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Forensic entomological case study and comparison of burned Forensic entomological case study and comparison of burned

and unburned Sus scrofa specimens in the biogeoclimatic zone of and unburned Sus scrofa specimens in the biogeoclimatic zone of

northwestern Montana northwestern Montana

Seth Barnes The University of Montana

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Forensic Entomological Case Study and Comparison of Burned and Unburned Sus Scrofa Specimens in the Biogeoclimatic Zone of Northwestern

Montana

Seth Bames

B.S., University of Wisconsin-Oshkosh

Presented in partial fulfillment of the requirements

for the degree of

Master of Arts

The University of Montana-Missoula

2000

by

Approved by:

£Chairman

Dean, Graduate School

, S" (*?" 2oooDate

UMI Number: EP40119

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Barnes, Seth, M.A., Spring 2000 Anthropology

Forensic Entomological Case Study and Comparison of Burned and Unburned Sus Scrofa Specimens in the Biogeoclimatic Zone of Northwestern Montana (75 pp.).

Chairperson: Randall SkeltonThe goal of this experimental case study is to provide baseline rural forensic

entomological data for the Interior Douglas-fir/Ponderosa Pine biogeoclimatic zone of Missoula County in Western Montana. The first hypothesis is that successional fauna will be similar to that of Dillon (1997) who conducted well-replicated forensic entomological studies in a similar biogeoclimatic zone located in British Columbia, Canada. The second hypothesis states that the burned and unbumed specimens will have different successional fauna specific to their physical states. The third hypothesis is that the insect successional pattern and species diversity on carcasses present in a mixed urban/rural and rural area will be distinct (Ternenyl997). This will provide the forensic investigator with a species diversity index indicating the abundance of urban and rural species in two biogeoclimatic zones of Montana.The first pig was burned with one gallon of gasoline and the second pig remained

unbumt to act as the control. Phormia regina (Meigan) adults and immatures dominated among the order Diptera throughout the study until the post-decay stage when they co­dominated with Lucilia illustris (Meigen). Protophormia terraenovae (Robineau- Desvoidy) also played a small role. During the post-decay stage Prochyliza brericornis (Melander) and Prochyliza xanthostoma (Walker) of the family Piophilidae made a significant contribution to the decomposition process. Among the Coleoptera Thanatophilus lapponicus (Herbst) and Necrodes surinamensis (Fabricius) were the most representative families.Many similarities were found between this study and that of Dillon (1997) in a similar

biogeoclimatic zone. The parallels in decay rates and species diversity for these two studies conducted in an interior Douglas-fir and Douglas-fir/Ponderosa Pine Rocky Mountain biogeoclimatic zone cannot be accounted for as coincidence. The insect successional pattern was very similar for both pigs. The burned pig showed only a slight difference in the rate of decay and successional pattern. The families of insects involved in the decomposition process were similar for both the urban/rural and rural areas. The insects were typed to genus or species in this study. If applicable to human decedents, this baseline data will aid in the interpretation of Montana forensic anthropology and autopsy results to provide the medico-legal community with a more lucid explanation concerning time since death in forensic cases.

Acknowledgements

I would like to thank all who made this study possible. I would like to thank Frank, Patty, and Chris of the Lubrecht Experimental Forest for the use of equipment, housing, and their generous help. I would also like to thank Diana Six of the School of Forestry, University of Montana, for use of her laboratory, equipment, and help in identification. Additional thanks to Douglas Emlen of the Division of Biological Sciences, University of Montana, for his advice and assistance. Extra thanks to Elena Ulev for assistance in completing fieldwork and to my committee members for guidance. Finally, a thank you to everyone who helped in their own w ay, including the two Sus scrofa who became test specimens.

