J. Aust. ent. SOC.,1983,22 137-148 137 THE SUCCESSION AND RATE OF DEVELOPMENT OF BLOWFLIES IN CARRION IN SOUTHERN QUEENSLAND AND THE APPLICATION OF THESE DATA TO FORENSIC ENTOMOLOGY M. A. O'FLYNN Department of Parasitology, University of Queensland, St Lucia, Qld 4067.*

Abstract The succession and rate of development of in carrion is potentially a useful and accurate tool for determining the length of time elapsed since death, but the accuracy of this method in Queensland has been severely limited by lack of data. The occurrence of the following species in carrion in the Brisbane district and at a site 450 km west of Brisbane from 1975 to 1979 is discussed: Lucilia cuprina (Wiedemann), Lucilia sericata (Meigen), augur (F.), Calliphora stygia (F.), Calliphora hilli (Patton), Chrysomya rufifacies (Macquart), Chrysomya varipes (Macquart), Chrysomya megacephala (F.), Chrysomya nigripes Aubertin, Chrysomya saffranea (Bigot), Hemipyrellia ligurriens Wiedemann, Chrysomya megacephala (F)., Tricholioproctia tryoni (J. and T.), Ophyra spinigera Stein and Australophyra rostrafa (R.-D.). Detailed observations at constant temperatures were made on rate of development of commonly infesting human cadavers. The duration of the egg, first and second larval instars, total feeding period, total larval period, pupal period and egg to adult period are given for the following species at the temperatures indicated: L. cuprina (15-34"C), C. augur (9-28"C), C. srygia (9-28"C), Ch. rufifcies (20-34°C) and A. rostrara (9-28°C). Limited data on rate of development of Ch. varipes, Ch. sajranea, Ch. nigripes and Ch. megacephala are also included. The application of these data to forensic entomology is discussed.

Introduction The use of blowflies to calculate time of death has been discussed by several authors, notably Megnin (1894), Easton and Smith (1970), Leclercq (1969, 1974,1976) and Nuorteva (1974, 1977). Useful information on succession and rate of development was provided by other authors, particularly Fuller (1934), Johnston and Tiegs (1922), Mackerras (1933), Norris (1959, 1965, 1966) and Waterhouse (1947) in Australia and by Bohart and Gressitt (1951), Deonier (1940), Payne (1965, 1967), Reed (1958), Roy and Siddons (1939) and Wijesundara (1957) overseas. However, the work of these authors was not carried out primarily for forensic purposes and therefore does not include such vital information as the duration of each larval instar. Furthermore, differences in the climate and in the composition of the blowfly populations between Queensland and southern Australia make much of the data from southern Australia inapplicable to the Queensland situation. Research was therefore undertaken from 1975 to 1979 to provide detailed information on distribution, behaviour and rate of development of carrion-breeding blowflies in the field in Queensland and on rate of development under controlled conditions in the laboratory, in order to improve the accuracy of estimates of length of time elapsed since death. Data on rate of development at constant temperatures must be used in conjunction with field data (O'Flynn 1976,1980; O'Flynn and Moorhouse 1979), data on the behaviour and distribution of the species concerned (Norris 1959, 1966; O'Flynn and Moorhouse 1979; Zumpt 1965) and the keys and descriptions of their immature stages (Cantrell 198 1; Fuller 1932; Kitching 1976; O'Flynn and Moorhouse 1980). Materials and methods Observations on carrion were carried out at 2 sites in southern Queensland: (I) the University of Queensland farm at Moggill on the outskirts of Brisbane (27" 25'S, 152" 54'E): (2) in grazing country in the Thallon district (28" 303, 148" 57'E), ca 450 km west-south-west of Brisbane. Climatic data (Bureau of Meteorology 1975) for these 2 sites are given in Table 1, the data for St George, approximately 38 km north of Thallon being the closest approximation to that for Thallon. Carcasses of large , mainly , pigs and dogs were used for field experiments. When possible, maggots were collected twice daily until no maggots remained on the carcass: however, shorter periods of observation were sometimes unavoidable. As small a sample as possible was collected to avoid disrupting the succession while still giving a representative sample of the maggots present at each site on the carcass; the sample size depended on the abundance of maggots at each site. Larvae were identified by the keys of * Present address: Department of Primary Industry, Edmurrd Barton Building, Barton, A.C.T. 2602. 138 M. A. OFLYNN

