provinces of Salta (24°54'40"S, 65°28'16"W, 1379 m altitude) and Jujuy (24°09'54.13"S, 65°18'37.73"W, 1383 m altitude), with mesic conditions. For collecting and conserving specimens the methodology followed was that by Centeno et al. (2002) and Aballay et al. (2008, 2012).

Other Histeridae specimens were obtained using three kinds of collecting procedures, the first was conducted on human corpses at different places in Mendoza province authorized by the Medical Forensic Committee of Mendoza; the second was conducted in field trips in different Argentinean provinces on carcasses of cow (Bos taurus), horse (Equus caballus), donkey (Equus asinus), dog(Canis familiaris), snake (not identified), Geoffroy´s cat(Leopardus geoffroyi), llama(Lama glama), guanaco (Lama guanicoe), vicuña (Vicugna vicugna), sheep (Ovis orientalis), fox (Lycalopex griseus), lesser rhea (Pterocnemia pennata), rat (Eligmodontia typus) all found outdoors; the third type of collection was using traps baited with rotting flesh of chicken, squid and sardine in different provinces of Argentina.

Voucher specimens are deposited in the entomological collections of the Instituto Argentino de Investigaciones de las Zonas Áridas (Mendoza, Argentina) and Museo Nacional de Historia Natural (Santiago, Chile).

Specimens were cleaned with water and detergent using a Haier ultrasonic cleaner. Diagnoses were made using a Bausch and Lomb stereomicroscope with magnification between 45× and 60×. Measurements (given in millimeters) were taken with an ocular micrometer. Body length was measured from anterior angle of pronotum to elytral apex, without including head and abdominal terga (propygidium and pygidium) and defined as follows: small 0.5–1.9 mm, medium 2.0–3.9 mm and large 4.0–8.0 mm. Body width was measured at maximum width of elytra, in humeral part. Terminology follows Lackner (2010). The main striae and parts of the body depicted in Figs 1 and 2 were taken from Lackner (2010). Digital photographs of the specimens were taken with a Canon S50 adapted to a Leica MZ6 stereomicroscope. Final images of the specimens (Figs 322) were produced with the image stacking freeware CombineZM (Hadley 2006).

Figure 1.

Saprininae, schematic. Pronotum and elytra, oblique lateral view (taken from Lackner 2010).

Figure 2.

Saprininae, schematic. Habitus, ventral view (taken from Lackner 2010).

Results

The list of histerids of forensic importance in Argentina comprises 16 species distributed in 13 provinces (Table I). In order to enable a more accurate use of the key, diagnosis of each species with habitus photographs are provided.

Key to species of Histeridae associated with carcasses in Argentina
1. Prosternal lobe present (Fig. 3) 2
Prosternal lobe absent (Fig. 4) 4
2. Labrum with setae (Fig. 5) Carcinops (Carcinops) troglodytes (Paykull)
Labrum without setae 3
3 Head prognathous, not retractile; mandibles long, prominent, as long as head; pronotum and elytra lacking punctures; length greater than 6.9 mm (Fig. 6) Hololepta (Leionota) reichii Marseul
Head hypognathous, retractile; mandibles short, as long as half of head; pronotum and elytra with finer and sparse punctation; length less than 1.5 mm (Fig. 7) Phelister rufinotus Marseul
4 Pronotal hypomeron setose in dorsal view 5
Pronotal hypomeron glabrous in dorsal view 10
5 Elytronwith five dorsal striae, the fifth between the fourth dorsal and sutural striae (Fig. 8) Euspilotus (sensu stricto) lacordairei (Marseul)
Elytronwith four dorsal striae, fifth stria absent (Figs 9, 10, 13–15) 6
6 Elytron black, lacking spots (Figs 9, 10) 7
Elytronblack with orange, yellow or white spots (Figs 13–15) 8
7 Outer margin of protibiae with teeth much expanded and 6 denticles (Fig. 11); elytron with coarse and dense punctation, with a shining area with finer and sparse punctation between the fourth dorsal and sutural striae, narrowed apically; length greater than 4.4 mm (Fig. 9) Euspilotus (sensu stricto) patagonicus (Blanchard)
Outer margin of protibiae with teeth moderately expanded and 7-8 denticles (Fig. 12); elytronwith very coarse and dense punctation, with a shining impunctate area between the fourth dorsal and sutural striae, wider apically; length less than 2.9 mm (Fig. 10) Xerosaprinus (Xerosaprinus) diptychus (Marseul)
8 Elytral spot with a digitiform projection towards apex (Fig. 13) Euspilotus (sensu stricto) richteri Lewis
Elytral spot straight on distal edge (Figs 14–15) 9
9 Elytral spot with two digitiform projections anterad, the outer one close to but not reaching the basal edge (Fig. 14) Euspilotus (sensu stricto) lepidus (Erichson)
Elytral spot with three digitiform projections anterad, away from basal edge (Fig. 15) Euspilotus (sensu stricto) ornatus (Blanchard)
10 Anterior half of elytronwith very coarse and dense punctation, with a shining impunctate area between the fourth dorsal and sutural striae (Fig. 16) Euspilotus (Hesperosaprinus) caesopygus (Marseul)
10´ Anterior half of elytronwith finer and sparse punctation, lacking shining impunctate areas (Figs 1722) 11
11 Dorsal elytral striae 3–4 present, well demarcated on anterior half (Figs 1720) 12
11´ Dorsal elytral striae 3 absent or marked as a row of impressed punctures on basal area, stria 4 present or reduced to a rounded arch basally connected to the sutural stria (Figs 21–22) 15
12 Pronotum with a single fovea on each side close to anterior angles or with a longitudinal lateral depression on each side close to lateral margins with coarse and dense punctation (Figs 17–18) 13
12´ Pronotum lacking fovea or longitudinal lateral depression (Figs 19–20) 14
13 Pronotum with a single depression on each side close to anterior angles, with coarse and dense punctation; distal half of elytra, propygidium and pygidium with ocellate punctation, a small puncture within a large puncture (Fig. 17) Euspilotus (Hesperosaprinus) strobeli (Steinheil)
13´ Pronotum with a longitudinal lateral depression on each side, with coarse and dense punctation; distal half of elytra, propygidium and pygidium with regular punctation (Fig. 18) Euspilotus (Hesperosaprinus) pavidus (Erichson)
14 Elytron with inner subhumeral stria; length greater than 2.5 mm (Fig. 19) Euspilotus (Hesperosaprinus) modestus (Erichson)
14´ Elytron lacking inner subhumeral stria; length less than 2.2 mm (Fig. 20) Euspilotus (Hesperosaprinus) parenthesis (Schmidt)
15 Pronotum with marginal stria away from lateral margin; pygidium with a transverse subapical groove not reaching lateral margins (Fig. 21) Euspilotus (Hesperosaprinus) connectens (Paykull)
15´ Pronotum with marginal stria very close to lateral margin; pygidium with a transverse subapical groove reaching lateral margins (Fig. 22) Euspilotus (Hesperosaprinus) azureus (Salberg)
Diagnoses of species
Carcinops (Carcinops) troglodytes (Paykull)

http://species-id.net/wiki/Carcinops_troglodytes

Figures 3, 5
Diagnosis.

