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This Article Appeared in a Journal Published by Elsevier. the Attached This article appeared in a journal published by Elsevier. The attached copy is furnished to the author for internal non-commercial research and education use, including for instruction at the authors institution and sharing with colleagues. Other uses, including reproduction and distribution, or selling or licensing copies, or posting to personal, institutional or third party websites are prohibited. In most cases authors are permitted to post their version of the article (e.g. in Word or Tex form) to their personal website or institutional repository. Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/authorsrights Author's personal copy Revista Brasileira de Entomologia 59 (2015) 132–137 REVISTA BRASILEIRA DE Entomologia A Journal on Insect Diversity and Evolution w ww.rbentomologia.com Biology, Ecology and Diversity Trophic roles of scavenger beetles in relation to decomposition stages and seasons a,b b a,∗ Noelia I. Zanetti , Elena C. Visciarelli , Néstor D. Centeno a Laboratorio de Entomología Aplicada y Forense, Departamento de Ciencia y Técnica, Universidad Nacional de Quilmes, Buenos Aires, Argentina b Cátedra de Parasitología Clínica, Departamento de Biología, Bioquímica y Farmacia, Universidad Nacional del Sur, Buenos Aires, Argentina a r a b s t r a c t t i c l e i n f o Article history: Carcasses represent a trophic and reproductive resource or shelter for arthropods, which are a repre- Received 29 November 2014 sentative component of the decomposition process. Four experiments, one per season, were conducted Accepted 9 February 2015 in a semi-rural area of Bahía Blanca, Argentina, to study the trophic roles of cadaveric beetles, evaluat- Available online 8 April 2015 ing the abundance, composition and dominance during all decomposition stages and seasons. Species Associate Editor: Rodrigo Kruger of necrophagous, necrophilous and omnivorous habits were found. Abundance, composition and domi- nance of beetles in relation to their trophic roles changed according to seasons and decomposition stages. Keywords: Guilds and patterns of succession were established in relation to those periods. Trophic roles could be an Carrion fauna indicator of beetle associations with decomposition stages and seasons. Ecological roles Forensic entomology © 2015 Sociedade Brasileira de Entomologia. Published by Elsevier Editora Ltda. All rights reserved. Insect succession Introduction amongst others (Greenberg, 1971; Catts and Goff, 1992; Goff, 1992; Kócarek, 2001; Centeno et al., 2002; Forbes et al., 2005; Faria et al., Carcasses represent a trophic and reproductive resource or 2007; Battán Horenstein and Linhares, 2011; Dekeirsschieter et al., shelter for a variety of animals such as arthropods, which are a 2011; Matuszewski et al., 2014). Thus, the study of these vari- representative component of the decomposition process. Leclercq ables is important for the estimation of postmortem interval (PMI) (1974) proposed an ecological classification based on the trophic and other applications in forensics (Keh, 1985; Anderson, 2001; roles of the cadaveric fauna: necrophages (feed from corpses), Centeno et al., 2002; Dourel et al., 2010; Battán Horenstein and necrophiles (predate on or parasitize other arthropods), omni- Linhares, 2011; Ortloff et al., 2012; Pers. comm., Zanetti et al., vores (feed from corpses and predate on other arthropods) and 2015). incidentals (look for refuge or appear at random). The car- Beetles are especially useful in forensic cases in which Diptera rion arthropod community is mainly represented by insects have not been sampled or when they have already left the body (Carvalho et al., 2000). These are capable of detecting chemical and only beetles can be found (Centeno et al., 2002). But also, changes, so their appearance in one particular stage of decom- some species of Coleoptera have been observed in earlier stages position is not likely to be accidental (Magana,˜ 2001; Archer and of decomposition and they can appear in different waves in a body, Elgar, 2003; Dekeirsschieter et al., 2011; Von Hoermann et al., thus, further studies on this order need to be conducted. 