Kitching (1976) and OFlynn and Moorhouse (1980) but if there was any doubt a number were reared to adults. Information on seasonal abundance, behaviour and development of blowflies in animal carrion was supplemented by observations of blowflies collected from human cadavers from throughout Queensland (Moorhouse and O’Flynn unpub. data). The species found to be the most common were cultured in the laboratory and detailed observations made on their rate of development at constant temperatures. Eggs were collected on liver and larvae reared on “pet mince”, a mixture of muscle and offal obtained from a local butcher. The age of the meat used for rearing was varied according to the preference of the species being reared, although most species could be reared successfully on either fresh or putrid meat. For Ausrralophyra rostrata (R. -D.),the meat in which a generation of Lucdia cuprinu (Wiedemann) had been reared, was used. In the early stages of development, twice-daily observations were made, being reduced to once-daily when the maggots reached the third instar. For each species, observations at each temperature were carried out on 2 batches of approximately 100 eggs, which were culled to approximately 30 larvae; further repetitions were made only if the first 2 sets of data did not agree.

Results Succession in field carcasses Data on decomposition and succession in carrion are summarised in Tables 2-4. Time since death is given in days, ‘day 1’ covering the period from death to 24 h after death. The stage described as ‘wandering larva’ is the post-feeding larva of Fraenkel and Bhaskaran (1973). The ‘pupa’ is the stage commencing with the formation of the ‘white pupa’ of Fraenkel and Bhaskaran (1973). The stages of decomposition are described according to the classification of Payne (1965) with the addition of the stage ‘slightly decomposed’. Payne’s stages after ‘advanced decay’ are combined into 1 as the carcass in those stages is not infested by blowflies. These stages are represented in Table 2 as follows: 1x-slightly decomposed, 2x-bloated. 3x-active decay, 4x-advanced decay, 5x-stages after advanced decay. The presence of a particular species on a given day, is indicated by an asterisk. In addition to the species mentioned in Tables 2-3 Tricholioproctia tryoni (J & T) was found as first instar larvae on day 1 in summer and on day 2 in winter; Calliphora hilli (Patton) was found as second instar larvae on day 6 in winter, on 1 occasion only;

L.cuprina 0 Ch.rufifacibs r Ch. varipws

’a-0

10 20 30 40 Iu 10 20 30 40 0 OC 0 FIG.1-The relationship between temperature and development from egg to adult for L. cuprina, C. stygiu, C. augur, Ch. rujifucies and Ch. varipes. TABLE1 CLIMATICDATA AT EXPERIMENTALSITES Jan Feb Mar APr May June July Aug Sept Oct Nov Dec 8 Si George 6 Rainfall (mm) 73.0 62.0 55.0 30.0 35.0 35.0 33.0 26.0 28.0 38.0 44.0 51.0 2 Temperature Mean daily % maximum("C) 34.2 34.2 31.4 28.4 23.0 20.3 19.3 21.0 24.8 29.0 31.9 33.4 Mean daily c minimum("C) 21.3 21.1 18.7 14.2 9.5 6.7 5.3 7.4 10.6 15.0 17.3 19.8 E

