Life table of blue bottle fly Lucilia cuprina (Wiedemann, 1830).
The life history of the L. cuprina was studied at 25[+ or -]2[degrees]C in the laboratory conditions on lamb meat as host. Egg we counted by daily meat 0life table by computer program. The obtained life history parameters of L. cuprina were as follows: the intrinsic rate of increase (r)= 0.231, the finite rate of increase ([lambda])= 1.260 day-1, the net reproduction rate (Ro) = 109.62 offspring per individual; and the mean generation time (T) =20.35 days. The maximum reproductive value of L. cuprina females was on the 14th day. The two-sex life table analysis showed a comprehensive description of the stage differentiation (age-stage specific survival rate, life expectancy of each age-stage, and reproductive value of each age-stage). This experiment has a significant impact on stage-specific control strategies and postmortem interval estimation.
KEYWORDS: Green Sheep Blowfly, Lucilia cuprina, life table, Ecology, Lamb meat.
Life table is often a simple method for keeping track of births, deaths, and reproductive output in a population of interest [1, 2]. Necrophagous Diptera belongs to several taxonomic groups such as blowfly (Calliphoridae), flesh fly (Sarcophagidae). Necrophagous Diptera has important forensic uses such as estimation of Postmortem interval based (PMI) on a composite of taphonomic factors that are affected by [3, 4] the geographic region or biogeoclimatic zone[5-7]. Lucilia cuprina is worldwide distributed and also found in the Mediterranean region including Egypt, Spain and Jordan [8-11]. Lucilia cuprina is a veterinary fly that induces sheep myiasis worldwide. L. cuprina is responsible for a condition known as "blow-fly strike" of sheep in several countries . Have a wide animal host such as sheep, goat, cow, donkey, and camel .
This species may be found on human corpse and have been used to estimate the postmortem interval (PMI) in forensic cases [14-16]. PMI is defined as the length of time between death and corpse discovery. The medical parameters begin to degrade that are needed to establish the cause, manner, and death time at the onset of death, [5, 17]. Study life history parameters of L. cuprina is basic to be used in PMI estimation [18, 19].
The objective of this study is to deliver two-sex life table analysis for L. cuprina isolate in Jordan, and to identify the meat host effect in life table parameters in comparison with other studies. Those parameters have useful and important applications in postmortem internal estimation in forensic field.
MATERIALS AND METHODS
The laboratory colony of Lucilia cuprina used in this study was established and the culture was initially obtained from dog carcass from Dead Sea. Adults from the stock colony of Lucilia cuprina were kept in cages (30 x 35 x 35 cm) at room temperature. The cages were made from a wooden floor, a gauze roof, and four wire glass sides. The gauze roof was supplied with an articulation to facilitate daily feeding, cleaning of the cage, and removal of eggs. Adults were supplied daily with granular sucrose, water, and sheep meat. Water was supplied by dipping a piece of cotton as a wick in a bottle filled with water, and the lamb meat was provided in a plastic Petri dish. Egg batches were removed daily and transferred to a fresh piece of lamb meat placed in a rearing modified plastic container (12 cm in diameter and 6cm depths with 9-12 small halls in the upper cover). At the prepupal stage, dry autoclaved sawdust was added to the Petri dish as a medium for pupation. Pupae were sieved or picked from the sawdust and transferred to cages for adult emergence.
Life table study and data analysis:
For the life table study, 58 eggs (laid during a 24 h period of L. cuprina were collected and put on a piece of fresh lamb meat for hatching. Each day, newly hatched larvae were transferred individually to 25 g lamb meat placed in a 100 ml beaker covered with muslin secured with a rubber band. At the pre-pupal stage, sawdust was added to the beaker for pupation. The emerged adults were paired (male, female) and kept in a muslin covered 1 L glass jar. Each jar was supplied with water, sucrose, and a piece of fresh lamb meat. Sucrose and water were checked daily and renewed as necessary, while the lamb meat was changed daily. Survival and fecundity were recorded for each individual until the death of the adult. The raw data for individuals were analyzed using the age-stage, two-sex life table approach [20, 21]. The computer program used in this analysis  is written in Visual BASIC for the Windows operating system and is available at http://184.108.40.206/Ecology/ (National Chung Hsing University, Taichung, Taiwan). The age-stage survival rate, the distribution of mortality rate, the age-stage life expectancy and stable age-stage distribution were calculated. Furthermore, the means and standard errors of the intrinsic rate of increase (r), the net reproductive rate (Ro), the mean generation time (T) and the finite rate of population increase ([lambda]), were calculated by using the Jackknife method [23, 24].