Small size (length: 2.1–2.3 mm, width: 1.4–1.6 mm). Body oval, elongated, parallel, black, shiny, with reddish legs. Pronotum with finer and sparse punctation, longer on lateral area, with a large puncture on medial part close to posterior margin. Pronotal hypomeron glabrous in dorsal view. Elytron with finer and sparse punctation in intervals; dorsal elytral striae 1–5 complete, well demarcated with punctures, sutural stria present, reduced on basal part. Pygidium without grooves. Protibiae with teeth expanded and 2 short, separated denticles and a long apical spur; proximal half of outer margin serrate, with small spurs.

Distribution.

Cosmopolitan (Mazur 2011). New record for Argentina.

Figures 3–4.

Prosternum in ventral view. 3 Carcinops (Carcinops) troglodytes 4 Euspilotus (Hesperosaprinus) modestus.

Hololepta (Leionota) reichii Marseul

http://species-id.net/wiki/Hololepta_reichii

Figure 6
Diagnosis.

Large size (length: 6.9 mm, width: 5.3 mm). Body black, shiny, depressed, elongated, parallel, head prognathous, not retractile, mandibles long, prominent, as long as head. Pronotum lacking punctures, with marginal stria well demarcated, in males ending in a fovea on anterior angles. Pronotal hypomeron glabrous in dorsal view. Elytron lacking spot and punctures, with only two dorsal striae, first stria reduced to anterior half, second complete, almost reaching apex. Propygidium larger than pygidium, pygidium without grooves. Protibiae with four teeth, the two distal ones longer.

Distribution.

Argentina, Brazil, French Guiana, Mexico and Central America (Mazur 1984, 2011).

Figures 5–10.

Habitus in dorsal view. 5Carcinops (Carcinops) troglodytes 6 Hololepta (Leionota) reichii. 7 Phelister rufinotus 8 Euspilotus (s. str.) lacordairei 9 Euspilotus (s. str.) patagonicus 10 Xerosaprinus (Xerosaprinus) diptychus. Scale bars: 2 mm. Scale bars: 2 mm.

Phelister rufinotus Marseul

http://species-id.net/wiki/Phelister_rufinotus

Figure 7
Diagnosis.

Small size (length: 1.5 mm, width: 1.3 mm). Body oval, black, shiny, with elytron reddish or black rufescent. Pronotum with finer and sparse punctation, larger on medial part close to posterior margin. Pronotal hypomeron glabrous in dorsal view. Elytron with finer and sparse punctation in intervals; dorsal elytral striae 1–4 complete, fifth present on distal half and with a large basal puncture; sutural stria present on distal half. Pygidium with finer and dense punctation and without grooves. Protibiae with outer margin not expanded and with 7 separated denticles.

Distribution.

Brazil (Mazur 2011). New record for Argentina.

Xerosaprinus (Xerosaprinus) diptychus (Marseul)

http://species-id.net/wiki/Xerosaprinus_diptychus

Figures 10, 12
Diagnosis.

Smallto medium size (length: 1.8–2.9 mm, width: 1.7–2.4 mm). Body oval, black to dark brown, shiny. Pronotum with coarse and dense punctation on anterior, lateral and basal areas, disc small, with finer and sparse punctation. Pronotal hypomeron setose in dorsal view. Elytron with coarse and very dense punctation seemingly rugose on distal half and on proximal half in intervals 1–3, with a smooth, shining area between the fourth dorsal stria, the sutural stria and the rounded arch; elytral dorsal striae 1–4 complete on anterior half, sometimes the first and third vestigial, fourth and sutural striae connected by a rounded arch. Pygidium without grooves. Protibiae with teeth moderately expanded and 7–8 denticles (Fig. 12).

Distribution.

Mexico (Mazur 2011) and Argentina (Aballay et al. 2008, 2012).

Figures 11–12.

Protibia in dorsal view. 11 Euspilotus (s. str.) patagonicus 12 Xerosaprinus (Xerosaprinus) diptychus.

Euspilotus (sensu stricto) lacordairei (Marseul)

http://species-id.net/wiki/Euspilotus_lacordairei

Figure 8
Diagnosis.

Medium to large size (length: 3.4–4.5 mm, width: 3.3–3.8 mm). Body black reddish. Pronotum with coarse and dense punctation on anterior, lateral and basal areas, disc small, with finer and sparse punctation. Pronotal hypomeron setose in dorsal view. Elytron with coarse and dense punctation on distal half, projecting anterad in intervals 1-3, shortest in interval 4; with five dorsal striae well demarcated, 1–4 complete on anterior half, fifth reduced between the fourth dorsal and sutural striae, fourth and sutural striae connected by a rounded arch. Pygidium without grooves. Protibiae with expanded outer margin and 10–11 short, reddish denticles.

Distribution: Argentina, Bolivia and Chile (Mazur 2011; Aballay et al. 2008, 2012).

Euspilotus (sensu stricto) patagonicus (Blanchard)

http://species-id.net/wiki/Euspilotus_patagonicus

Figures 9, 11
Diagnosis.