Moggill Rainfall (mm) 167.0 212.0 81.0 87.0 96.0 19.0 82.0 53.0 22.0 155.0 137.0 148.0 1 Temperature Mean daily maximum("C) 30.1 28.6 28.6 26.9 24.2 21.0 20.8 22.6 24.0 26.7 27.4 28.8 Mean daily minimum("C) 19.2 19.0 17.0 13.9 11.6 8.8 6.8 8.6 10.3 14.1 14.9 17.4 140 M.A. OFLYNN and Hemipyrellia ligurriens Wiedemann was found once, as first instar larvae on day 2 in summer. All of those species appeared to be rare in the experimental areas. Australophyra rostrata was, however found in several carcasses at Moggill; during winter it was found as third instar larvae from day 8 to day 27, and in summer it persisted for up to 8 w in large carcasses. It also occurred in winter carcasses at Thallon; third instar larvae were found in the rumen contents of a sheep 8 w after death, and in the burned remains of a dog 7 w after death (5.5 w after burning) and first and third instar larvae were found in the creamy flesh of an old sheep carcass. Data derived from laboratory studies at constant temperatures The duration of each stage of development over the range of temperatures for which complete development was observed, are given in Tables 5-7 for Lucifia cuprina, Calliphora augur (F.), Calliphora stygia (F.), Chrysomya ruJifacies (Macquart), Chrysomya varipes (Macquart) and Australophyra rostrata. The relationship between temperature and development from egg to adult for the first 5 of these species is shown in Fig, 1. Tables 5-7 include data for Chrysomya nigripes Aubertin, Chrysomya saJranea (Bigot) and Chrysomyamegacephala (F.), at 28 “Conly. The data in Tables 5- 7 are expressed as follows: “n.d.” means no data were available, “1-4 d” means that the duration of the stage was not known exactly but was between 1 and 4 d, “1 to 4 d” means that the minimum observed duration of that stage was 1 d and the maximum 4 d; the symbol “-” indicates the figure is approximate only, a line extending through 2 or more stages of development indicates that the period stated represents the duration of those stages combined. Eggs of L. cuprina failed to hatch at 9 “C.At 40T,L. cuprina eggs hatched and the larvae reached the third instar but failed to pupate; the total feeding period occupied less than 48 h and the egg and first instar together, less than 23 h. Approximately 10% of the C. stygia eggs hatched at 5 “C and the larvae reached the third instar but none pupated. All the larvae died without leaving the food, excepting 1 mature larva which left the food 90 d after hatching. The first instar lasted between 20 and 26 d; some larvae reached the third instar after 23 d but some remained in the second instar more than 38 d after the first moult commenced. At 34”C, few C. stygia pupated and no adults emerged. The total feeding period lasted only 2.5 to 3.5 d, but some larvae did not pupate until 8 d after the eggs hatched. C. augur is ovoviviparous. At 5°C the first instar lasted less than 3 d and the second, from 14 to 18 d, the total of these 2 instars occupying between 17 and 21 d. The minimum period from larviposition to commencement of wandering was 54 d; some larvae survived for 110 d but none pupated. At 34”C, the first and second instars together occupied 24 h, and the total larval period lasted 4.5 d of which the larvae fed for 2.5 d. Some larvae died in the food but mQst pupated, although no adults emerged. At 40”C,larvae commenced to leave the food after 30 h, but all died without pupating. Eggs of Ch. rufijacies failed to hatch at 9 “C.At 15 “C, eggs hatched after 2 d and most larvae reached the third instar but none pupated. Some larvae entered the second instar after 3 d but some not for 4 d or more; the second instar lasted 5 d. At 40T, the eggs commenced to hatch within 20 h, and the first 2 larval stages together lasted between 20 and 30 h, the total larval feeding period occupying 2.5 to 4 d. No larvae pupated at that temperature. At 45°C all larvae died within 45 h of hatching, most during the second instar. At 34”C, A. rostrata larvae commenced to leave the food after a minimum of 3 h, but none pupated. Comparison ojjeld and laboratory data To estimate the effectiveness of using data on development at constant temperature, combined with air temperatures at the carcass site, to estimate rate of development in a carcass, the actual and predicted development of Ch. rufifacies and Ch. varipes in 3 carcasses was compared. The data are presented in Table 8; the discrepancy between actual duration of development in a carcass and the duration predicted by present methods, is clear from that table. TABLE2 CARRION DECOMPOSITION ANDBLOWFLY SUCCESSION IN CARRION AT MOGGILL IN SUMMER (OCT-APR), 1975-1979 Days after death