RESULTS AND DISCUSSION
Stage developmental time, adult longevity, and fecundity of L. cuprina on lamb meat are given in Table 1. Out of 58 eggs, 55 eggs were hatched successfully with an average time of 18.5 hour and with mortality rate at egg stage of 5.17%. the data showed that from 78.2% emerged adult, a 38.18% were males and 40.0% females (a 21.8% pre-adult mortality rate). Other laboratory studies of L. cuprina reared on beef meat showed 45:42 female: male sex ration 1:1 . Table 1 clearly indicated the absence of significant differences in the development times of the egg, larval, pre-pupa and pupal stage between both sexes. However, the adult longevity of female 19.73 days is significantly (P<0.05) longer than male 11.33 days by suing t-test. This finding is also applicable in the total longevity between female 33.27 days and male 23.52 days. These results are in agreement to that reported for other Calliphorid flies, Gabre found that longevity of the adult Chrysomya megacephala was 26.13 days .
The age-stage specific survival rate (sij) of L. cuprina (Figure 1) gives the probability that a newly laid egg will survive to age i and stage j. Moreover, this figure shows the survivorship and stage differentiation as well as the variable developmental rate. The probability that a new-born egg survives to adult stage is 0.38 for males and 0.4 for females.
Age of first reproduction by female has an important impact on population growth, the adult pre-oviposition period (APOP) and total pre-oviposition period illustrated in (Table 2). The maximum daily fecundity of L. cuprina female was 58 egg/female with maximum fecundity female of 394 egg/female during female life span. This results match the maximum daily fecundity of 50.3 eggs/female that obtained. This is not only due to the different geographical regions, but also to the different type of diet used in the study conducted .
Life table analysis parameters were calculated by using Jackknife method (Table 3). These results showed that the intrinsic rate of increase (r) of L. cuprina scored (0.2312), while the finite rate of increase ([lambda]) reached 1.2601. The net reproduction rate ([R.sub.0]) and the mean generation time (T) scored 109.62 and 20.35, respectively. Moreover, these results showed that the coefficient of variation is higher than that of the Egyption study. For instance, [R.sub.o] was higher by 12.1% in my study than that of the Egyption study Zied et al (Table 3). This finding also shows that L. cuprina is r-strategist, and characterized by a high rate of increase with a large fecundity and short generation time [27-29].
Age-specific survival rate ([l.sub.x]), the female age-specific fecundity ([f.sub.ix]), the age-specific fecundity ([m.sub.x]), and the age-specific maternity ([l.sub.x][m.sub.x]) of L. cuprina are shown in Figure 2. Since the egg batches were laid at discrete time intervals (8 peaks of females), mean daily fecundity ([f.sub.ij]) was 25.05 eggs. This result agrees with previous findings that showed L. cuprina and some other dipterans displayed discrete ovarian cycles . The age-specific survival rate ([l.sub.x]) is the probability a newborn will survive to age x. It is calculated by including all individuals of both sexes. The age-specific survival rate ([l.sub.x]) was 25% dead after 10 days of L. cuprina population in all stages.
Mortality rate and life expectancy of L. cuprina are shown in Figure (3). This figure showed the distribution of mortality during the whole life span. The highest mortality percentage of egg stage was represented by the first peak. The highest mortality percentage of larval stage was represented by the second peak. However, the highest mortality percentage of both prepupa and pupa were represented by the fourth peak (7th day). 22 females, and 21 males out of the 43 survival insects. The stage mortality rate was 5.2%, 10.3%, 5.2% 5.2% 37.9% and 36.2% for egg, larva, prepupa, pupa, female, and male, respectively.
The life expectancy ([e.sub.ij]) of each age-stage of L. cuprina was shown in Figure 4. [e.sub.ij] is used to predict the time individuals estimate of age i, and stage j. For instance, the life expectancy of a new egg was 22.3 days with decreasing the life expectancy with age. |It is clearly shown that the life expectancy value of the female adult is higher than that of the male adults (Table 1); whereas the adult longevity value of male is shorter than that of the female.
