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ARTHROPOD SUCCESSION ON EXPOSED AND SHADED MAMMALIAN CARCASSES IN NSUKKA, NIGERIA

ONYISHI, Grace Chinenye , OSUALA, Fredrick, AGUZIE, Ifeanyi Oscar , OKWUONU , Elijah Sunday and ORAKWELU, Chinemerem Hodges Department of Zoology and Environmental Biology, U niversity of Nigeria, Nsukka, Enugu State, Nigeria.

Corresponding Author: Onyishi, G. C. Department of Zoology and Environmental Biology, University of Nigeria, Nsukka, Enugu State, Nigeria. Email: grace.ony [email protected] Phone: +234 7064609656

Received February 26, 2020 ; Revised March 08, 2020; Accepted October 26, 2020

ABSTRACT

Ten Wistar rats were used as model to study succession on carcass under shade and exposed conditions . Carcass took longer periods under shade than on the exposed site . Four decomposition stages ( fresh, bloated, decay and dry ) were observed. A total of 164 were c ollected belonging to three classes : Insecta, Arachnida and Crustacean. I nsects formed 95.1 2 %, arachnids 1.22 % and crustaceans 3.66 %. Large numbers of arthropods were found on the exposed carcass than the shaded carcass . C alliphorids and S arcophagids were the primary colonizers observed breeding on the carcasses. The most abundant include Chrysomya albiceps (13.4 %) and serricata (10.3 7 %). Other dipterans including , Phoridae, Sepidae, Fannidae and A nthomyiidae were found during the bloated and decay stages. Famil ies of the coleopterans including; , Silpgidae, Staphylinidae and H isteridae were observed during the bloated to the dry stages and fed on the immature dipterous and remains. Hymenopterans were observed throughout the process of decomposition and played a vital role in carcass decomposition. Other arthropods arrived mostly during the decay stage. Species richness on the carcasses peaked during the decay stage. The rate of decomposition of carcasses is affected by environmental fac tors such as temperature, relative humidity, precipitation and abundance.

Keywords: Post mortem interval, Carcass decomposition, Insect abundance, Forensic

INTRODUCTION According to Bornemissza (1957), the decomposer community of a carcass goes Decomposition is a natural and necessary through a process of ecological succession. The process responsible for the return of organic in sects arrive in a determined sequence, material to the environment through the producing an addition or substitution of species. breakdown of dead plant or matter. It The potential of insects as forensic indicators represents a temporary and changing food has stimulated detailed research regarding the source for varied and distinct community of role of fauna in carcass decomposition in organisms (Putman, 1983). Insects, especially various regions and countries (Tomberlin and coleopteran s and dipteran s are the major Alder, 1998; Centeno et al. , 2002; Kočárek , components of insect community which are 2003 ; Okiwelu et al. , 2008). The basic reason primarily involved in the decomposition process for using insects in criminal investigations ( Kočárek , 2003 ). resides in the fact that insects are usually the