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28

State of decomposition lx 2x 3x 4x 5x

Species L. cuprina First instar * Third instar *** m Ch. ruffacies * Eggs Second instar ** Third instar **** Q Wandering larvae * 4 Pupae * Ch. varipes Third instar **** Wandering larvae * Pupae * Teneral adults * Ch. saffranea * Eggs Third instar * Ch. nigripes' Second instar * Third instar ...... Pupa ************** Teneral adults * 0. spinigera' First instar * Second instar **************** Third instar ****************

0. spinigera weredeveloping in froth in the mouth, and in froth-soaked litter under body of dead cow. When the rumen contents were exposed in the third week, 0. spinigera,Ch. nigripesand A.rostrata developed in this material. e- 142 M. A. O'FLYNN

*** * * * * * * * * * * * * * * * * * ** * * ** ** * ** * ** * * ** * * * ** ** * * * ** * * * ** *

** * * *** I** *

** * ** TABLE4 CARRION DECOMPOSITION AND BLOWFLY SUCCESSION IN CARRION AT THALLON IN SUMMER (NOV-APR) AND WINTER (MAY-OCT), 1975-1979 Days after death SUMMER 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 State of decomposition lx 2x 2x 3x

Ch. ruJJacies Eggs *** Third instar ** Ch. varipes * Eggs Third instar ** Ch. saffranea Eggs * Wandering larvae * Days after death WINTER I 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28

~ ~ State of decomposition' lx lx 2x 3x 4x 5x E L. cuprina 0 First instar * z C. augur First instar ********* Second instar ******** Third instar ******* C. srygia First instar ****** Second instar ******** Third instar ******* Ch. ruJJacies Second instar * Third instar ***** Pupae * Teneral adults * C. varipes Teneral adults *

1 One carcass remained in stage lx until day 9 144 M. A. OFLYNN

TABLE5 RATE OF DEVELOPMENT OF COMMON BLOWFLIES AT CONSTANT TEMPERATURE

Temp First larval Second larval Total larval "C Egg stage stage stage Pupa

Lucilia cuprina 15 -45h -48 h -48 h 18->40d -25 d 20 26 h 2.5-3 d 7-10 d 13-14 d 25 Y 13-19h n.d.' n.d. n.d. n.d. 28 n.d. -12h -12h 4-10 d' 7d 34 n.d. n.d. n.d. 3.5-6 d 6d

Calliphora augur (ovoviviparous) 9 - t4d 34- > 55 d2 47 d -6 d 15 - -2.5 d 11 d 25 d 20 - t24h t24 h 7-10 d 14 d -47 h 25 - t20h 5-9 d 8d -33 h 28 - n.d. n.d. 6-9 d 6-7 d

Calliphora stygia 9 <3 d3 -5 d -4 d >28 d4 37-40 d 15 n.d. n.d. n.d. 12-14 d 19 d 20 24 h 2d 9d 13-14 d 3d 25 >11 h -20 h <24 h I->9 d -10d 2-3 d 28 n.d. n.d. n.d. 5.5-10 d 9d

Chrysomyia rujifacies 20 -24h 24-48 h n.d. n.d. 6-8 d 3d 25 16-22 h n.d. n.d. n.d. n.d. 28 7-20 h <20 h 5.5-6 d 4-5 d

~ -28h 34 n.d. -20 h 4-5.5 d 3d

~ -23h 42-46 h

Chrysomya varipes 20 n.d. n.d. n.d. 14 d 10 d 25 n.d. n.d. n.d. 6-9 d 5d

28 ~ >24h n.d. n.d. - 48h 34 n.d. n.d. n.d. -6 d -3 d

Chrysomya nigripes 28 n.d. 2-5 d 5-10 d 4-5 d Chrysomya sajianea j 28 8-10 h -12h n.d. 3-4 d 4-5 d -42 h

Chrysomya megacephala 28 n.d. n.d. n.d. 4d 4d

Australophyra rostrata 9 16 d 13 d 73 d6 42 d 15 3 d- -4 d -4 d 25->49 d 20-26 d 20

most pupated after 4-5 d. most pupated during 7th week. larvae visible in some eggs at 40 h. most pupated during 5th week. data for Ch. saffiunea includes data for development at room temperature (mean 28 "C, range 20-35 "C). periods quoted include the egg period. ' no data available. BIOLOGY OF CARRION FLIES 145