Table of Contents

Title Page AbstractAcknowledgmentsTable of ContentsList of TablesList of FiguresChapter 1-IntroductionChapter 2-Materials and MethodsChapter 3-ResultsChapter 4- DiscussionChapter 5-ConclusionsTablesFiguresReferences

Table NumberList o f Tables

Page Number

1. Insect Succession for the burnt Sus scrofa----------------------------------- 46-49

2. Insect Succession for the control S. scrofa----------------------------------- 50-53

3. Decay stage lengths from Dillon (1997) and the present study-------------54

4. Decay stage lengths from Temeny (1997) and the present study-----------54

Figure Number

List of Figures

Page Number

1. Diptera Species Diversity, Fresh: Burnt and Control S. scrofa ---------55

2. Diptera Species Diversity, Bloated: Burnt and Control S. scrofa------------ 56

3. Diptera Species Diversity, Decay: Burnt and Control S. scrofa---------------57

4. Diptera Species Diversity, Post-Decay: Burnt and Control S. scrofa---------58

5. Diptera Species Diversity, Remains: Burnt and Control S. scrofa-------------59

6. Coleoptera Species Diversity, Bloated: Burnt and Control S. scrofa----------60

7. Coleoptera Species Diversity, Decay: Burnt and Control S. scrofa------------61

8. Coleoptera Species Diversity, Post-Decay: Burnt and Control S. scrofa 62

9. Coleoptera Species Diversity, Remains: Burnt and Control S. scrofa---------- 63

10. Ambient and Internal Temperatures, days 1-30: Burnt and Control S. scrofa—64

Introduction: Chapter 1

The benefits and uses of forensic entomology are many. Insects have been used

in forensic science on nearly every continent in a number of diverse climates. This

variety of biogeoclimatic zones ensures that comparisons of data will show striking

contrasts as well as remarkable similarities. The widespread application of forensic

entomology is not surprising because it is typically used for conducting criminal

investigations. Every year forensic entomology plays an increasing important role in the

pursuit of justice.

The characteristic role of the forensic sciences is to provide law enforcement

agencies with comprehensive examinations of evidence provided by law enforcement

personnel. This evidence is commonly examined by crime laboratory sections such as

serology and latent prints. Unfortunately these sections rarely, if ever, examine

entomological evidence. Only the coroner or medical examiner frequently have the

opportunity to inspect such evidence, and in most cases, these individuals do not have an

extensive background in the study of insects. At this point, the entomologist is able to

contribute their scientific observations to the overall examinations of case evidence. This

data may provide law enforcement officials with the information they need to

successfully bring closure to open cases. Several areas of entomology may serve this

end.

Lord and Stevenson (1986) recognize three areas of forensic entomology: urban,

stored-product, and medicolegal. Urban and stored-product investigations usually

involve lawsuits pertaining to civil and consumer domains, respectively. Medicolegal

forensic entomology examines arthropod association with illegal activities, such as

1

murder. Utilization of this type of entomology involves determining the time since death,

or Post Mortem Interval (PMI). The PMI may be estimated in two ways. The first

method is to estimate the development time of each species collected at the scene. This

estimate is primarily used in the early stages of decomposition. Ordinarily those insects

exhibiting the longest developmental period of growth are used to ascertain the PMI.

The second method involves corpses in an advanced stage of decomposition.

This method involves observing arthropod successional patterns and compares them with

preexisting data for the same or similar arthropods in the relevant biogeoclimatic zone in

question. These methods complement examinations made by other forensic investigators,

including forensic anthropologists.

Both methods require the determination of behavior of the arthropods present.

Goff (1993) presents four types of arthropod feeding behavior, all of which may be found

during forensic investigaitons. The first type of behavior is necrophagy, feeding directly

upon the corpse. The second types of behavior is predation and parasitism. This group

frequently consists of individuals in the orders Coleoptera, Hymenoptera, and Diptera,

and frequently affect eggs and larva. The third type of behavior is omnivory, feeding

upon both corpses and other arthropods present. The fourth type of species is adventive

with use of the corpse possible in many different ways. Some of these species may feed

upon mold or fungi growing on the corpse while others simply extend their normal

habitat to include the microenvironment provided by the corpse.

Forensic entomology and anthropology share an interest in the circumstances and

processes surrounding a corpses process of decay. The study and application of forensic

entomology has, until recently, been relatively uncommon in the United States of

2

America. The only institution legally sanctioned to use human corpses is the

Anthropological Research Facility (ARF) in Tennessee. However, most researchers

frequently use animal models for surrogate humans. At ARF, corpeses are allowed to

decompose under a variety of conditions to simulate actual medicolegal forensic cases.