ISSN: 1597 – 3115 ARI 2020 17( 3 ): 3869 – 3877 www.zoo - unn.org Onyishi et al . 3870 first organisms to detect and find a cadaver and later reappear in the course of decomposition are involved in all stages of decomposition. (Wolff et al. , 2001). It is important to observe Furthermore, some insect species are specific that not all the i nvertebrates found near a for certain areas and seasons (Carvalho et al. , corpse are fe eding on it. Based on this, four 2000). Also, it has been o bserved that ecological categories have been recognized for oviposition can occur minutes after death the insect community found around corpses. (Smith, 1986). The knowledge of this succession These include: (i) necrophagous species which pattern as well as their preference for the are flesh feeding species. They constitute the di fferent stages of decomposition, and the m ost important category to establish the time of meteorological data ( temperature and death. (ii) Predators and parasites of humidity), have been applied in necrophagous species , (i ii ) omnivorous species to determine the post mortem interval (PMI) of and the (iv) a dventitious species that use the a corpse ( Kočárek, 2003; Okiwelu et al. , 2008 ) . cadaver like an extension of their ecological This field of involves the niche (Smith, 1986). use of the knowledge of insects in investigation The stud y of the entomofauna of crimes or even ci vil disputes. I t also includes associated with cadavers has been an extremely the application of the study of insects and other effective tool to clarify numerous cases of arthropods to legal i ssues, especially in the homicides, sexual abuses and traffic of organs court of law (Okiwelu et al. , 2008). Some (Catts and Haskell, 1990 ). The types of insect examples of criminal cases where forensic present in a decomposing corpse are specific for entomology was successful in determining the c ertain areas and seasons. Invariably , certain outcome includes, one involving a young groups will colonize first, such as blowflies prostitute and a thirty - year - old member of the ( ) and flesh (S arcophagidae), Army ( Lord and Rodriguez, 1989 ) . Her family but the species involved will usually vary. It is members had informed the police of her therefore necessary , to study each region for vanishing approximately four days before her arthropods associat ed with mammalian body was found. She had last been seen in the carcasses. This is a major factor that has company of this military man on May 31. A hindered the application of forensic entomology number of adult flies and maggots were noticed in Nigeria, as only few studies has been carried and gath ered inside and around the wounds of out in the different geographical location of the the young girl. A number of these maggots country which include works by Okiwelu et al. were kept and grown to be adult flies, and (2 008) in Rivers State, Ekanem and Dike (2010) others were taken in their present state. in Uyo, Ekrakene and Iloba (2009) in Benin City Pictures were taken and enlarged showing the and Abajue et al. (2014) in Okija. Therefore, it flies around the body when it was found. Using is necessary to carry out this study, as this will this evidence along with associating weather add to the available data on entomofauna conditions known to be current around the time, present in mammalian c arcasses in Nigeria. The an entomologist was able to give a PMI . The objective of this study was to investigate insects evidence showed that the flies had arrived on visiting exposed and shaded decomposing rat May 31, which implied a time period of four carcasses to establish relationship between days since she had died. This information insect s and carcasses which may be of forensic indicated the army man to be the last person to importance in Nsukka, Enugu State, Nigeria. be with the girl. When presented with the evidence the authorities were able to get a MATERIALS AND METHODS confession out of the man. The use of forensic entomology was a strong factor in this case S ite D escription : The study was carried out ( Lord and Rodrigue z, 1989 ). within the University of Nigeria, Nsukka Invasion time of the corpse is usually environs. The climate in Nsukka is tropical with considered important since some insect species monthly temperatures ranging between 24 ° C are always present, while others disappear and and 29 ° C (Inyang, 1978 ). There are two