Large size (length: 4.4–5.8 mm, width: 3.9–4.7 mm). Body black. Pronotum with large, shiny disc, with finer and sparse punctation, lateral and basal areas with coarse and dense punctation, with a punctate depressed area on anterior angles, without punctures behind anterior margin. Pronotal hypomeron setose in dorsal view. Elytronwith coarse and dense punctation on distal half, projecting towards anterior half in intervals 1-4, not reaching inner subhumeral stria, the basal area of fourth and sutural striae, with a shining area with finer and sparse punctation between the fourth dorsal and sutural striae, narrowed apically; elytral dorsal striae 1–4 complete on anterior half, sutural stria sometimes absent in(on) basal part. Pygidium without grooves. Protibiae with teeth much expanded and 5–6 short denticles wider on base (Fig. 11).

Distribution.

Argentina, Bolivia and Chile (Mazur 2011).

Euspilotus (sensu stricto) richteri Lewis

http://species-id.net/wiki/Euspilotus_richteri

Figure 13
Diagnosis.

Medium size (length: 2.3–3.8 mm, width: 2.1–3.4 mm). Body black, elytron with yellow or white spot. Pronotum with finer and sparse punctation, with a longitudinal lateral area on each side with coarse and dense punctation reaching the marginal stria, with two rows of large punctures on base. Pronotal hypomeron setose in dorsal view. Elytron with punctation coarse and dense on posterior half, finer and sparser on anterior half between intervals 2, 3 and 4; elytral dorsal striae 1–4 complete on anterior third, third stria sometimes reduced in basal area, fourth and sutural striae connected by a rounded arch; elytral spot with a digitiform projection towards apex, with two digitiform projections anterad, the outer one between the first and third dorsal elytral striae, the inner one between the fourth dorsal and sutural striae, sometimes between anterior margin, first and fourth dorsal striae and humerus with small yellow spots, making the anterior margin of the large elytral spot fuzzy. Pygidium: female with subapical groove V-shaped, male without grooves. Protibiae with outer margin expanded and 11–12 short, reddish denticles.

Distribution.

Argentina, Chile and Paraguay (Mazur 2011).

Figures 13–18.

Habitus in dorsal view. 13 Euspilotus (s. str.) richteri 14 Euspilotus (s. str.) lepidus 15 Euspilotus (s. str.) ornatus 16 Euspilotus (Hesperosaprinus) caesopygus 17 Euspilotus (Hesperosaprinus) strobeli 18 Euspilotus (Hesperosaprinus) pavidus. Scale bars: 2 mm.

Euspilotus (sensu stricto) lepidus (Erichson)

http://species-id.net/wiki/Euspilotus_lepidus

Figure 14
Diagnosis.

Medium size (length: 2.3–3.3 mm, width: 1.86–2.3 mm). Body black, elytron with yellow or white spot. Pronotum with finer and sparse punctation, with a shining area on disc, with a longitudinal lateral area on each side with coarse and dense punctation, with two rows of large punctures on base. Pronotal hypomeron setose in dorsal view. Elytron with punctation coarse and dense on posterior half, finer and sparser on anterior half defining a shining area between intervals 2, 3 and 4; elytral dorsal striae 1, 2 and 4 complete on anterior half, third stria reduced to a short row of punctures on basal area, fourth and sutural striae connected by a rounded arch; elytral spot with distal margin straight and two digitiform projections anterad, the outer one between the first and second (or third) dorsal striae, the inner one towards the fourth dorsal elytral stria. Pygidium without grooves. Protibiae with outer margin expanded and 10–13 denticles.

Distribution.

Argentina, Bolivia, Chile, and Peru (Mazur 2011).

Euspilotus (sensu stricto) ornatus (Blanchard)

http://species-id.net/wiki/Euspilotus_ornatus

Figure 15
Diagnosis.

Medium size (length: 2.5–3.5 mm, width: 2.3–3.2 mm). Body black, elytron with yellow or orange spot. Pronotum: disc with finer and sparse punctation, lateral areas and base with coarse and dense punctation. Pronotal hypomeron setose in dorsal view. Elytron with punctation coarse and dense on posterior half, finer and sparser on anterior half defining a shining area between intervals 3 and 4; elytral dorsal striae 1–2 and 4 complete on anterior half, third interrupted, fourth and sutural striae connected by a rounded arch; elytral spot occupying the distal half of elytron with distal margin straight and three digitiform projections anterad, the outer one between the first and second dorsal striae, the medial one between the third and fourth dorsal elytral striae, and the inner one close to the sutural elytral stria. Pygidium without grooves. Protibiae with outer margin expanded and 8–10 short, reddish denticles.

Distribution. Argentina and Chile (Mazur 2011; Aballay et al. 2008, 2012).

Euspilotus (Hesperosaprinus) caesopygus (Marseul)

http://species-id.net/wiki/Euspilotus_caesopygus

Figure 16
Diagnosis.

Medium to large size (length: 3.2–4.3 mm, width: 2.7–3.7 mm). Body black. Pronotum with coarse and dense punctation, disc small, with finer and sparse punctation. Pronotal hypomeron glabrous in dorsal view. Elytron with coarse and very dense punctation seemingly rugose, dorsal elytral striae 1–4 absent or vestigial, sutural stria present, lacking rounded arch, with a shining area on anterior half between the fourth dorsal and sutural striae which presents a finer and sparse punctation visible only at 60× magnification. Pygidium: female with subapical groove, male without grooves. Protibiae with outer margin expanded and 10 short, reddish denticles.

Distribution.

Argentina and Bolivia (Mazur 2011).

Euspilotus (Hesperosaprinus) strobeli (Steinheil)

http://species-id.net/wiki/Euspilotus_strobeli

Figure 17
Diagnosis.

Medium to large size (length: 3.5–4.0 mm, width: 2.9–3.9 mm). Body black to metallic blue. Pronotum with a large, shiny disc with finer and sparse punctation, with coarse and dense punctation on lateral area and in a single depression on each side close to anterior angles. Pronotal hypomeron glabrous in dorsal view. Elytron with finer and sparse punctation in the intervals on proximal half; distal half with coarse and dense ocellate punctation, a small puncture within a large puncture, with a smooth, shining area between the fourth dorsal and sutural striae; elytral striae 1-2 almost complete, 3-4 reduced but surpassing the middle of elytron on posterior half, fourth dorsal and sutural striae connected by a rounded arch, lacking inner subhumeral stria. Pygidium with ocellate punctation and with a complete subapical groove in the middle with internal ramifications. Protibiae with outer margin expanded and 7–8 short, reddish denticles.

Distribution.

Argentina and South Brazil (Mazur 2011).