Animal models have been used in temperate environments (Dicke and Eastwood 1952,

Reed 1958, Payne 1965, Johnson 1975, Baumgartner 1988, Blackith and Blackith 1990,

Dillon 1997, Temeny 1997, and Adair 1999), tropical environments (Early and Goff

1986, Goff et. al. 1986, Goff and Odom 1986 and 1987, Goff 1992b) and arid

environments (Baumgartner 1986, Galloway et. al. 1989).

Entomological investigations are common outside the United States. A few of the

earliest works in Europe include those by Redi (1668), Bergeret (1855), Brouardel

(1879), Megnin (1888, 1894), and Motter (1898). Additional case studies and

experiments from the twentieth century include those conducted in such diverse countries

as Finland (Nuorteva et. al. 1967 and Nuorteva 1970,1974,1977 and 1987), Italy

(Introna et. al. 1998), Australia (Palmer 1980), South Korea (Rueda et. al. 1997), Egypt

(Tantawi et. al. 1996 and 1997), and Japan (Nishida 1984). Many investigations are not

listed here but may be found in sources such as Meek et al (1983), Vincent et al (1985)

and Smith (1986).

One of the earliest works completed by Redi (1668) consisted of leaving meat

exposed to, or protected from, flies. This work showed that spontaneous generation of

flies did not occur, and that flies contributed to the decay process. Bergeret's (1855) later

medicolegal forensic entomology work provided insight into similar methods and

materials still used in today's investigations. But it was Megnin's work from 1883-1898

3

which suggested for the first time that the process of decomposition occurred in a

predictable manner. These early studies gave status to the budding branch of entomology

among the forensic sciences. More recent and broad examinations of the subject have

been published by Smith (1986) and Catts and Haskell (1990).

The detection of various pharmaceuticals in forensic cases has also received

treatment in recent studies. Goff et. al. (1989,1991, 1993 and 1994) have examined the

evidence of the effects of cocaine, heroine, and other various chemicals upon decaying

tissues and the arthropods associated with them. Goff and Lord (1994) refer to this new

field as entomotoxicology and believe it has good prospects of becoming a standard

forensic technique when insects and chemicals are found together.

Forensic anthropology is similar to forensic entomology in that both attempt to

ascertain data about the deceased, frequently with only a small amount of material to

work with. The main contrast is that forensic anthropologists focus upon the bones of the

individual to determine aspects such as age, stature/weight, paleopathology, and sex. It is

difficult for the anthropologist to determine how long the individual has been dead, apart

from cultural markers. This is where entomology contributes as a partner in solving a

forensic case. With the entomologist's estimate of time since death (PMI), the

information gathered by the entomologist and anthropologist may be combined to form a

comprehensive evaluation of the activities surrounding the individual before and after

death. Until now the only forenstic entomological study from Montana was conducted by

Temeny (1997). Apart from several forensic cases (Barnes, unpublished data), forensic

entomology has not been used extensively within Montana.

4

Within the realm of forensic entomology one may discern:

• Whether the body was moved after death. Many insects are limited to particular

environments and are rarely found elsewhere (Goff 1992a). For example, aquatic insects

found on a corpse in an arid environment would indicate that the deceased spent some

amount of time in an aquatic environment after death.

• Whether the individual was killed inside or outside a structure (Meek, unpublished,

Haskell, unpublished). If insects typically found in urban environments, such as those

feeding upon stored goods, are found upon a corpse in a rural environment, it could be

deduced that the individual was killed inside a building and then transported away from

the urban environment.

• Whether the individual was killed during the day or the night (Nuorteva 1977, Smith

1986 and Benecke 1996). Species of insects have distinct diurnal or nocturnal activity

patterns.

• The presence of antemortem drugs (Catts and Goff 1992b) by conducting tests upon

larvae.

• The possible effect of insect activity on mimicking sexual assault clothing patterns

(Komar and Beattie 1998).

• Patterns of insect succession in wildlife death investigations (Dillon 1997).

• Evidence of child abuse or neglect (Goff et.al. 1991 and Lord 1990).

• Evidence of insect interaction with a crime suspect (Webb et.al. 1983 and Prichard

et.al. 1986). Suspect may have insect bites or insects on their person which are consistent

with entomological evidence found at the crime scene.