Animal Research International (2020) 17( 3 ): 3869 – 3877 Arthropod succession on exposed and shaded ma mmalian carcasses 3871 seasons: the dry and rainy seasons. The dry RESULTS season is from November to March , while the rainy season is from April to October. Two sites, Phases of Carcass Decomposition : During an open grass area filled with Pennisetum the observation periods, the ambient purpureum (elephant grass), and abandoned temperature for both sites ranges from 25 ° C to building, 50 m apart opposite the maize 28 ° C (26.8 ± 0.53 ° C) on the exposed site and farmland o f the Faculty of Agriculture, University 25 ° C to 29 °C (26.9 ± 1.05 ° C) on the shaded of Nigeria, Nsukka were used . site. Relative humidity ranges from 80 to 100 % (98 ± 5.05) on both sites, and a total of 5550 Study Design : Ten carcasses of laboratory rats mm 2 of precipitation was recorded. ( Rattus norvegicus : Berkenhout 1769) each Decomposition period lasted for 23 days weighing ~ 200 g were used as model . The rats on the exposed carcasses and 25 days on the were sacrificed by strangulation and shaded carcasses. Fou r carrion decomposition immediately transported to the study area. Five phases , namely fresh, bloated, decay and dry carcasses were placed on each site in a cage were observed. The fresh stage lasted for less (120 mm x 100 mm x 30 mm). The cages were than 24 hours on the exposed carcasses, while covered with a 5 cm mesh to allow colonization on the shaded carcasses it was about 30 hours. of carcasses by insects, while preventing Bloating occurred on the second day on both scavengers attack (Sukon tason et al. , 2003 ) . carcasses. It was characterized by the swelling Samples were collected twice a day of the and this lasted for two days on (9:00 and 13:00) during the first (fresh) and the exposed carcasses while on the shaded second (bloated) stages of decomposition and carcasses it lasted for three days. The decay thereafter once a day for up to the end of stage started on the 5 th day, with the breaking - decomposition according to Mabika et al. up of the at the eye and anal region. The (2014). Flying insects were caught using an abdomen later began to tear - up and the i nsect net, while the immature insects and other carcasses were relaxing a strong foul odour. arthropods were collected using forceps. The Lots of maggots activities were observed on arthropods were killed with ethyl acetate and both carcasses, although, the maggots present preserved either dry, or in 70 % alcohol for on the exposed carcasses were greater in subsequent taxonomic determination. Samples number than those of the shaded carcasses. of larvae were reared and id entified (Mabika et The dry stage began when the maggots were al. , 2014). observed moving out from the carcasses and Arthropods were identified to specie s lasted until carrion fauna were no longer found level using keys of Elzinga (2004), Ge nn ard associated with the remains. By the end of the (2007) and Carvalho and Mello - P atiu (2008). experiment, only dry skin, fur an d bones were The ambient air temperature was recor d ed left of the carcasses. using a mercury column thermometer held inside the cage, 5 cm above ground (Feugang et Visiting Arthropod Species Abundance : A al. , 201 2 ). total of 164 arthropods belonging to 3 c lasse s were collected. These include: Insecta, Data analysis : Data was analyzed using Arachnida and Crustaceans. Insects formed Microsoft Office Excel® 2013 (Microsoft 95.12 % of the total arthropods collected , while Incorporation, Redmond, USA). Arthropod t he Crustacea contributed 3.66 % and the relative abundance was computed in Arachnida 1.22 % (Table 1). Eight dipteran percentages as fraction of number of individuals families (Calliphoridae, Sarcophagidae, in a given group divided by total number of Muscidae, Phoridae, Fannidae, , individuals in groups of interest. Sepsidae and Culicidae) were recovered from the exposed carcasses (Table 2), while six di pteran families (Calliphoridae, Sarcophagidae,

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Table 1: Overall arthropod relative abundance on exposed and shaded mammalian carcasses in Nsukka, Nigeria Class Order Family Species Abundance (%) Insecta Diptera Calliphoridae Chrysomya albiceps 22(13 .4) Chrysomya megalocephala 9(5.49) Chrysomya chloropyga 7(4.27) Lucilia serricata 17(10.37) Lucilia illustris 7(4.27) Phormia regina 8(4.88) 70(42.68) Muscidae Musca domestica 10(6.10) stabulans 4(2.44) 14(8.54 ) Sarcophagidae Sarcophaga spp. 6(3.66) Anthomyiidae Anthomyia spp. 1(0.61) Fannidae Fannia spp. 5(3.05) Sepsidae Sepsis spp. 15(9.15) Phoridae Megaselia scalaris 3(1.83) Culicidae Culex pipiens 1(0.61) 115 (70.13) Coleoptera Histe ridae Hister spp. 3(1.83) Hypocalus spp. 1(0.61) Silphidae Nicrophoru spp. 3(1.83) Staphylinidae Ocypus spp. 2(1.22) Dermestidae lardarius 1(0.61) 10(6.1) Hymenoptera Formicidae Messor spp. 10(6.10) Camonutus spp. 10(6.10) Pheidole spp. 5(3.05) 25(15.24) Orthoptera Acrididae Pododula ancisa 4(2.44) Gryllidae Gymnogrylus lucens 2(1.22) 6 (3.66) Total 156(95.12) Arachni d a Araneae Araneidae Aranea spp. 2(1.22) Crustacean Isopoda Armadillididae Armadill idium spp. 6(3.66) 164(100)