TABLE6 DURATION OF TOTAL LARVAL FEEDING PERIOD OF COMMON BLOWFLIES AT CONSTANT TEMPERATURES

Species Temp. Duration Species Temp. Duration “C “C

L. cuprina 15 9-15 d Ch. rufifbcies 20 9d 20 3.5-4 d 25 n.d. 25 n.d. 28 3.3-5 d 28 2->4 dl 34 3-4 d 34 2.5-4 d Ch. varipes 20 12-13 d C. augur 9 -14d 25 5-6 d 15 6-1 d 34 >3 d 20 4-5 d 25 3-4 d Ch. nigripes 28 4-1 d 28 2.5-4 d Ch. sajraneu2 28 3d C. stygiu 9 16-18 d 15 6d Ch. megarephala 28 3d 20 5-> 10 d 25 3-6 d A. rostruta 9 70 d3 28 3-5 d IS.. 10 d 20 8-9 d 28 5-6 d

most larvae fed for 2-3 d. the data for Ch. saflranea include data for development at room temperature (mean 28”C, range 20- 35°C). most of the larvae pupated in the meat so the actual feeding period could have ended earlier than the time indicated. The period quoted includes the egg period.

Discussion In forensic investigations, our estimates of date of death have been based on mean air temperature, derived from daily maxima and minima recorded over the relevant periods at the Meteorological Stations nearest to the sites. As shown in Table 8, estimates based on overall mean may be erroneous. Estimates based on mean maximum air temperature would appear to be more accurate for Queensland, except during winter. The temperature at which the maggots develop in a corpse is often much higher than air temperature, and varies according to the state of decomposition and the location within the corpse (Deonier 1940; O’Flynn 1980; Reed 1958; Waterhouse 1947). The size of the body should be taken into consideration, as in small cadavers such as babies, temperatures would not reach the high levels found in large cadavers. Pupae and post-feeding larvae may be some distance from the corpse, and the corpse may be reduced almost to a skeleton by the time those stages are reached. Those stages would therefore be subjected to the diurnal fluctuations in air temperature, and the rate of development at a fluctuating temperature may differ from that at a constant temperature even if the mean temperatures are the same (Ahmad 1936). In the laboratory experiments, the maggots were culled to prevent crowding, which would have raised the temperature of the culture above that of the incubator. Although this would have created an unrealistic situation by comparison with a carcass, it was considered that these preliminary investigations should be conducted at controlled temperatures rather than under field conditions, to minimise the number of uncontrolled variables. As expected (Nagasawa and Kishino 1965; Pradhan 1946; Wardle 1930)the curve representing the relationship between temperature and the duration of the egg to adult period (Fig. l), is not a straight line. No attempt has been made to analyse this data mathematically as the quantity and precision of the data were not considered sufficient, but it is clear from Fig. 1 that care is required in interpolating the data in Tables 5-7, , particularly at temperatures close to the extremes of the favourable range for development of each species. 146 M. A. O'FLYNN

TABLE1 DURATION OF DEVELOPMENT OF COMMON BLOWFLIES FROM EGG TO ADULT AT CONSTANT TEMPERATURES Species Temp. "C Duration Species Temp. "C Duration

L. cuprina 15 -43 d Ch. rufijacies 20 16-17 d 20 21-22 d 28 10-14 d 28 12-13 d 34 8d 34 10-11 d Ch. varipes 20 24 d C. augur' 9 83-94 d 25 11 d 15 36-38 d 34 9d 20 21-23 d 25 13 d Ch. nigripes 28 10-14 d 28 12-16 d Ch. saffranea2 28 1-9 d C.stygia 9 66 d 15 33 d Ch. megacephala 28 8-9 d 20 24 d 25 -17d A. rostrata 9 115d 28 16-20 d 15 49-70 d 20 28 d 28 18 d - I C.augur is ovoviviparous, therefore the figures for this species are for development from first instar larva to adult. data for Ch. saffranea include data for development at room temperature (mean 28 "C, range 20-35°C).