The reproductive value ([[nu].sub.ij]) is the expectation of offspring of age i and stage j (Figure 5). In this study, the reproductive value for a new egg ([[nu].sub.01]), which is the finite rate of increase ([lambda]) equals 1.2, while the highest reproductive value was at day 14. This implies that, female individuals of age 19 scored the highest contribution to the population 75 egg, compared to other ages, If the pre-oviposition period is counted as time from birth to fist reproductions in female (the total oviposition period, TPOP), the mean TPOP for all L. cuprina was 14.68 [+ or -] 2.59 days, a value close to the age peak reproductive value (Figure 5).
The estimation of PMI is dependent on the time the insect spent for development. This time is not only determined by ecological factors, but also on the type of diet the insect feeds. This means that PMI estimation is different from geographical region and another geographical region. This means that it is important to obtain local parameters of the developmental rate of forensic dipteral applications. This can be explained due to different ecological parameter between Jordan and other regions in the world. For instance, the time needed for the insect to develop becomes shorter when temperature becomes higher . Moreover, the type of diet affects the developmental rate parameters and fecundity. In conclusion, I showed, in this study, that the species lives in Jordan has different ecological parameters. For instance, the No. of eggs laid in the Jordanian species is 56. However, the No. of eggs laid of the Egyptian species is 105 eggs. This is not only due to the different geographical region, but also to the different type of diet used in the Egyptian study. This makes this study important of forensic purposes in Jordan. Since the PMI estimate of the local species is longer compared with the global PMI estimate based on the parameters for the same species.
Necrophagous diptera is an important model for forensic entomology in specific, and as part of forensic science in general. The importance of this study is in providing a practical, measurable, and creditable tool in criminal investigations. This study showed the role of model organism in forensic science is not limited to single host. Therefore, each forensic model organism parameters needs to be verified locally to optimize the output results. In fact, this study contributes to the forensic knowledge not only in finding the local variables parameters for a forensic model organism, but also in emphasizing the importance and the power of forensic entomology.
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Hazem S. Hasan
Department of Plant Production and Protection, Faculty of Agricultural Technology, Al-Balqa' Applied University, Al-Salt, 19117, Jordan.
Email: Hazem@bau. edu.jo
Address For Correspondence:
Hazem S. Hasan, Department of Plant Production and Protection, Faculty of Agricultural Technology, Al-Balqa' Applied University, Al-Salt, 19117, Jordan.
This work is licensed under the Creative Commons Attribution International License (CC BY).
Received 18 September 2016; Accepted 1 January 2017; Available online 16 January 2017
Table 1 : Basic statistics of life history data for Lucilia cuprina reared on lamb under laboratory conditions. Stage Sex No. Mean (Day) S.D S.E 55 1.00 0 0 Larva 49 4.24 0.97 0.14 PrePupa All 46 1.89 0.97 0.14 Pupa 43 5.65 1.66 0.25 Adult 43 15.63 6.21 0.95 Egg 22 1.00 0 0 Larva 22 4.55 0.67 0.14 PrePupa Female 22 2.00 1.31 0.28 Pupa 22 6.00 1.93 0.41 Adult 22 19.73 5.43 1.16 21 1.00 0 0 Larva 21 4.14 0.96 0.21 PrePupa Male 21 1.76 0.54 0.12 Pupa 21 5.29 1.27 0.28 Adult 21 11.33 3.51 0.77 Egg 12 1.00 0 0 Larva 6 3.5 1.52 0.62 PrePupa Unknown 3 2.00 0 0 Pupa - - - - Adult - - - - Table 2: The development time (days) for Lucilia cuprina reared on lamb under laboratory conditions. Parameter No Mean SEM Range Adult pre-oviposition period 22 1.14 0.64 1-3 (APOP) of female (days) Total pre-oviposition period 22 14.68 2.59 11-20 (TPOP) of female (days) Female Table 3: The population parameters of Lucilia cuprina reared on lamb under laboratory conditions. Population parameter Estimated by using Jackknife method Mean[+ or -]SE CV% The intrinsic rate of 0.2312[+ or -]0.013 5.62 increase r ([d.sup.-1]) The finite rate of 1.2601[+ or -]0.016 1.27 increase [lambda] (d.sup.-1) The net reproduction rate 109.62[+ or -]20.14 18.37 [R.sub.o] (offspring/individual) The mean generation time T (d) 20.35[+ or -]0.75 3.69
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|Author:||Hasan, Hazem S.|
|Publication:||Advances in Environmental Biology|
|Date:||Jan 1, 2017|
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