Table 2: Arthropod relative abundance on exposed mammalian carcasses in Nsukka, Nigeria Class Order Family Species Abundance (%) Insecta Diptera Calliphoridae Chrysomya albiceps 12(12.37) Chrysomya megalocephala 6(6. 19) Chrysomya chloropyga 5(5.15) Lucilia serricata 11(11.34) Lucilia illustris 4(4.12) Phormia regina 8(8.25) 46(47.43) Muscidae Musca domestica 7(7.22) 2(2.06) 9(9.28)

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Sarcophagidae Sarcophaga spp. 3( 3.09) Anthomyiidae Anthomyia spp. 1(1.03) Fannidae Fannia spp. 3(3.09) Sepsidae Sepsis spp. 9(9.28) Phoridae Megaselia scalaris 2(2.06) Culicidae Culex pipiens 1(1.03) 19(19.59) Coleoptera Histeridae Hister spp. 2(2.06) Silphidae Nicrophoru spp. 1(1.03) Staphylinidae Ocypus spp. 1(1.03) Dermestidae Dermestes lardarius 1(1.03) 5(5.15) Hymenoptera Formicidae Messor spp. 5(5.15) Camonutus spp. 5(5.15) 10(10.31) Orthoptera Acrididae Pododula ancisa 2(2.06) Gr yllidae Gymnogrylus lucens 1(1.03) 3(3.09) 92(94.85) Arachnida Araneae Araneidae Aranea spp. 2(2.06) Crustacean Isopoda Armadillididae Armadillidium spp. 3(3.09) Total 97(100)

Muscidae, Phoridae, Fannidae and Sepsidae) (Culicidae) were observed on the exposed were recovered fro m the shaded carcasses carcasses during the dry stage. Immature of (Table 3). Calliphorids were observed on both carcasses during the decay stage. Arthropod Succession on Carcasses : During the study, four families of Primary colonizers on the exposed carcasses (Coleoptera) were ob served. These were during the fresh stage were the Calliphorids and Dermestidae, Silphidae, Staphylinidae and Sarcophagids (Figure 1) , while on the shaded Histeridae. Dermestes lardarius (Dermestidae) carcasses only the Calliphorids were observed was the first to arrive at the exposed carcasses fr om the fresh stage up to the decay stage on the 5 th day (decay stage), while on the (Figure 2) . Adult Sarcophagids were obs erved shaded carcasses, Nicrophoru species (Silphidae) from the fresh stage up to the decay stage on was the first to arrive on the 7 th day (decay the exposed carcasses, while from the shaded stage). Dermestids were absent on the shaded carcasses it was observed from the bloated carcasses while Silphids and Staphylinids were stage to the decay stage. found on both carcasses during the decay and The family Muscidae and Sepsidae were dry stages. Histerids were only found during the observed during the bloated and decay stages dry stage on both carcasses. on both carcasses. Adult Fannidae, Anthomyiidae An ts (Hymenoptera) were actually the and Phoridae were observed during the decay first group of insects to have invaded both stage on the exposed carcasses. This was carcasses. They were found in less than one similar to those found on the shaded carcasses hour on the first day of the research. Three exc ept for Anthomyia sp ecies which were species of ants were observed, namely, Messor absent. The sepsid flies were also observed on sp p. , Camonutus sp p. and Pheidole sp p . The y both carcasses during the dry stage. The occurred throughout the process of Calliphorids were the highest number of flies decomposition. from both the exposed and the shaded Other arthropods observed on bo th carcasses (42.68 %), followed by the sepsids carcasses include; Aranea sp . (spiders) and (9. 15 %) and muscids (8.54 %). Mosquitoes terrestrial crustaceans ( Armadillidium sp . ).