5

Erzinclioglu (1985) stated that each forensic case must be treated as unique and

that no basic rules of forensic entomology can be formulated. Yet, baseline data are

required in each biogeoclimatic zone if forensic entomology cases are to produce positive

and useful information. The intention of this study is to confirm baseline data previously

collected in the northern region of the Rocky Mountains in a similar biogeoclimatic zone

(Dillon 1997). If the fauna are similar, a forensic investigator maybe able to extrapolate

Dillon's data to forensic cases in a similar biogeoclimatic zone in Montana and vice

versa. A biogeoclimatic zone may be defined as a geographic region which has a

distinctive climate and supports organisms specific to that region when compared with

bordering regions or zones.

Data from this study Will also be compared with the only other "bum study"

published in forensic entomology to this date (Avila and Goff 1998). The burned

specimen is meant to simulate an accidental death or homicide victim burned with an

accelerant. The standards for the extent of the bum have been set by Glassman and Crow

(1996) who divide the identification of fire injury to an individual into five levels. The

final aim of the project is to complement the only existing forensic entomological study

conducted in Montana, which was performed in the same biogeoclimatic zone at a lower

elevation in a mixed urban/rural setting (Temeny 1997). This will provide the forensic

investigator with a species diversity index indicating the abundance of urban and rural

species in the interior Douglas-fir/ponderosa pine biogeoclimatic zone of Montana.

Results of this study may also be valuable in determining if the decedent was physically

moved after death and possibly by whom. These three hypothesis may be stated as:

6

(1) The successional fauna of the pig model will be similar to that found by Dillon

(1997) who conducted well replicated forensic entomological studies in a similar

biogeoclimatic zone located in British Columbia, Canada.

(2) The burned and unbumed specimens will have different successional fauna

specific to their physical states.

(3) The insect successional pattern and species diversity on carcasses present in a

mixed urban/rural and rural area will be distinct (Temeny 1997).

It has been established that pigs, being omnivores, have similar gut fauna as well as

roughly an equal amount of body hair as humans. Accordingly, the use of pigs as

surrogate humans in decay studies is well established (Payne 1965, Payne et. al. 1968,

Payne and King 1970 and 1972, Smith 1986, Tullis and Goff 1987, Haskell 1990,

Hewadikaram and Goff 1991, Shean et. al. 1993, Dillon 1997, Temeny 1997). Therefore,

the animal model chosen for this experiment was the common pig, Sos scrofa.

While pigs are viewed as suitable animals for forensic studies, the size of a pig can

have important effects on insect colonization. Previous experiments have used animal

models ranging from a small size of approximately 25 kilograms or less (Hackman 1963,

Comaby 1974, Blackith and Blackith 1989 and Patrican and Vaidyanathan 1995), a

medium size of approximately 50 kilograms (Fuller 1934, Bomemissza 1957, Reed 1958,

Burger 1965, Johnson 1975, Jiron and Cartin 1981, Early and Goff 1986, Tantawi 1996,

LeClercq 1997), to a relatively large animal model, such as an elephant (Coe 1978 and

Braack 1986). Arguments for or against a particular size commonly revolve around

decay rate as opposed to insect succession or species.

7

In a study pertaining to effects of specimen size, Hewadikaram and Goff (1991)

concluded that successional patterns and developmental rates of insects were not

dependent upon specimen size. Their largest pig was twice the size of the smallest and

attracted a greater number of taxa, with a marked difference between days 5-16 of the

study. This attraction of a greater number of taxa is probably related to increased surface

area and mass available for oviposition. The authors found no difference between carcass

size and the rate of development for arthropods, which are used for determining the PMI.

They deduced that arthropod succession patterns and developmental rates are

independent of the size of the animal model. However, Komar and Beattie (1998) used

pigs ranging in size from 19-162 kilograms and determined that larger carcasses attract a

greater number of arthropods and support larger maggot masses than do small carcasses.

Therefore, while development and succession may not differ in pig carcasses of differing

size, relative abundance of insects may.

