International 166 (2007) 182–189 www.elsevier.com/locate/forsciint

Succession pattern of -feeding in Paramo, Colombia Efrain Martinez, Patricia Duque, Marta Wolff * Grupo interdisciplinario de Estudios Moleculares (GIEM). Universidad de Antioquia. AA, 1226 Medellı´n, Colombia Received 8 April 2004; accepted 10 May 2006 Available online 21 June 2006

Abstract The minimum postmortem interval can be estimated based on knowledge of the pattern of succession on a corpse. To use this approach requires that we take into account the rates of insect development associated with particular climatological conditions of the region. This study is the first to look at insect succession on decomposing carcasses in the high altitude plains (Paramo) in Colombia, at 3035 m above sea level. Five stages of were designated with indicator species identified for each stage: Callı´phora nigribasis at the fresh stage; Compsomyiops verena at the bloated stage; Compsomyiops boliviana during active decay; Stearibia nigriceps and Hydrotaea sp. during advanced decay and Leptocera sp. for dry remains. A succession table is presented for carrion-associated species of the region, which can be used for estimating time since death in similar areas. Compsomyiops boliviana is reported for the first time in Colombia. # 2006 Elsevier Ireland Ltd. All rights reserved.

Keywords: ; Paramo; Insect succession; Neotropics

1. Introduction change drastically over short distances. In any study of this type we would expect to find a similar process of insects being Forensic entomology is a frequently used tool to estimate the involved in the recycling of , but the occurrence of the time interval between death and the discovery of the body. This species involved in this process should vary throughout period is known as the postmortem interval or PMI. For different climactic regions. To date most of the neotropical intervals greater than 72 h, forensic entomology can be more studies have been done in warm tropical conditions, and none accurate in determining PMI than traditional techniques, and has looked at the process at high altitudes. sometimes is the only available method [1]. This study addresses our lack of knowledge in this area as The carrion-feeding invertebrate fauna is principally made part of a larger ongoing effort to identify the carrion-feeding up of insects and the estimation of PMI using the succession of entomofauna and associated insect succession patterns at insects on cadavers and age necrophagous larvae has been used altitudes ranging from 0 to >3000 m above sea level. extensively [2–8]. The first records of applied forensic entomology occurred in China in the 13th Century [9]. 2. Materials and methods Forensic entomology was used for the first time as a legal instrument in a court of law in France in 1850 [10]. Since then The study was carried out in the Paramo region of Chingaza the number of studies carried out in this field has increased National Park, located in the Eastern range of the Colombian significantly, particularly in , United States, Canada and Andes, (738300 to 738550W; 48200 to 48500N) at an altitude of in some parts of the subtropics [1,2,10,11–16]. 3035 m above sea level. Average annual temperatures fluctuate In the Neotropics, relatively few studies have been carried between 4.5 and 21.4 8C and the annual relative humidity is out [17,18,3,4,19,6,20–22]. In countries such as Colombia, above 80% [23]. The landscape of the Paramo is characterized by environmental conditions and regional climatic conditions can the presence ‘‘ fraylejones’’ (Espeletia spp.), ‘‘chuscales’’ (Chusquea tessellate) and ’’pajonales (Calamagrostis spp.) [24]. Three pigs (Sus scrofa L.) were used as models. Each pig * Corresponding author at: Instituto de Biologı´a, Universidad de Antioquia, weighed approximately 10 kg and was killed by cardiac puncture A.A. 1226, Medellı´n, Colombia. Tel.: +57 4 210 5662; fax: +57 4 233 01 20. on September 12, 2002. Immediately after death, the pigs were E-mail address: [email protected] (M. Wolff). placed in individual metal cages (60 cm 40 cm 40 cm)

0379-0738/$ – see front matter # 2006 Elsevier Ireland Ltd. All rights reserved. doi:10.1016/j.forsciint.2006.05.027 E. Martinez et al. / Forensic Science International 166 (2007) 182–189 183