Euspilotus (Hesperosaprinus) pavidus (Erichson)

http://species-id.net/wiki/Euspilotus_pavidus

Figure 18
Diagnosis.

Medium size (length: 2.4–3.8 mm, width: 2.1–3.2 mm). Body black with elytron dark reddish. Pronotum with large, shiny, and smooth disc with finer and sparse punctation; anterior, lateral and basal areas with coarse and dense punctation, with two longitudinal, lateral, depressed punctate areas. Pronotal hypomeron glabrous in dorsal view. Elytron with coarse and dense punctuation on distal third from interval 2 to sutural stria, on proximal half with finer and sparse punctation in intervals 1-4; elytral striae 1-2 almost complete, second longer than first, 3-4 surpassing the middle of elytron on posterior half, with inner subhumeral stria well demarcated, sometimes reduced. Pygidium with punctures, without grooves. Protibiae with outer margin expanded and 7–8 short, reddish denticles.

Distribution.

Argentina, Bolivia, Brazil, French Guiana, Paraguay, Uruguay, Suriname, and Central America (Arriagada 1987; Mazur 2011; Aballay et al. 2008, 2012).

Euspilotus (Hesperosaprinus) modestus (Erichson

http://species-id.net/wiki/Euspilotus_modestus

Figures 4, 19
Diagnosis.

Medium to large size (length: 2.5–4.0 mm, width: 2.4–2.7 mm). Body black to reddish. Pronotum with fine and sparse punctation on disc, larger and deeper on lateral area. Pronotal hypomeron glabrous in dorsal view. Elytron with finer and sparse punctation in the intervals on proximal half; distal half with coarse and dense punctation, apically the punctures form elongate wrinkles; elytral striae 1-4 well demarcated, 1-2 surpassing the middle of elytron on posterior half, 3-4 reduced to anterior half; with inner subhumeral stria well demarcated. Pygidium with coarse and dense punctation, with two short transverse grooves or two longitudinal depressions. Protibiae with outer margin expanded and 8–9 short, reddish denticles, the basal fourth very small.

Distribution.

Argentina, Brazil, French Guiana, Paraguay, Uruguay and Venezuela (Mazur 2011; Aballay et al. 2008, 2012).

Figures 19–22.

Habitus in dorsal view. 19 Euspilotus (Hesperosaprinus) modestus 20 Euspilotus (Hesperosaprinus) parenthesis 21 Euspilotus (Hesperosaprinus) connectens 22 Euspilotus (Hesperosaprinus) azureus. Scale bars: 2 mm.

Euspilotus (Hesperosaprinus) parenthesis (Schmidt)

http://species-id.net/wiki/Euspilotus_parenthesis

Figure 20
Diagnosis.

Smallto medium size (length: 1.7–2.2 mm, width: 1.3–1.8 mm). Body black reddish. Pronotum with fine and sparse punctation over the whole surface area, larger on lateral area. Pronotal hypomeron glabrous in dorsal view. Elytron with coarse and dense punctation on distal half, with finer and sparse punctation in the intervals on proximal half; elytral striae 1-2 almost complete, 3-4 reduced but surpassing the middle of elytron on posterior half; lacking inner subhumeral stria. Pygidium with coarse and dense punctation with or without a short subapical groove, if present it is parenthesis-shaped and concave anterad, not reaching lateral margin of pygidium. Protibiae with outer margin expanded and 7–8 short, reddish denticles.

Distribution. Brazil (Mazur 2011) and Argentina (Aballay et al. 2008, 2012).

Euspilotus (Hesperosaprinus) connectens (Paykull)

http://species-id.net/wiki/Euspilotus_connectens

Figure 21
Diagnosis.

Medium to large size (length: 2.6–3.8 mm, width: 2.2–3.2 mm). Body black. Pronotum with a large, shiny disc with finer and sparse punctation, with coarse and dense punctation on lateral and basal areas and in a single rounded, shallow depression on each side close to anterior angles; with marginal stria away from lateral margin. Pronotal hypomeron glabrous in dorsal view. Elytron with proximal 2/3 lacking punctures, distal third with coarse and dense punctation between the second elytral dorsal and sutural striae; elytral dorsal striae 1–2 almost complete, second larger, third absent or reduced to a short row of punctures on basal area, fourth absent or reduced to a short row of punctures on basal area connected by a rounded arch with sutural stria. Pygidium with punctures and with a transverse subapical groove not reaching lateral margins. Protibiae with outer margin expanded and 7–8 short, reddish denticles.

Distribution.

Argentina, Brazil and Uruguay (Mazur 2011).

Euspilotus (Hesperosaprinus) azureus (Sahlberg)

http://species-id.net/wiki/Euspilotus_azureus

Figure 22
Diagnosis.

Medium to large size (length: 2.9–5.5 mm, width: 2.5–4.7 mm). Body black or metallic blue. Pronotum with a large, shiny disc with finer and dense punctation visible only at 60× magnification, larger on lateral areas and in a single depression on each side close to anterior angles; with marginal stria very close to lateral margin. Pronotal hypomeron glabrous in dorsal view. Elytron with finer and sparse punctation in the intervals on proximal half; distal half with coarse and dense punctation between the second interval and sutural stria; elytral dorsal striae 1–2 almost complete, third absent or reduced to a short row of punctures on basal area, fourth complete on anterior half, fourth and sutural striae connected by a rounded arch. Pygidium with punctures and with a transverse subapical groove reaching lateral margins. Protibiae with outer margin expanded and 7–13 short, reddish denticles, the most basal ones very small.

Distribution.

Argentina, Brazil and Venezuela (Mazur 2011).

Figure 23.