8

Chapter 2: Materials and Methods

Study Area

The study site was located in northwestern Montana (46’53'N, 113'27'E) at the

Lubrecht Experimental Forest (LEF) in Missoula County. Lubrecht is approximately 30

miles (53 kilometers) east of Missoula, MT. The forest comprises 28,000 acres (10,927

ha) of zoned and managed areas with uses ranging from recreational to resource

conservation and range management. The study site is found within an area zoned for

grazing, hunting and nature study (Maus, personal communication). It is in range and

township of R15W T13N on the northern border of section 15. The site is located at the

end of an unimproved secondary road, located 1 mile southwest of the Lubrecht

Experimental Forest headquarters off Highway 200. The road proceeds northwest of

Highway 200 for approximately 2 miles. The elevation of the study site is nearly 4,400

feet and is located on the southward facing slope.

Vegetation consists predominantly of coniferous species in low to middle

elevation categories (Hitchcock and Cronquist 1973; Eyre 1980). The primary forest

cover is a mix of ponderosa pine (Pinus ponderosa) and Douglas-fir (Pseudotsuga

menziesii). Understory vegetation consists of snowberry (Symphoricarpos duhamel),

ninebark (Physocarpus maxim.), various grasses, and other vegetation common to these

stands.

Animal Cadavers

Two pigs were used in this study and therefore observations were not replicated.

Replication is preferable in experiments of this nature. Therefore, this project should be

9

viewed more as a long-term case study with the intention of gathering baseline data for

the biogeoclimatic zone of northwestern Montana.

Two mature 180-pound (81.65-kilogram) pigs were selected for the animal model in

this study. Each was killed with a single gunshot to the head, producing only a small

entrance wound. Both pigs were placed in separate bear proof cages measuring 150

centimeters in length by 90 centimeters in height by 90 centimeters in width and

constructed with re-bar steel. The control pig (CSS) and cage were placed 30 meters

from the pig to be burned (BSS).

Once in the cage, one pig was sprinkled with 3.8 liters (1 gallon) of gasoline and set

afire. After the pig had been allowed to bum for approximately 30 minutes, flames began

to wane and small cracks appeared upon the skin surface with a small amount of charring

(Figure 1). Once the flames dissipated, the level of bum apparent on the pig was

evaluated using the Crow-Glassman scale (Glassman and Crow 1996).

Climatology

The experiment began on June 17, 1999. Temperatures were measured using a hand­

held thermometer at the following locations: internal temperature (approximately 3

inches under the skin), the skin surface, the ground surface and between the specimen and

the ground surface in the area of the abdomen. Measurements were taken twice a day,

once in mid-morning and once again in mid-afternoon for the first 30 days of

decomposition, then similarly every two days for a period of approximately 60 days, and

finally one day a week until December 1st, 1999, when travel to the site was no longer

possible due to snow. Ambient local site temperatures and humidity were recorded

continuously from June 17,1999 to December 1,1999 using a hygrothermograph.

10

Rainfall was recorded using a standard rain-gauge located at the site and monitored at

the same time as the hand-held readings discussed above. Local cloud cover conditions

were noted visually. All ambient temperature and rain measurements were compared

with information from the nearest National Weather Service (NWS) station located at the

Missoula County Airport in Missoula, MT.

Decomposition Stages

The framework selected for this experiment was designed by Early and Goff

(1986) and reviewed by Goff in 1993. The classification system used consists of five

stages, which indicate arthropod successional patterns, chemical decay processes, and

physical transformations in the carcass or corpse. Transitions from one stage to the next

are continuous. The five stages are summarized briefly as follows:

1. Fresh Stage

This stage begins at death and ends when bloating is visible. The first arthropods

to colonize the specimen usually consist of the families Calliphoridae (blowflies) and

Sarcophagidae (flesh flies). Abundance and species diversity during this stage is

dependent upon the biogeoclimatic zone and a host of environmental variables. Flies

present at this stage will deposit eggs or larvae on natural orifices or open wounds of

the specimen.

2. Bloated Stage

At this stage, anaerobic bacterial action produces gasses that inflate the carcass.

The feeding activity of dipteran larvae and putrefaction increase the carcasses

temperature. The combination of carcass fluid seepage-and the ammonia of dipteran

11

larval activity cause the soil under, and immediately surrounding, the carcass to

become more alkaline, thus changing the faunal content of the soil.