Table 1 Succession of insects associated with exposed carcasses on Paramo. Ecological Order Family Species Fresh Bloated Active Advanced Remains category ALEALEALEALPALP Necrophagous Diptera Calliphora xxxxxxxxxxxxxxx nigribasis Compsomyiops xxxxxx boliviana Sarconesiopsis xx magellanica Compsomyiops x xxxxxxx xxxxx verena Sarcophagidae Helicobia sp. x x Microcerella sp x Unident x Predators Diptera Unident x x x x Coleoptera Histeridae Neopachylopus sp. x x x x Carabidae Carabus sp. x x x x Staphylinidae Lathropinus sp. x x x x Dianous sp.xxxxx Anacyptus sp. x Stenus sp. x Melyridae Dasyrhadus sp. x x x x Dytisidae Copelatus sp. x x Predators/ Hymenoptera Sphecidae Unident x x x x parasitois Ichneumonidae Unident x x x Braconidae Unident x Encyrtidae Unident x Chalcididae Unident x Predators Dermaptera Forficulidae Unident x x x x Hemiptera Enicocephalidae Unident x Reduviidae Unident x x x Saprophagous Diptera Stearibia xx xx nigriceps Azelia sp. xxxx (probably) Dasymorellia xxxx seguyi sp. x x Helina sp. x Hydrotaea sp. x x x x x Limnophora sp.xxxxx Syllimnophora xxxx atroviatta Muscina x x xx xx stabulans Leptocera sp.xxxxxx Beckerina sp. x x x Dohrniphora sp. x x Metopina sp. x x x Diploneura sp. x Borophaga sp. x Unident x x Orygma sp. x x Drosophila sp. x x Speolepta sp. x Coleoptera Ptillidae Actinopteryx sp.xxxxx Omnivorous Hymenoptera Vespidae Unident x x x Dyctioptera Blattellidae Unident x x Incidental Diptera Ischnomyia sp. x x Unident x x Unident x Unident x x 184 E. Martinez et al. / Forensic Science International 166 (2007) 182–189 Table 1 (Continued ) Ecological Order Family Species Fresh Bloated Active Advanced Remains category ALEALEALEALPALP Meromyza sp. x x x Dolycopodidae Unident x x Otitidae Cephalia sp. x x Psycodidae Psychoda sp. x x Unident x x x Sciomycidae Unident x Syrphidae Eristalis sp.xxxx Tabanidae Tabanus sp. x x x Coleoptera Cantaridae Cantharis sp. x Chrysomelidae Unident xxxxx Coccinelidae Unident x Curculionidae Unident x Lampyridae Pollaclasis sp. x x Photuris sp. x x Lycidae Unident x Melolontidae Unident x Nitidulidae Conotelus sp. x x Oedemeridae Unident x Hemiptera Cercopidae Unident x Cicadelidae Unident xxxxx Lygaeidae Unident x x Miridae Unident x x x Pentatomidae Unident x Pyrrhocoridae Unident x A: Adult; L: Larvae; E: Eggs; P: Pupe. made with 2 cm 2 cm wire meshing. This allowed access to The taxonomic identification of the adults and larvae was the carcasses by insects while preventing disturbance by carried out using the following keys; Greenberg and Szyska vertebrate scavengers [6]. The three cages were placed 50 m [18,21], Borror et al. [27], Carvalho [28], Dear [29], Liu and from each other. Greenberg [30], Mariluis [31], Mc Alpine et al. [32,33], The cadavers were observed for 8 h immediately following Queiroz and Carvalho [34], Smith [2,35], Stehr [36], Wells sacrifice. Subsequently, the process of decomposition was et al. [37] and White [38]. When necessary, the larvae were observed during daylight hours from September 12 to December cleared with KOH and permanently mounted in Canada balsam. 4 2002. Samples were taken three times a day (7.00, 13.00 and All the specimens were deposited in the Universidad de 19.00) for the first eight days; twice a day (at 11.00 and 17.00) for Antioquia Entomological Collection. the next eight days and subsequently once a day (at 12.00) until the dry remains stage had been reached 83 days after death. At 3. Statistical analysis each sampling time, flying over or perched on the carcass were collected with an entomological net before the This study was designed to evaluate the succession pattern of carcass was moved. Then, insects in natural cavities (eyes, nose, insects during the decomposition of the three cadavers, and was mouth, anus) and the cardiac puncture wound were collected. not designed for a quantitative analysis. However, insect Finally, three times a day the insects were collected underneath species were assigned a number and the presence or absence of the carcass and in the soil to a depth of 10 cm. Immature each species on each on each day was recorded and the specimens were fixed in 80% alcohol and the adults were killed data analysed with a Kruskal–Wallis test P > 0.05. with ethyl acetate and mounted with entomological pins. Ambient and body (rectal) temperature were taken at the same 4. Results time as collecting the specimens. Observations were made on the physical changes of the carcass over time (colour, degree of (dipterans) and beetles (coleopterans) constituted swelling, discharge of liquids and gases, etc). The carcass was 99.6% of the individuals collected. There were no significant weighed once a day with a scale [25]. differences between the three cadavers used with respect to the The first eggs mass founded, the colonising species, were presence and absence of species on each day (Kruskal–Wallis identified by collecting eggs from the carcasses. These were test, P > 0.05). The data then were combined to define the placed in plastic containers (three per pig) containing succession pattern. approximately 200 g of raw pig liver. The containers were In total, 36,892 individuals were collected (larvae, nymphs immediately covered with muslin and placed in styrofoam and adults) belonging to 6 orders, 53 families, 42 genera and 98 boxes. The development of the insects that hatched was species and morph species (Table 1). Of these, 34,832 (94.4%) monitored until the adult stage was reached [26]. were flies (Diptera), 1,881 (5.1%) beetles (Coleoptera), 73 E. Martinez et al. / Forensic Science International 166 (2007) 182–189 185