Geographical distribution of sixteen species of Histeridae in Argentina. Provinces: 1 Jujuy: Euspilotus (Hesperosaprinus) caesopygus, Euspilotus (s. str.) lacordairei, Euspilotus (s. str.) lepidus 2 Salta: Euspilotus (Hesperosaprinus) caesopygus, Euspilotus (Hesperosaprinus) strobeli 3 Chaco: Euspilotus (s. str.) lacordairei 4 Catamarca: Euspilotus (Hesperosaprinus) caesopygus, Euspilotus (Hesperosaprinus) pavidus, Euspilotus (s. str.) lacordairei, Euspilotus (s. str.) richteri 5 La Rioja: Euspilotus (Hesperosaprinus) caesopygus, Euspilotus (s. str.) lacordairei, Euspilotus (s. str.) lepidus, Euspilotus (s. str.) richteri 6 San Juan: Euspilotus (Hesperosaprinus) modestus, Euspilotus (Hesperosaprinus) parenthesis, Euspilotus (Hesperosaprinus) pavidus, Euspilotus (s. str.) lacordairei, Euspilotus. (s. str.) ornatus, Xerosaprinus (Xerosaprinus) diptychus 7 Córdoba: Euspilotus (Hesperosaprinus) pavidus 8 Entre Ríos: Euspilotus (Hesperosaprinus) pavidus 9 San Luis: Euspilotus (Hesperosaprinus) caesopygus, Euspilotus (Hesperosaprinus) pavidus, Euspilotus (s. str.) lacordairei, Euspilotus (s. str.) ornatus 10 Mendoza: Carcinops (s. str.) troglodytes, Euspilotus (Hesperosaprinus) azureus, Euspilotus (Hesperosaprinus) caesopygus, Euspilotus (Hesperosaprinus) connectens, Euspilotus (Hesperosaprinus) modestus, Euspilotus (Hesperosaprinus) parenthesis, Euspilotus (Hesperosaprinus) pavidus, Euspilotus (Hesperosaprinus) strobeli, Euspilotus (s. str.) lacordairei, Euspilotus(s. str.) lepidus, Euspilotus (s. str.) ornatus, Euspilotus (s. str.) patagonicus, Euspilotus (s. str.) richteri, Hololepta (Leionota) reichii, Phelister rufinotus, Xerosaprinus diptychus 11 Buenos Aires: Euspilotus (s. str.) patagonicus 12 Neuquén: Euspilotus (s. str.) patagonicus 13 Chubut: Carcinops (s. str.) troglodytes, Euspilotus (Hesperosaprinus) modestus, Euspilotus (s. str.) lacordairei, Euspilotus (s. str.) ornatus, Euspilotus (s. str.) patagonicus, Euspilotus(s. str.) richteri.

Discussion

The 16 Histeridae species collected in this study on carcasses in Argentina are grouped into three of the 11 subfamilies: Saprininae (twelve species of Euspilotus Lewis and one species of Xerosaprinus Wenzel), Histerinae (one species of Hololepta Paykull and one species of Phelister Marseul) and Dendrophilinae (one species of Carcinops Marseul).

Species of Euspilotus, Xerosaprinus and Phelister have been recorded as attracted by carcasses (Kovarik and Catherino 2001), and species of Hololepta and Carcinops are associated with rotting vegetation, especially cacti and bromeliads (Arriagada 1986, Kovarik and Catherino 2001). The species Carcinops (Carcinops) troglodytes has been found to be an effective natural enemy of synanthropic muscoid Diptera: Musca domestica Linnaeus (Muscidae) and Chrysomya putoria (Wiedemann) (Calliphoridae), and considered a potential biological control agent for the coleopteran Alphitobius diaperinus Panzer (Tenebrionidae) that develops in chicken droppings in Brazil (Lopes et al. 2006, Santoro et al. 2010); in Chile it was collected on dry goat’s dung (Arriagada 1986). Hololepta (Leionota) reichii has been considered a predator of larvae and pupae of Melipona compressipes manaosensis Schwarzand Melipona seminigra merrillae Cockerell(Hymenoptera: Apidae)inside bee hives (Coletto-Silva and Freire 2006). In this study, Carcinops troglodytes was collected on pig and Hololepta reichii on human carcasses, in both cases with presence of Calliphoridae larvae.

Nine species of Histeridae constitute new records from the cadaveric fauna in Argentina: Euspilotus caesopygus, Euspilotus connectens, Euspilotus lepidus, Euspilotus richteri, Euspilotus strobelis, Euspilotus azureus, Hololepta reichii, Phelister rufinotus and Carcinops troglodytes. All of them were collected mostly on human and pig carcasses. The remaining seven species associated with carcasses listed in this key were recorded previously for the country in Buenos Aires (Centeno et al. 2002), Neuquén (Oliva and Ravioli 2004) and San Juan provinces (Aballay et al. 2008, 2012). Two species are new records for Argentina: Phelister rufinotus and Carcinops troglodytes.

Histeridae of forensic importance were already cited in the literature, for instance in Central Europe adults of Saprinus planiusculus Motschulsky and Saprinus semistriatus (Scriba) are predictable at a specific time period in the cadaver succession because they have a short period of residency in the carcasses depending on their specialized feeding habits, therefore they are good tools for estimating PMI indicators (Matuszewski et al. 2010).

Operclipygus hospes (Lewis) was recorded from Brazil in buried bodies of rabbits in summer and autumn, and it was suggested that this species plays an important role in forensic entomology as a seasonal indicator (Corrêa et al. 2012).

Further research is necessary to establish the specific time period in the cadaver succession for which the species cited in the present article can be predictable and could be used to estimate PMI indicators based on succession patterns. In addition, immature stages can be useful in forensic entomology because they are reared within the body and collected in advanced stages of decomposition (Aballay pers. obs.) but the duration of larval development is variable and depends on the species (Kovarik and Catherino 2005).

Due to the limited information concerning development of larvae of Histeridae species (Kovarik and Catherino 2005), research studies should be conducted on their life cycle and to this end it is essential to achieve a correct identification of the adult necrophilous histerids. In this sense we consider that the present paper is a basic tool for undertaking these studies.

Table 1.

List of Histeridae species collected on vertebrate carcasses and from baited traps in Argentina and their geographic distribution by provinces. * = baited traps.