3. Decay Stage

Gasses begin to be released and the carcass ceases to be bloated. The large

number of larvae present increase the presence of predaceous arthropods. The end of

this stage is indicated when the majority of dipteran larvae have migrated away and

the carcass has been mostly consumed.

4. Post-Decay Stage

This stage finds a transition of the carcass into primarily skin, connective tissue,

and bones. Coleoptera as well as predatory and parasitic species increase in number.

5. Remains Stage

The only portions of the carcass remaining at this stage are bones and hair. Soil

samples indicate a return to normal fauna. The length of this stage is extremely variable

depending upon the biogeoclimatic zone. Times may vary from years in an extremely

dry environment to months in a tropical environment with high scavenging activity.

Insect Sampling

Insects were collected only from the external surface of the pigs to avoid altering

conditions of the cadavers. Detailed notes were taken of where the insects were located

on the body according to procedures outlined by Lord and Burger (1983) and Borror et.

al. (1989). Photographs were taken each day to illustrate locations of insects and the

status of the carcasses. Samples were collected from each carcass daily during the first

30 days of the experiment. Once into the decay stage, samples were collected once every

3 days until the post decay stage was apparent. Sampling was decreased to once every 7

12

days when the post decay stage occurred. This rate of sampling continued until snow and

low temperatures hindered arthropod activity as well as collection procedures.

Collection procedures for each decomposition stage were limited to the mode of

locomotion of the arthropods present. During the bloated and decay stages many fly

larvae and adults were collected. The procedures for the collection and storage of these

different stages of Diptera development are distinct. Furthermore, the collection

techniques are also different depending upon the size of the Diptera being captured.

These procedures and techniques are detailed below.

Insects: Collection Techniques

Adult Insects

This experiment used procedures basic for both ground-dwelling as well as flying

specimens. Adult ground-dwelling insects were collected in two ways. Pitfall traps were

used to capture adults as well as migrating larvae. Each pitfall trap consisted of a

medium sized glass jar approximately 15 centimeters (6 inches) tall and 7.5 centimeters

(3 inches) wide at the mouth. A piece of metal screening (.625 centimeter or 1/4 inch

mesh) material measuring 2.5 centimeters by 2.5 centimeters (1 inch by 1 inch) was fitted

over the mouth of the jar. Soapy water was placed in the bottom of the jar to ensure

insects could not escape. A single pitfall trap was placed approximately 30 centimeters

(1 foot) from each carcass inside the cage to prevent small mammals being captured or

injured. Forceps and fingers were also used to capture many ground-dwelling insects.

A net was utilized for large flying insects such as Calliphoridae and

Sarcophagidae. Two-dozen sweeps were made each day over each cadaver to capture a

representative sample of insects. An aspirator was used for smaller specimens such as

13

Phoridae and Piophilidae. Captured specimens were killed in ethyl acetate killing jars

and then transferred to vials containing 70% ethyl alcohol. Labels for vials included the

site name, carcass type, county, state, date, time, sample number, and location of insect.

Larval Insects

Collection of larvae was accomplished using forceps. Samples were taken every

day larvae were present, with a small portion of each sample saved for rearing purposes.

Portions not used for rearing were placed in vials containing Pampel's solution, a

preservative that helps retain the larvae's size and natural coloration (Smith 1986).

Larvae migrating from the carcasses were collected and labeled separately from those

collected on the cadavers with migratory distances noted. Migration routes and distances

are important in finding specimens at a death scene, therefore, empty pupal cases were

also collected from near the cadavers.

Pupal Collections

The use of pupae in the forensic sciences has not been extensive (Malloch 1917,

Gilbert and Bass 1967, Okely 1974, Chu and Wang 1975 and Nuorteva 1987). Several

authors have published repports indicating that the age determination of pupae is

significant to forensic investigations (Reiter and Wollenek 1983, Nuorteva 1987,

Greenberg 1988 and Catts 1991). Once the age of the pupae has been established, it

lends credence to the estimation of the PMI. Therefore, the discovery and collection of

pupae is an important part of the overall forensic entomological data gathered at a crime

scene. Archaeological methods were employed here in the hope that using a specific

technique of sampling and excavation would yield better results than using non­

archaeology techniques.