TABLE8 COMPARISON OF EXPECTED AND ACTUAL DURATION OF EGG TO ADULT PERIODS FOR CH. RUFIFACIES AND CH. VARIPES IN CARCASSES IN THE FIELD

Temperature ( C) Duration of development (days) Species Mean Range Actual Expected' min max overall min max overall

Ch. rujijacies 6 20 13 2-24 <222 nil 15-16 nil 14 22 18 10-24 12 nil 15-16 17-18 20 34 21 14-40 t12 16-17 8 14-16 Ch. varipes 14 22 18 10-24 12 n.d. n.d. 24 19 28 24 18-32 < 9 24 10 11 20 34 27 14-40 t12 24 9 10

I based on constant temperature equal to mean. teneral adults were found 22 d after the infestation of the carcass, no observations having been made during the intervening period. n.d. = no data.

The period from death to oviposition varies not only between species (Fuller 1934), but also within the 1 species (O'Flynn and Moorhouse 1979). The length of the pre-infestation period as well as the rate of development, may be affected by micro- environmental conditions in the vicinity of the corpse (Norris 1965, 1966; Nuorteva 1977; Payne 1967; Reed 1958) and these factors are not shown in the meteorological records. Tables 2-4 summarise data presented in detail by O'Flynn (1976, 1980). The variations between different types of animal carcass (Fuller 1934) and between animal and human remains are recognised. The table should, however, serve as a useful guide to the relationship throughout the year between state of decomposition, carrion fauna and length of time since death in the Brisbane area, in southern Queensland and also in northern New South Wales, where the carrion fauna is similar. Where several species and several stages of development are present, the combination of all the information from all of these will reduce uncertainty and give a BIOLOGY OF CARRION FLIES 147 more accurate estimate than would be possible if only 1 or more species were present. This highlights the importance of collecting a representative sample, and this should be stressed to police and others involved in collecting samples for forensic examination. It may be necessary in some cases to rear a sample through to adults in order to identify them or to infer from the duration of the subsequent development, how long the larvae had been developing in the corpse. Disadvantages of this procedure are that it prolongs the time taken to estimate date of death; it is virtually impossible to ensure that development is proceeding at the same rate as it would have, had the larvae been left undisturbed on the corpse; and other species, particularly Sarcopghagids, may invade the culture, leading to a false conclusion. When the specimens are collected as pupae, there is no alternative to the rearing method until a reliable method of determining the age of pupae of carrion flies is developed. Also, there is currently no way of knowing how long a period has elapsed since wandering (i.e. post-feeding) larvae have ceased to feed; this could introduce an error of weeks or even months at low temperatures. The age of these stages when collected, can be deduced only be rearing them to adults. A rapid and accurate identification of most species can now be made from the immature stages of the common species using the keys of Cantrell (1981), Kitching (1976) and O’Flynn and Moorhouse (1980); rearing to determine age at time of collection is essential only in the case of post-feeding larvae or pupae, or of those species for which insufficient data on rate of development are available. However, as species about which little is known may sometimes occur in corpses, it is always advisable to rear out a sub-sample in order to confirm the findings based on the immature stages. The importance of presenting specimens in a viable state, not preserved or moribund from anoxia or refrigeration as is often the case, should therefore be stressed to those collecting specimens for forensic examination. These data on rate of development and succession of the common carrion species of southern Queensland have considerably increased the accuracy of estimating date of death for human remains in that area. Further information on rate of development, especially of Chrysomya nigripes and of carrion-breeding Sarcophagids, is required in view of their frequent occurrence in carrion in that area. Emphasis should however be on field, rather than laboratory, studies.

Acknowledgments The research for this paper was carried out in the Departments of Parasitology and Entomology at the University of Queensland, under a grant from the Australian Wool Corporation. I wish to thank Dr D. E. Moorhouse. Mr G. Wolf, Mr L. Siddell and Mr G. Kirkup for their assistance and Mr R.Storey and Mr D. Morgan for their advice. Thanks are also due to my family for their assistance with the collection of data of Thallon.

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