Species Substratum /carcasses Province Geographic Coordinates Altitude (m) Collector/ reference
Carcinops (s. str.) troglodytes 4 Pig Mendoza 32°53'49.3"S, 68°52'23.9"W 839 Aballay (2012)
Carcinops (s. str.) troglodytes 4 Sheep Chubut 43°16'18.2"S, 65°26'23.3"W 39 Arriagada G.
Euspilotus (Hesperosaprinus) azureus 192 Pig Mendoza 32°53'58.4"S, 68°52'22.1"W 841 Aballay (2012)
Euspilotus (Hesperosaprinus) caesopygus 5 Pig Mendoza 32°53'53.3"S, 68°52'26.2"W 850 Aballay (2012)
Euspilotus (Hesperosaprinus) caesopygus 2 Human, Mendoza 32°32'07.5"S, 68°58'42.8"W 1424 Aballay F. (forensic cases)
Euspilotus (Hesperosaprinus) caesopygus 3 Pig Jujuy 24°09'54.1"S, 65°18'37.7"W 1383 Quiroga N.
Euspilotus (Hesperosaprinus) caesopygus 1 Pig Salta 24°54'40"S, 65°28'16"W 1379 Ayón R.
Euspilotus (Hesperosaprinus) caesopygus 121 Squid  La Rioja 29°10'45.3"S, 67°37'33.9"W 1806 Arriagada G.
Euspilotus (Hesperosaprinus) caesopygus 4 Squid * Catamarca 27°36'35.4"S, 67°41'48.5"W 1752 Arriagada G.
Euspilotus (Hesperosaprinus) caesopygus 1 Dog San Luis 32°37'37.8"S, 66°54'35.5"W 744 Arriagada G.
Euspilotus (Hesperosaprinus) connectens 12 Pig Mendoza 32°53'57.6"S, 68°52'32.4"W 847 Aballay (2012)
Euspilotus (Hesperosaprinus) modestus 21 Human Mendoza 32°49'18.4"S, 68°52'38.9"W 788 Aballay F. (forensic cases)
Euspilotus (Hesperosaprinus) modestus 4 Cow San Juan 31°59'51.1"S, 68°03'20.3"W 541 Arriagada G
Euspilotus (Hesperosaprinus) modestus 2 Pig San Juan 31°32'34.1"S, 68°34'38.2"W 673 Aballay et al. (2008, 2012)
Euspilotus (Hesperosaprinus) modestus 91 Pig Mendoza 32°53'53.3"S, 68°52'26.2"W 850 Aballay (2012)
Euspilotus (Hesperosaprinus) modestus 91 Sardine* Chubut 43°16'37.1"S, 65°29'49.8"W 68 Arriagada G.
Euspilotus (Hesperosaprinus) parenthesis 2 Pig San Juan 31°32'34.9"S, 68°34'35.9"W 674 Aballay et al. (2008, 2012)
Euspilotus (Hesperosaprinus) parenthesis 30 Pig Mendoza 32°53'49.3"S, 68°52'23.2"W 850 Aballay (2012)
Euspilotus (Hesperosaprinus) pavidus 5 Human Mendoza 32°56'14.2"S, 68°36'32.9"W 653 Aballay F. (forensic cases)
Euspilotus (Hesperosaprinus) pavidus 63 Pig San Juan 31°32'32.1"S, 68°34'44.8"W 675 Aballay et al. (2008, 2012)
Euspilotus (Hesperosaprinus) pavidus 163 Pig Mendoza 32°53'58.4"S, 68°52'22.1"W 841 Aballay (2012)
Euspilotus (Hesperosaprinus) pavidus 70 Donkey Catamarca 26°59'22.1"S, 66°08'42.1"W 2121 Arriagada G.
Euspilotus (Hesperosaprinus) pavidus 15 Horse San Luis 32°38'43.4"S, 66°53'52.7"W 717 Arriagada G.
Euspilotus (Hesperosaprinus) pavidus 45 Chicken * Córdoba 30°44'39.8"S, 64°48'35.5"W 480 Arriagada G.
Euspilotus (Hesperosaprinus) pavidus 100 Cow Entre Rios 32°08'39.1"S, 58°13'04.3"W 31 Arriagada G.
Euspilotus (Hesperosaprinus) strobeli 1 Pig Salta 24°54'40"S, 65°28'16"W 1379 Ayón R.
Euspilotus (Hesperosaprinus) strobeli 1 Cow Mendoza 34°03'18.1"S, 67°49'13.8"W 537 Flores G.
Euspilotus (Hesperosaprinus) strobeli 1 Chicken * Mendoza 34°03'25.1"S, 67°49'11.8"W 534 Arriagada G
Euspilotus (s. str.) lacordairei 25 Pig San Juan 31°32'34.1"S, 68°34'38.2"W, 673 Aballay et al. (2008, 2012)
Euspilotus (s. str.) lacordairei 2 Pig San Juan 30°07'01.1"S, 68°39'43.9"W 1144 Aballay F.
Euspilotus (s. str.) lacordairei 867 Pig Mendoza 32°53'57.6"S, 68°52'32.2"W 850 Aballay (2012)