14

The migration of larvae from a carcass occurs at varying times and ways

depending upon the insect taxon and stage. The last larval stage often pupates in the soil

directly under or adjacent to their food source. However, some larvae often "wander"

(between 3 and 10 meters) from a carcass to avoid competition and to seek dry soil for

pupation (Smith 1986). After emergence, the empty pupal case remains.

Haskell and Williams (1990) found that samples should be collected up to one

meter from the body for good coverage. If maggots have left the corpse it is important to

track down and capture puparia, inactive larvae, and prepupae. Smith (1986)

reccommends making shallow 7 centimeter (3 inch) transects in several compass

directions of up to 20 feet from the corpse.

Puparia were sampled using systematic sampling (Renfrew and Bahn 1991). A

datum point was selected and a grid system formed with respect to an N-S/E-W axis.

The grid system consisted of one meter plots. Alternating squares were chosen for

sampling. A 50 centimeter2 test unit was excavated in each of the alternate plots. A large

number of the plots were located in rough terrain consisting of large stones and extremely

rocky soil. Small samples from each 50 centimeter test unit were screened through 1/8

inch mesh and collected at the site. Larger samples were bagged and returned to the

laboratory for screening. Rocks were overturned at random in the selected one meter

squares to search for puparia.

Excavations were also conducted under the cages. On December 1st, the final day

of the experiment, the cage containing the burned specimen was moved and a one meter

excavation unit was established where the cage had been located. Using standard

15

archaeological techniques (Renfrew and Bahn 1991), the sod layer and subsoil were

removed, screened, and the insect associated materials collected for laboratory analysis.

Insects: Identification

Insects were identified in the laboratory using standard keys such as Hall (1947),

Cole (1969), Borror and White (1970), Greenberg (1973), Oldroyd and Smith (1973),

Anderson and Peck (1985), and Borror et al (1989). Adult insects were then pinned

while larvae were kept in Pampel's solution.

16

Results: Chapter 3

The insect succession results for this experiment are summarized below by the

stage in which they occurred. Each decay stage is divided. The commencement and

termination of the remains stage was the most variable and was determined by the

relative amount of flesh remaining upon the pig. Species diversity of insects utilizing

each of the stages are shown in Figures 7-11 for Diptera and Figures 12-15 for

Coleoptera. Insects present during each stage for each pig are listed in Tables 1 and 2.

The level of burning obtained for BSS was consistent with a Crow-Glassman Scale

(1996) (CGS) Level #1 bum with some elements of a CGS #2 bum. A level #1 bum has

the characteristics of epidermal blistering and singed hair, typical of a smoke-inhalation

death. A level #2 bum displays a degrees of charring. The bum results from the present

study are in slight contrast to Avila and Goff (1998) who obtained only a level #2 bum

only. Once the bum was completed, the smell of roasted skin was readily apparent,

unfortunately providing an attractant to large carnivores such as bears and scavenging

birds.

Fresh Stage (day 1)

Odor associated with this stage was consistent with that of fecal matter and urine

as the subjects bowels discharged upon death. Anthomyiidae, Otitidae, Phoridae,

Fannidae, and Cynomyopsis cadaverina (Robineau-Desvoidy) (Family Calliphoridae)

were present and are typically attracted to fecal matter (Borror and White 1970). More

than one inch of rain fell during day one. Flying insects in general were not visible

during rainfall but increased markedly once the sun shone. Any eggs laid were likely

washed away during the intermittent rains as none were present in the evening.

17

Adults of Phormia regina (Meigen) and a species of Sarcophaga were the most

abundant decomposers sampled on the pigs during this stage, with adults of Eucalliphora

(Townsend) and Lucilia illustris (Meigen) contributing roughly an equal ratio. Flies of

the species Protophormia terraenovae (Robineau-Desvoidy) were also present in low

numbers at the burnt carcass while Anthomyiidae were present only at the control

carcass. Additional Diptera present, albeit contributing very few adults to the whole

community, were Phaenecia cadaverina, several species of Fannia, and members of the

families Tachinidae, Heleomyzidae, and Phoridae. See Tables 1 and 2 for records of

Diptera present or absent on each respective carcass. The orders Hymenoptera and

Lepidoptera were observed flying around the pigs. Members of the families

Megachilidae and Vespidae were observed in the area but did not interact directly with

the carcasses. The bulk of the fauna collected were present on both carcasses.