Fig. 1. Daily temperature variations related with decomposition phase. Fig. 2. Daily weight variations related with decomposition phase.

(0.2%) bugs (Heteroptera), 73 (0.2%) wasps (Hymenoptera), 26 remains (Table 1), which were determined by the physical and (0.07%) earwigs (Dermaptera) and 7 (0.02%) cockroaches body temperature changes of the carcass [6]. (Dyctiopetera). Within the Diptera two families were pre- dominant: blowflies (Calliphoridae) (91% of all Diptera) and 5.1. Fresh (0–3 days) small dung flies (Sphaeroceridae) (6.4% of the Diptera). The other 2.6% corresponded to individuals belonging to another 20 A sharp decrease in body temperature was observed families. Of the coleopterans, four of the 15 families reaching levels below ambient temperatures (Fig. 1). represented were most abundant: featherwing beetles (Ptilli- Oviposition was observed in natural orifices, on the neck, dae) (46%), rove beetles (Staphylinidae) (24%), hister beetles around the wound and on the side of the carcass touching the (Histeridae) (19%) and soft winged flower beetles (Melyridae) ground. (8.6%). Immediately after death, the first adults of Muscidae According to the classification of ecological categories by (Dasymorellia seguyi, Fannia sp., Limnophora sp.) and Smith [2] and Magan˜a [39], the following groups were present Calliphoridae (Calliphora nigribasis and Compsomyiops among the entomofauna collected: necrophagous species, verena) arrived. From 4 h after death and until day 18, predators, parasites of necrophagous species, saprophagous Calliphora nigribasis were observed ovipositing. Three days species, omnivorous species and incidental or local fauna after death, 1st instar larvae of Calliphora nigribasis were (Table 1). collected in natural orifices and around the wound (Table 3 and Appendix A). The second species to oviposit was Compso- 5. Stages of decomposition myiops verena from day 3 until day 7. Other abundant adult insects during this stage were Five different stages of decomposition were observed during Leptocera sp., Ichnomyia sp. and coleopterans Actinopteryx the study: fresh, bloated, active decay, advanced decay and dry sp. and Dianous sp.

Table 2 Percentage of adult dipterans collected at different stages of decomposition Family Species Fresh Bloated Active Advanced Remains Calliphoridae Calliphora nigribasis 31.08 33.80 26.03 0.21 0.27 Compsomyiops verena 1.35 20.21 42.47 0.00 0.14 Sarconesiopsis magellanica 0.00 1.38 2.05 0.00 0.00 Muscidae Limnophora sp. 16.22 3.05 1.37 0.07 0.14 Dasymorellia seguı´ 6.76 1.66 4.11 0.14 0.00 Muscina stabulans 4.05 17.17 12.33 0.21 0.00 Syllimnophora atroviatta 4.05 3.32 0.68 0.34 0.00 Fannia sp. 2.70 0.28 0.00 0.00 0.00 Hydrotaea sp. 0.00 0.83 3.42 0.34 0.00 Sarcophagidae Helicobia sp. 1.35 0.55 0.00 0.00 0.00 Microcerella sp. 0.00 0.28 0.00 0.00 0.00 Piophilidae Stearibia nigriceps 0.00 0.00 0.00 0.69 0.41 Sphaeroceridae Leptocera sp. 20.27 13.30 3.42 95.54 98.10 Anthomyzidae Ischnomyia sp. 12.16 1.11 0.68 0.00 0.00 Syrphidae Eristalis sp. 0.00 0.83 0.68 0.07 0.00 Tachinidae Unident 0.00 1.67 2.68 0.14 0.00 Dolychopodidae Unident 0.00 0.00 0.00 1.78 0.81 Sciaridae Unident 0.00 0.28 0.00 0.48 0.14 186 E. Martinez et al. / Forensic Science International 166 (2007) 182–189