Euspilotus (s. str.) lacordairei 2 Horse San Luis 32°38'34.4"S, 66°53'35.7"W 720 Arriagada G.
Euspilotus (s. str.) lacordairei 2 Cow San Luis 32°22'08.2"S, 67°09'37.3"W 556 Aballay F.
Euspilotus (s. str.) lacordairei 4080 Sardine*, Squid* Chubut 43°16'37.1"S, 65°29'49.8"W 68 Arriagada G.
Euspilotus (s. str.) lacordairei 20 Rat Chubut 42°24'11.1"S, 63°57'25.4"W 6 Cheli G
Euspilotus (s. str.) lacordairei 2 Lesser rhea Chubut 42°20'21.8"S, 64°49'11.2"W 50 Flores G;.
Euspilotus (s. str.) lacordairei 2 Sheep Chubut 42°20'28.8"S, 64°49'09.2"W 48 Flores G.
Euspilotus (s. str.) lacordairei 5 Vicuña Jujuy 22°44'52.4"S, 65°53'12.9"W 3667 Arriagada G.
Euspilotus (s. str.) lacordairei 166 Squid * La Rioja 29°10'43.5"S, 67°31'49.9"W 1196 Arriagada G.
Euspilotus (s. str.) lacordairei 2 Donkey Catamarca 26°59'22.1"S, 66°08'42.1"W 2121 Arriagada G.
Euspilotus (s. str.) lacordairei 1 Snake Chaco 26°30'16.3"S, 61°11'15.2"W 124 Arriagada G.
Euspilotus (s. str.) lepidus 7 Pig Jujuy 24°09'54.1"S, 65°18'37.7"W 1383 Quiroga N.
Euspilotus (s. str.) lepidus 55 Pig Mendoza 32°53'49.3"S, 68°52'23.9"W 839 Aballay (2012)
Euspilotus (s. str.) lepidus 10 Squid * La Rioja 28°34'17.9"S, 66°47'07.4"W 812 Arriagada G.
Euspilotus (s. str.) ornatus 2 Pig San Juan 31°32'34.9"S, 68°34'35.9"W 674 Aballay et al. (2008, 2012)
Euspilotus (s. str.) ornatus 48 Pig Mendoza 32°53'58.4"S, 68°52'22.1"W 841 Aballay (2012)
Euspilotus (s. str.) ornatus 30 Rat Chubut 45°49'04.7"S, 67°55'59.6"W 680 Cheli G.
Euspilotus (s. str.) ornatus 30 Sardine* Chubut 43°16'30.1"S, 65°29'26.8"W 66 Arriagada G.
Euspilotus (s. str.) ornatus 3 Geoffroy´s cat San Luis 33°08'07.5"S, 66°30'27.9"W 551 Arriagada G.
Euspilotus (s. str.) patagonicus 4 Rat Chubut 42°47'07.5"S, 65°00'43.8"W 9 Cheli G.
Euspilotus (s. str.) patagonicus 4 Guanaco Mendoza 36°03'27.5"S, 68°47'11.1"W 1684 Flores G., Ruiz Manzanos E.
Euspilotus (s. str.) patagonicus 1 Pig Buenos Aires 34°47'13.2"S, 58°26'33.1"W 17 Centeno et al. (2002)
Euspilotus (s. str.) patagonicus 1 Human Neuquén 38°53'54.8"S, 69°56'54.2"W 962 Oliva and Ravioli (2004)
Euspilotus (s. str.) richteri 178 Pig Catamarca 26°01'38.2"S, 67°20'31.6"W 3595 Aballay (2012)
Euspilotus (s. str.) richteri 95 Llama Catamarca 26°01'33.4"S, 67°20'42.5"W, 3585 Aballay (2012)
Euspilotus (s. str.) richteri 85 Squid * Catamarca 27°36'30.1"S, 67°41'04.7"W 1750 Arriagada G.
Euspilotus (s. str.) richteri 8 Pig Mendoza 32°53'57.6"S, 68°52'32.4"W 847 Aballay (2012)
Euspilotus (s. str.) richteri 1 Rat Chubut 42°16'10.4"S, 63°45'32.2"W 40 Cheli G.
Euspilotus (s. str.) richteri 76 Squid * La Rioja 28°34'17.9"S, 66°47'07.4"W 812 Arriagada G.
Hololepta (Leionota) reichii 1 Human Mendoza 32°56'14.2"S, 68°36'32.9"W 653 Aballay F.(forensic cases)
Phelister rufinotus 11 Pig Mendoza 32°53'53.3"S, 68°52'26.2"W 850 Aballay (2012)
Xerosaprinus diptychus 72 Pig San Juan 31°32'32.1"S, 68°34'44.8"W 675 Aballay et al. (2008, 2012)
Xerosaprinus diptychus 2 Horse San Juan 30°13'52.3"S, 67°42'33.8"W 1261 Aballay F.
Xerosaprinus diptychus 2 Fox San Juan 30°19'01.3"S, 68°41'42.3"W 673 Aballay F.
Xerosaprinus diptychus 114 Pig Mendoza 32°53'49.3"S, 68°52'23.9"W 839 Aballay (2012)
Acknowledgements