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Table 3: Insect species relative abundance on shaded mammalian carcasses in Ns ukka, Nigeria Class Order Family Species Abundance (%) Insecta Diptera Calliphoridae Chrysomya albiceps 10(14.93) Chrysomya megalocephala 3(4.48) Chrysomya chloropyga 2(2.99) Lucilia serricata 6(8.96) Lucilia illustris 3(4.48) 24(35.8 2) Muscidae Musca domestica 3(4.48) Muscina stabulans 2(2.99) 5(7.46) Sarcophagidae Sarcophaga spp. 3(4.48) Fannidae Fannia spp. 2(2.99) Sepsidae Sepsis spp. 6(8.96) Phoridae Megaselia scalaris 1(1.49)

Coleoptera Histeridae Hister spp. 1(1.49) Hypocalus spp. 1(1.49) Silphidae Nicrophoru spp. 2(2.99) Staphylinidae Ocypus spp. 1(1.49) 5 (7.46) Hymenoptera Formicidae Messor spp. 5(7.46) Camonutus spp. 5(7.46) Pheidole spp. 5(7.46) 15(22.39) Insecta Orthoptera Acrididae Pododula ancisa 2(2.99) Gryllidae Gymnogrylus lucens 1(1.49) 3(4.48) Crustacean Isopoda Armadillididae Armadillidium spp. 3(4.48) 67(100)

The Arachnids were found during the fresh and different from the present observations only bloated stages while the crustace ans were based on nomenclature; their observed stages observed during the decay up to the dry stages could be categorized in to the four stages fresh, on both carcasses. Orthoptera (grasshoppers bloated, decay and dry. and crickets) were also observed on both Colonization species of greatest carcasses. imp ortance in the early stages of decomposition usually are those from the three dipteran DISCUSSION families: Calliphoridae, Sarcophagidae and Muscidae ( Carvalho and Mello - Patiu, 2008 ). This The sequence of the different stages observed was confirmed in the study as the calliphorids during the study (fresh, bloated, decay and dry) and sarcophagids were the first to arrive on was similar to that report ed by previous studies both exposed and shaded carcasses. However, ( Kočárek , 2003 , Ekanem and Dike, 2010; the muscids first appeared on the third day Mabika et al. , 2014). However, Okiwelu et al. (bloated stage). (20 08 ) observed fresh, bloated, wet, dry and Generally the sequence and duration of skeletonized , while Feugang et al. (2012) noted insect succession on the exposed and shaded five stages of decomposition; fresh, bloated, sites followed the same general pattern active deca y, advanced decay and dry. The previously rep orted ( Centeno et al. , 2002; observations made by the two later studies are Ekanem and Dike , 2010; Okiwelu et al. , 20 08 ;

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ARTHROPODA Stages and Days Fresh Bloated Decay Dry 1 2 3 - 4 5 - 6 7 - 12 13 - 15 16 - 20 21 - 30 Diptera Calliphoridae Sarcophagidae Muscidae Fannidae Anthomyiidae Sepsidae Phoridae Culicidae Coleoptera Dermestidae Staphtlinidae Silphidae Histeridae Hymenoptera Formicidae Orthoptera Acrididae Gryllidae Arachnida Aranaea Crustacea Isopoda

Key: Fresh Bloated Decay Dry Figure 1: Arthropod succession on exposed mammalian carcasses in Nsukka, Nigeria

ART HROPODA Stages and Days Fresh Bloated Decay Dry 1 - 2 2 - 3 3 - 4 5 - 8 9 - 12 13 - 15 16 - 20 21 - 30 Diptera Calliphoridae Sarcophagidae Muscidae Fannidae S epsidae Phoridae Coleoptera Staphtlinidae Silphidae Histeridae Hymenoptera Formicidae Or thoptera Acrididae Gryllidae Crustacea Isopoda