5.2. Bloated (4–16 days) Table 3 Percentage of dipteran larvae activity at each stage of decomposition Body temperature began increasing during this stage Family Species Bloated Active Advanced Remains (Fig. 1) as a result of insect activity and the putrefaction Calliphoridae Callı´phora 89.24 81.55 51.07 31.89 process [40]. The carcass lost 10% of its initial weight nigribasis (Fig. 2). Abundant masses of larvae were observed through- Compsomyiops 0.00 8.23 44.41 62.52 out the carcass. First and 2nd instar larvae belonging to boliviana Calliphoridae, Sarcophagidae and Muscidae were found as Compsomyiops 10.14 8.77 1.43 0.17 verena well as 3rd instar larvae of Calliphora nigribasis, Compso- myiops verena,Muscidae(Azelia sp. probably) and Sarco- Muscidae Helina sp. 0.00 0.00 0.06 0.00 Hydrotaea sp. 0.00 0.00 0.48 1.37 phagidae (Appendix A). Azelia sp. 0.51 1.27 2.20 0.70 A large number of adults of Calliphora nigribasis, (probably) Compsomyiops verena, Leptocera sp. and Muscina stabulans Muscina sp. 0.00 0.18 0.01 0.00 were attracted to the carcass during this stage (Table 2). Piophilidae Stearibia 0.00 0.00 0.32 2.01 Ovipositing individuals of these species were also collected. nigriceps A surge in predators was evident at this stage. New groups of Sphaeroceridae Leptocera sp. 0.00 0.00 0.00 1.34 coleopterans appeared: Neopachylopus sp., Dasyrhadus sp.; Sarcophagidea Indent 0.10 0.00 0.00 0.00 Anacytyus sp., Dianous sp., Lathropinus sp., Carabus,in Tabanidae Indent 0.00 0.00 0.02 0.00 addition to other groups of dipterans, dermapterans, hyme- nopterans and hemipterans (Table 1). predation) and of the other species (inter-specific predation). New saprophagous species were first observed at this stage such 5.3. Active decay (17–30 days) as: Stearibia nigriceps (Piophilidae), collected after day 40 and Helina sp. and Hydrotaea sp. (Muscidae), collected after day 34 The temperature of the cadaver remained low and only (Appendix A). increased in the last days of active decay (Fig. 1). A sharp and Pupae of Calliphora nigribasis, Compsomyiops boliviana rapid loss of weight occurred (Fig. 2) in which 52% of the total and some Muscidae were found in the remains of the carcass. body weight was lost. Pupae of Calliphora nigribasis and Compsomyiops verena were With respect to the insects present, all three larval instars of collected at a distance of 2 m from the carcass among layers of Calliphora nigribasis and 2nd and 3rd instar larvae of moss on the ground. These were placed in rearing containers Compsomyiops boliviana and 3rd instar larvae of Compso- and raised to adults (Table 1). myiops verena were found. No individuals of the family The number of adult Calliphoridae dropped considerably, Sarcophagidae were observed (Table 1). Larval activity mainly being replaced by small dipterans such as Leptocera sp. and took place in large masses concentrated towards the side of the Stearibia nigriceps which may have been attracted by the carcass touching the ground. This activity continued to be decomposing tissues and the BOD (Table 2). dominated by Calliphora nigribasis and Compsomyiops verena. Ovipositing specimens of the latter species were 5.5. Dry remains (52–83 days) collected on days 18 and 19 (Table 2). Other predators present were individuals of hymenoptera The last collections were made during this stage. By the end, (Ichneumonidae and Vespidae) and coleoptera (Staphylinidae) 83% of the initial body weight had been lost (Fig. 2). Only (Table 1). remains of skin and bones were left. The BOD had dried and some of this had mixed with the soil. 5.4. Advanced decay (31–51 days) The small numbers of larvae found were concentrated in the extremities (hooves), places providing shelter and where small Peaks in cadaver temperature were reached during this stage amounts of soft tissue remained. Second instar larvae of with a maximum value of 100.40 8F(Fig. 1). By the end of Calliphora nigribasis and Compsomyiops boliviana were found advanced decay, the body had lost 75% of its initial weight until days 57 and 64, respectively. Third instar larvae and pupae (Fig. 2) and a mucilaginous material was found on and around were found until the end, as was the case with 3rd instar larvae the cadaver. This by product of decomposition (BOD) consists of Stearibia nigriceps (Appendix A). some of the internal tissues with insect material and other products of decomposition [41]. 6. Discussion Larval activity in the carcass was notably reduced although it continued to be dominated by Calliphoridae (Table 3). The Five stages of decomposition of the cadaver were observed: majority of the larval masses were concentrated in the muddy fresh, bloated, active decay, advanced decay and dry remains. ground under the body. Several dead larvae were found on the The different stages were categorized according to changes in body, underneath it and in the immediate vicinity. We colleted appearance and body temperature. The fresh stage lasted four Calliphora nigribasis and Compsomyiops boliviana larvae days and the bloated stage lasted 13 days. This was longer that predating individuals of its own species (intra-specific the corresponding periods reported in other studies [1,6,41], E. Martinez et al. / Forensic Science International 166 (2007) 182–189 187 where the fresh stage lasted for 2 days and the bloated stage observed ovipositing until the first few days of the active lasted from 4 to 8 days. In our study, subsequent stages stage while in Hawaii, Chrysomya megacephala and Chryso- conformed to the time periods given in the aforementioned mya rufifacies were only observed ovipositing during the fresh studies. This situation may be