2011, 2012). Segura et al. (2011) suggested that decomposition The aim of this study was to analyze the abundance, composition stages are a primary factor in determining the insect commu- and dominance of cadaveric beetles per season and decomposition nity, because the species exhibit adaptations that allow them to stage in accordance with their trophic roles (guilds). exploit and use corpses under different conditions and periods of time. Materials and methods The decomposition process and the cadaveric fauna succes- sion can be influenced by circumstances that cadavers go through This study is a part of a greater study of decomposition after death and many variables have to be considered in each and cadaveric succession (Pers. comm., Zanetti et al., 2015). The situation: climatic conditions, season, geographical region, sun experiments were carried out in a field property of the Uni- ◦ exposure, synanthropy, latitude, substrate type, larval migration, versidad Nacional del Sur, located at Bahía Blanca (38 41 41 S, ◦ 62 15 10 W), Buenos Aires province, Argentina. Twelve domestic pigs (Sus scrofa L.) weighing 15–16 kg were ∗ used. Three of them were killed each season by a stab to the heart Corresponding author. E-mail: [email protected] (N.D. Centeno). (the usual procedure of commercial butchers) 1 h before exposure, http://dx.doi.org/10.1016/j.rbe.2015.03.009 0085-5626/© 2015 Sociedade Brasileira de Entomologia. Published by Elsevier Editora Ltda. All rights reserved. Author's personal copy N.I. Zanetti et al. / Revista Brasileira de Entomologia 59 (2015) 132–137 133 Table 2 and were kept inside a plastic bag to keep away insects until the Species/morphospecies of necrophagous habits. beginning of the experiment. The Ethical Commission of the Uni- versidad Nacional del Sur approved this procedure. Each carcass Families Species/morphospecies was laid inside a cage to protect it from scavenger vertebrates, and Dermestidae Dermestes maculatus (De Geer, 1774) under direct sun along transects separated 100 m from each other. Tenebrionidae Scotobius clathratus (Guérin-Méneville, 1834) To complement the sampling, six pitfall traps were placed around Scotobius miliaris (Billberg, 1815) Scotobius muricatus (Guérin-Méneville, 1834) each cage at a distance of 50 cm from the carcass (experimental Scarabaeidae Aphodius militaris (LeConte, 1858) traps); two per each long side and one per each wide side. For con- Aphodius pseudolividus (Balthasar, 1941) trol purposes, other six pitfall traps with the same spatial pattern Ataenius platensis (Blanchard, 1844) were placed at 15–30 m from the last cage. Canthon ornatus bipunctatus (Burmeister, 1873) Degallieridium sp. Literature typically refers to decomposition as a process of Pseudocanthon sp. five stages that are not necessarily discrete (Bornemissza, 1957; Onthophagus sp. Schoendly and Reid, 1987; Moura et al., 2005) but which can Trogidae Omorgus batesi (Harold, 1872) be used as a guide or reference points. We followed the crite- Polynoncus gemmingeri (Harold, 1872) rion established by Centeno et al. (2002) to define the stages of decomposition. Carcasses were visited daily until concluding the experiment (several days after the carcass reached the remains Table 3 Species/morphospecies of omnivorous habits. stage). Beetles from on, under and inside the corpses were col- lected manually with entomological forceps. Beetles were taken Families Species/morphospecies from cadavers (per day) only if 10% of the individuals (this percent- Anthicidae Anthicidae sp. age was considered for each family) were available in order to avoid Cleridae Necrobia rufipes (De Geer, 1775) any alteration in the decomposition and/or succession processes Nitidulidae Nitidula carnaria (Schaller, 1783) due to loss of insects, and preserved into plastic containers with ethanol 70%. Some larvae were reared for identification of the spec- imens and cultures with adults were established for future research Abundance of guilds (Pers. comm., Zanetti et al., 2015). Taxonomical determination or corroboration of some species/morphospecies was performed by Necrophiles some specialists. The specimens were deposited dry-mounted or It was the most abundant group. More than 38 species were in alcohol at Cátedra de Zoología de Invertebrados II, Departa- observed. In cadavers, in order of frequency and abundance, mento de Biología, Bioquímica y Farmacia, Universidad Nacional they were Carpelimus spp., Aleocharinae spp., Philonthus flavolim- del Sur. batus (Erichson, 1753), Creophilus maxillosus (Linnaeus, 1758) Species and morphospecies were grouped in accordance with (Staphylinidae); Euspilotus (Hesperosaprinus) pavidus (Erichson, their trophic roles. Firstly, we evaluated the composition and abun- 1834), Phelister sp., Euspilotus (sensu stricto) ornatus (Blanchard, dance of each group, calculating the mean and SE of each one per 1843), Euspilotus (sensu stricto) lacordairei (Marseul, 1855) and sampling unit (observations and/or capture on carcass or capture Euspilotus (sensu stricto) patagonicus (Blanchard, 1843) (His- in six pit fall traps) and day, during each season and decomposition teridae); while in traps, Hylitus tentyroides (Lacordaire, 1830) stage. Secondly, we analyzed the dominance of each group. (Tenebrionidae) and the tribes Harpalini, Cyclosomini,
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