We acknowledge Hugo San Martino and Eduardo Berté (Medical Forensic Committee of Mendoza) for the collecting permission on human carcasses; Rosana Ayón, Nancy Quiroga and Germán H. Cheli for his collecting efforts in Salta, Jujuy and Chubut provinces; Tomás Lackner for authorizing the utilization of Figs 1 and 2; Nelly Horak for correction of the English language; Mariana Chani Posse for suggestions improving the manuscript; Ana María Scollo for mounting specimens; Germán San Blas and Federico Agrain for help in taking photographs; Rodolfo Carrara for GIS assistance; Michael S. Catherino (Editor Histeridae) and two anonymous reviewers for suggestions for improving this paper. Financial support was provided by grants of the Agencia Nacional de Promoción Científica y Tecnológica (ANPCYT), Argentina (“Estudio de aspectos biológicos de la entomofauna cadavérica aplicados a la entomología forense”, PICT 26000), Universidad Nacional de Quilmes to FHA and Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET, Argentina).

References
Aballay FH (2012) Estudios estacionales en composición, colonización y asociación de la entomofauna cadavérica, en relación a los estadios de descomposición en zonas áridas de llanura y de altura. PhD Thesis, Universidad Nacional de Quilmes, Buenos Aires province.
Aballay FH, Murúa AF, Acosta JC, Centeno N (2008) Primer registro de artropodofauna cadavérica en sustratos humanos y animales en San Juan, Argentina. Revista de la Sociedad Entomológica Argentina 67: 157-163.
Aballay FH, Murúa AF, Acosta JC, Centeno ND (2012) Succession of carrion fauna in the arid region of San Juan province, Argentina: its forensic relevance. Neotropical Entomology 41: 27–31. doi: 10.1007/s13744-011-0005-9
Almeida LM, Mise KM (2009) Revista Brasileira de Entomologia 53 (2): 227-244. doi: 10.1590/S0085-56262009000200006
Arnaldos MI, García MD, Romera E, Presa JJ, Luna A (2005) Estimation of postmortem interval in real cases based on experimentally obtained entomological evidence. Forensic Science International 149: 57-65. doi: 10.1016/j.forsciint.2004.04.087
Arriagada G (1986) Histéridos chilenos (Coleoptera: Histeridae). Primera parte. Revista Chilena de Entomología 14: 71-80.
Arriagada G (1987) Notas sinonímicas y datos distribucionales de Saprininae Neotropicales (Coleoptera: Histeridae). Revista Chilena de Entomología 15: 61-70.
Battán Horenstein M, Linhares AX (2011) Seasonal composition and temporal succession of necrophagous and predator beetles on pig carrion in central Argentina. Medical and Veterinary Entomology 25: 395-401. doi: 10.1111/j.1365-2915.2011.00969.x
Battán Horenstein M, Rosso B, García DM (2012) Seasonal structure and dynamics of sarcosaprophagous fauna on pig carrion in a rural area of Cordoba (Argentina): Their importance in forensic science. Forensic Science International 217: 146-156. doi: 10.1016/j.forsciint.2011.10.043
Benecke M (1998) Six forensic entomology cases: description and commentary. Journal of Forensic Sciences 43: 797-805.
Carvalho LML, Thyssen PJ, Lindares AX, Palhares FAB (2000) A checklist of arthropods associated with pig carrion and human corpses in southeastern Brazil. Memórias do Instituto Oswaldo Cruz 95: 135-138. doi: 10.1590/S0074-02762000000100023
Centeno ND, Maldonado M, Oliva A (2002) Seasonal patterns of arthropods occuring on sheltered and unsheltered pig carcasses in Buenos Aires Province (Argentina). Forensic Science International 126: 63-70. doi: 10.1016/S0379-0738(02)00037-3
Coletto-Silva A, Freire DCB (2006) Hololepta (leinota) reichii Marseul (Coleoptera, Histeridae), un nuevo enemigo natural para la meliponicultura en la Amazonía Central, Brasil. Revista Brasileira de Zoologia 23 (2): 588-591. doi: 10.1590/S0101-81752006000200036
Corrêa RC, Moura DP, Leivas FWT, Almeida LM (2012) Operclipygus hospes (Lewis) (Coleoptera, Histeridae): a beetle of potential forensic importance for buried bodies. Neotropical Entomology 41: 254-256. doi: 10.1007/s13744-012-0032-1
Goff ML (1993) Forensic Science Review 5 (2): 81-94.
Hadley A (2006) CombineZM public domain image processing software. Available on: http://www.hadleyweb.pwp.blueyonder.co.uk/CZM/combinezm.htm [downloaded on 11/June/2009].
Kovarik W, Catherino M (2001) In: Arnett RJ, Thomas MC (Eds.). American Beetles. Volume 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, FL: 213-223.
Kovarik W, Catherino M (2005) In: Beutel RG, Leschen RAB (Eds.). Handbook of Zoology, Coleoptera, Beetles. Volume 1. Morphology and Systematics (Archostemata, Myxophaga, Adephaga, Polyphaga partim). De Gruyter, Berlin: 190-222.
Kulshrestha P, Satpathy DK (2001) Use of beetles in forensic Entomology. Forensic Science International 120: 15-17. doi: 10.1016/S0379-0738(01)00410-8
Lackner T (2010) Review of the Palaearctic genera of Saprininae (Coleoptera: Histeridae). Acta Entomologica Musei Nationalis Pragae 50: 1-154.
Lopes ZWD, Zanetti Lopes WC, Costa FH, Carvalho Balieiro JC, Prado ÂP (2006) Abundância e sazonalidade de histerídeos (Coleoptera) associados ao esterco de granja aviária da Região Nordeste do Estado de São Paulo, Brasil. Revista Brasileira de Entomologia 50 (4): 492-497. doi: 10.1590/S0085-56262006000400009
Mariani R, Varela GL, Demaría M, Rossi Batiz MF (2010) Registro de la artropodofauna cadavérica asociada a restos humanos en situaciones forenses en la provincia de Buenos Aires, República Argentina. Libro de Resumenes XI Congreso Nacional de Criminalística; VII Congreso Internacional de Criminalística y Ciencias Forenses; IX Congreso Iberoamenicano de Criminalística y Ciencias Forenses. Misiones (Argentina), September 2010, 1–7.
Matuszewski S, Bajerlein D, Konwerski S, Szpila K (2010) Insect succession and carrion decomposition in selected forests of Central Europe. Part 2: Composition and residency patterns of carrion fauna. Forensic Science International 195: 42-51. doi: 10.1016/j.forsciint.2009.11.007
Mazur S (1984) A world catalogue of Histeridae. Polskie Pismo Entomologiczne 54: 1-379.
Mazur S (2011) A concise catalogue of the Histeridae (Insecta: Coleoptera). Warzaw University of Life Sciences, SGGW Press, 332 pp.
Mise KM, Massutti L, Moura MO (2007) Levantamento da fauna de Coleoptera que habita a carcaça de Sus scrofa L., em Curitiba, Paraná. Revista Brasileira de Entomologia 51(3): 358–368. doi: 10.1590/S0085-56262007000300014
Mise KM, Barros de Souza AD, Campos CM, Ferreira Keppler RL, Massutti de Almeida L (2010) Coleoptera associated with pig carcass exposed in a forest reserve, Manaus, Amazonas, Brazil. Biota Neotropica 10 (1): 321-324. doi: 10.1590/S1676-06032010000100027
Nuorteva P (1970) Histerid beetles as predator of blowflies (Diptera, Calliphoridae) in Finland. Annales Zoologici Fennici 7: 195-198.
Oliva A, Ravioli JA (2004) Conscripto Carrasco: A peacetime casualty. In: Benecke M (Ed.) Forensic Entomology Special Issue, Anil Aggrawal´s Internet. Journal of Forensic Medicine and Toxicology, 45–49.
Özdemir S, Sert O (2009) Determination of Coleoptera fauna on carcasses in Ankara province, Turkey. Forensic Science International 183: 24-32. doi: 10.1016/j.forsciint.2008.09.018
Santoro PH, Neves PMOJ, Alexandre TM, Gavaguchi SA, Alves LFA (2010) Carcinops troglodytes (Erichson) (Coleoptera: Histeridae) predando larvas de Alphitobius diaperinus (Panzer) (Coleoptera: Tenebrionidae) en aviários. Neotropical Entomology 39 (5): 831-832. doi: 10.1590/S1519-566X2010000500026
Schroeder H, Klotzbach H, Oesterhelweg L, Püschel K (2002) Larder beetles (Coleoptera, Dermestidae) as an accelerating factor for decomposition of a human corpse. Forensic Science International 127: 231-236. doi: 10.1016/S0379-0738(02)00131-7
Segura NA, Usaquén W, Sánchez MC (2009) Succession pattern of cadaverous entomofauna in a semi-rural area of Bogotá, Colombia. Forensic Science International 187: 66-72. doi: 10.1016/j.forsciint.2009.02.018
Turchetto M, Vanin S (2004) Forensic entomology and climatic change. Forensic Science International 146: 207-209. doi: 10.1016/j.forsciint.2004.09.064
Wolff M, Uribe A, Ortiz A, Duque P (2001) A preliminary study of forensic entomology in Medellín, Colombia. Forensic Science International 120: 53-59. doi: 10.1016/S0379-0738(01)00422-4