explained by the particular stage. This late oviposition could also explain the presence of conditions of cooler temperatures in our studies compared with 2nd instar larvae of Calliphora nigribasis and Compsomyiops the other reports. boliviana during the dry remains stage and 3rd instar of Calliphora nigribasis and Compsomyiops verena were the the same species being present until the last day of sampling first species to colonise the carcasses. This differed from the (Appendix A). study carried out by Wolff et al. [6] in Colombia at 1450 m Larvae of Piophilidae (Stearibia nigriceps = P. foveolata) above sea level where Phaenicia sericata was the first species and Muscidae (Hydrotaea sp.) were founded in advanced and to arrive. remains. Stearibia nigriceps important forensic indicator During the bloated stage and active decay, Calliphora because arrives frequently after saponification in the last nigribasis, Compsomyiops boliviana and Compsomyiops decomposition stages [1,43]. In the present and other studies, at verena were mainly responsibleforthelossofcarcass Colombia, Stearibia nigriceps presented the same behaviour tissue. The presence of a large number of individuals of these [6,22]. The individuals identified as Azelia sp. and Helina sp., species is the main reason for the rapid loss of weight during following Smith [35], were all larvae. Due to the early age advanced decay and consequently a reduced availability of stages and the lack of a larvae key for this group, we do not give food (Table 2, Fig. 2). This competition for resources may, in a definitive identification for these individuals. turn, may have contributed to the interspecific predation In this region, and with the specific environmental between larvae of Calliphora nigribasis and Compsomyiops conditions, we can designate indicator species for the different boliviana. stages of decomposition, which were determined by the The succession pattern observed shows similarities between physical and body temperature changes of the carcass and those described by Anderson and VanLaerhoven [1], Wolff based on the presence of the species for the first time on each of et al. [6] and Tullis and Goff [41], in those two principal groups the different decomposition stages, as follows: fresh - (Diptera and Coleoptera) were represented. Diptera, especially Calliphora nigribasis; bloated – Compsomyiops verena; active Calliphora nigribasis and Compsomyiops boliviana continued decay – Compsomyiops boliviana; advance decay – Stearibia their necrophagous activity during the whole decomposition nigriceps and Hydrotea sp.; dry remains – Leptocera sp. process (Table 3). which normally is not the case in most reports Although indicator species are not used for the estimation of in which Calliphoridae are present only up to the first 3–4 stages PMI, they are useful to know the stage of decay when of decomposition, but are not found on dried. combinations of species occur. The environmental conditions of the study area, at an This is the first report of Compsomyiops boliviana in altitude of 3035 m and with environmental temperatures Colombia. fluctuating between 9 and 23 8C during the day and 1to 7 8C at night, influenced the process of decomposition. The 7. Conclusions decomposition process of the carcasses took 83 days as opposed to the 45 days reported by Carvalho and Linhares Altitude, bio-climatic factors and vegetation determine [19] in Brazil during the wet season where the same 10 kg species distribution in the tropics. This is clearly shown by this model was used. Although the weight of the pigs was equal, study carried out on Paramo at 3035 m above sea level where the rate of decomposition may have been influenced by the the Andean species, Calliphora nigribasis, Compsomyiops environmental temperature. This study was done at 3035 m boliviana and Compsomyiops verena were found. These species above sea level, whereas the study by Castillo-Miralbes [42] had not previously been reported in studies of forensic was done at 300 m above sea level and at higher ambient entomology in Colombia, conducted in different climactic temperatures. zones. These species may be important indicator species of The rapid weight loss of the cadavers was a result of the specific stages of decomposition in this bio-climatic zone. conversion of carcass biomass into larval biomass and the Finally, a succession table is presented for insects on a carcass subsequent exit of insects from the body during pupation [41]. in Paramo. This aims to be a useful instrument for other The percentage of carrion removed over time was much less researchers and forensic scientists working in similar areas than a study carried out in Hawaii [41] with pigs of a similar (Appendix A). size. At the end of the active stage in Hawaii, 80% of body weight had been lost and subsequently only a small reduction Acknowledgements was observed. At the end of this study [41] 18% of the body weight remained, whereas in our study only 52% of the body This study received financial support from Colciencias weight had been removed by the end of the active stage. At the (Instituto Colombiano para el Desarrollo de la Ciencia y la advanced stage, 75% had been lost and only 13% was left at the Tecnologı´a) project number 1115-05-11503 and from the dry remains stage (Fig. 2). This may be explained by the Universidad de Antioquia, Medellı´n, Colombia and from Drs marked difference in the ovipositing habits of the dipterans. Jeffrey Wells and Carl Lowenberger for the revision of Calliphora nigribasis and Compsomyiops verena were manuscript. 188 E. Martinez et al. / Forensic Science International 166 (2007) 182–189