Key: Fresh Bloated Decay Dry Fi gure 2: Arthropod succession on shaded mammalian carcasses in Nsuk ka, Nigeria

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Mabika et al., 2014). E xposed carcasses therefore the spiders probably did not prey on decomposed faster than the shaded carcasses. any adult flies. This was as a result of the large number of insects that visited the exposed carcasses. Conclusion: The rate of decomposition of Maggots were observed on the fifth day of the carcasses is affected by environmental factors study, was in contrast with the study of Mabika such as temperature, relative humidity, et al. (2014), where maggots were observed on precipitation and insect abundance. Four distinct the fourth day on rabbit carcasses. Feugang et stages off carcass decomposition were observed al. (2012) also observed maggots from rat during the study. Succession of arthropods was carcasses on the second day of decomposition. attracted on the decomposing carcasses, These differences could be attributed to the size particular ly insects. Their breeding biology led to of the carcasses, vegetation and weather the breakdown and exposure of the carcasses condition of the area (Grassberger and Frank, soft tissues. Of the insect species collected 2004). migration from the carcasses during the study, Chrysomya albiceps and was observed on day 9, and th is observation Lucilia serricata could be important for further was almost similar to th at of Mabika et al. forensic studies since they were m ore numerous (20 14 ) who observed maggot migration o n the and observed breeding on the carcasses. 8 th day. The earliest coleopterans which were ACKNOWLEDGEMENTS members of Dermestid ae , Staphylind ae and Silphid ae were observed during the decay stage The authors are grateful to the Department of of the study , and correspond ed to that of Zoology and Environmental Biology, University Centeno et al. (2002) , but differ ed from that of of Nigeria, Nsukka, Enugu State, Nigeria for Ekanem and Dike (2010), w ho observed provision of the equipment used for conduct of members of the Staphylind ae , Scarabid ae , the study. Clerid ae and Chrysomelid ae during the bloated stage. Th ese difference s may probably be due REFERENCES to the differences in vegetation of the a r e a s . The beetles (Coleoptera) were observed ABAJUE, M. C., EWUIM, S. C. and AKUNNE, C. consuming soft tissues of the decomposing E. (2014). Preliminary checklist of flies carcasses, as well as preyed on the abundant associated with pig carrion decomposition dipterous larvae present on the carcasses. The in Okija, Anambra State, Nigeria. Animal late coleopterans arrivals e.g. the Histerid ae fed Research International , 11(1): 1899 – on the dry carcass tissues. 1904. The ants (F ormicidae) played a major BORNEMISSZA, G. F. (1957). An analysis of role in decomposition of the car casses, since arthropod succession in carrion and the they were present in large numbers throughout effect of its decomposition on the soil the period of the experiment. Their invasion, fauna. Australian Journal of Zoology , especially as they came in colonies, may have 5 (1): 1 – 12. affected the carcasses decomposition rate, CARVALHO, C. J. B. D. and MELLO - PATIU, C. A. either by predation on immature stages of D. (2008). Key to the adults of the most dipterous flies, or by removal of portions of the common forensic species of Diptera in carcasses (Ekrakene and Iloba, 2010). Several South America. Revista Brasileira de other arthropods were observed on the Entomologia , 52 (3): 390 – 406. carcasses, these include; crickets, grasshoppers, CARVALHO, L. M. L. D., THYSSEN, P. J., spiders, and terrestrial crustaceans. These LINHARES, A. X. and PALHARES, F. A. formed part of the local entomofauna. For B. (2000). A checklist of arthropods insta nce, the arachnids in this study were only associated with pig carrion and human encountered at the exposed sites. Cobwebs corpses in Sout heastern . Memórias were not observed around the enclosure;

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