Appendix A

Succession table (presence–absence) Species Dev Fresh Bloated Active

0123456789101112131415161718192021222324252627272930

A L100011111111 1 1 1 1 1 1 1 1111111111110 L200000111111 1 1 1 1 1 1 1 1111111111111 L300000000001 1 1 1 1 1 1 1 1111111111111 B L200000000000 0 0 0 0 0 1 1 1111111111111 L300000000000 0 0 0 0 0 0 0 0001111111111 C L300000000011 1 1 1 1 1 1 1 1111111111111 D L200000000001 1 1 0 0 0 0 0 0111110100000 L300000000000 0 0 0 0 1 0 1 1011011110111 E L300000000000 0 0 0 0 0 0 0 0000000000000 F L200000000000 0 0 0 0 0 0 0 0000000000000 L300000000000 0 0 0 0 0 0 0 0000000000000 G L300000000000 0 0 0 0 0 0 0 0000010001110 H L200000000000 0 0 0 0 0 0 0 0000000000000 L300000000000 0 0 0 0 0 0 0 0000000000000 I L100000011110 0 0 0 0 0 0 0 0000000000000 L200000001011 0 0 1 0 0 0 0 0000000000000 L300000000001 0 1 0 0 0 0 0 0000000000000 J L300000000000 0 0 0 0 0 0 0 0000000000000

Species Dev Advanced Remains

31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67

A L11000000000000000000000000000000000000 L21111111111111111110111100010000000000 L31111111111111111111111111111111111111 B L21111111111111111111111111111111111000 L31111111111111111111111111111111111111 C L31111110110000010111011110100000000000 D L21010010011110110000100000000000000000 L31111111111111111111111111110111100000 E L30000110000001000001000000000000000000 F L20001000000000100110110000100000000000 L30000100101111101101111011111111101100 G L31000000000000000000000000000000000000 H L20000000000000000101100000000000000000 L30000000001001011111111111111111011011 I L10000000000000000000000000000000000000 L20000000000000000000000000000000000000 L30000000000000000000000000000000000000 J L30000000000000000000001000000010000001 L: larva, Species A: Calliphora nigribasis,B:Compsomyiops boliviana,C:Compsomyiops verena,D:Azelia sp., E: Helina sp., F: Hydrotaea sp., G: Muscina sp., H: Stearibia nigriceps, I: Sarcophagidae (unidentified), J: Leptocera sp.

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