A slight difference in carcass color was apparent on the abdomen of the burnt

carcass, which exhibited a large turquoise green di scoloration and lack of hair on the

singed skin. In the evening, this discolored skin began to split and crack.

Bloated Stage (days 2,3-8)

The bloated stage commenced as gasses began to inflate the carcasses after day

two. This stage began about a day earlier, and ended roughly a day sooner, for the burnt

carcass possibly due to cracks in the skin assisting in carcass deflation. The axial portion

of the burnt carcass became extremely dry and firm following the bum and displayed the

least amount of insect activity. Unidentified eggs were present in the early days of this

stage. Larvae of Ph. regina, the only immatures found during this stage, were present by

18

day five. Eggs and larvae were encountered in the mouth, ears, eyes and open body

cavity (at the end of the bloat).

Adults of Eucalliphora and Calliphora terraenovae were present in low numbers

while adults of Prot. terraenovae, two species of Sarcophaga and Fannia (burnt carcass)

were slightly more abundant. As bloating proceeded, Piophilidae adults of the species

Prochyliza brericomis (Melander) were attracted to each carcass. Sepsids, Sepsis spp.

and Meroplius Stercoronius (Robineau-Desvoidy), were also present on the burnt carcass.

Individuals of the genus Nemapoda were especially attracted to the control, albeit in low

numbers. A low number of Heleomyzids continued to show attraction to the control.

Coleoptera associated with the bloated stage were diverse. The burned skin on

the burned pig cracked and a small amount of fluid seeped from various cracks in the

abdomen and thorax. Members of the species Thanatophilus lapponicus (Herbst) were

active at these cracks in the skin and may have been feeding upon Diptera eggs. T.

lapponicus present at the control carcass were active on skin which began sloughing off

toward the ground at the end o f the bloat stage. Seven to eight days after death they were

seen mating on the burnt carcass.

High numbers of the genus Dermestes were found on the burnt carcass throughout

the bloat period and were observed moving on the burnt carcass through the nose and

mouth. Dermestes appeared to be attracted to fluid seepage from the nose and mouth of

both carcasses. Dermestes were found in slightly lower numbers on the control.

Coleoptera present in moderate numbers included the families Histeridae and

Staphylinidae. Numbers of Chrysomelidae, Tenebrionidae, Elateridae (species 4), and

Scarabaeidae, subfamily Aphodiinae were low and nearly insignificant on the burnt

19

carcass. Of these, only Chrysomelidae and Scolytidae were present on the control.

Overall, more species of Coleoptera were associated with the burnt carcass. However,

Coleoptera were found in higher numbers on the control. The only Hymenoptera found

on the control carcass were from the family Formicidae. Hymenoptera collected from the

burnt carcass included Formicidae, Megachilidae (subfamily Megachilinae), and

Vespidae (subfamily Vespinae).

Red mites (order Acari) were first observed during this stage and were present on

both carcasses for the remainder of the study. The mites may be phoretic on Histeridae

as they were often observed on hister beetles as well as moving over the carcass.

Springett (1968) found that the mite Poecilochirus necrophori may have a symbiotic

relationship with the beetle Necrophorus. Mites used Necrophorus adults as transport to

a carcass and fed upon dipteran larvae alongside Necrophorus. By aiding in reduction of

Dipteran larvae, Necrophorus was more successful in breeding. In experiments where

mites were absent, Necrophorus was entirely unsuccessful in reproducing. The mites

were not observed to directly feed upon Dipterous larvae in the present study but may

have a similar relationship with hister beetles.

Decay Stage (days 9-13)

Diptera associated with this stage were predominantly larval Ph. regina (Tables 1

and 2). Prot. terraenovae were present in very low numbers on both carcasses.

Sarcophaga (species 6) and Piophilidae (species 2) were found on the control carcass. In

comparison, Fannia (species 1) was present only upon the burnt carcass.

Day 9 was marked by a very low temperature of 3°C. At this time adult Diptera

activity was extremely low and the majority of the larvae present were in the body cavity

20

of the both pigs, generating enough heat to stay alive (Figure 2). Previous research has

shown that larval movement frequently raises internal temperatures above the ambient

temperature (Payne 1965, Williams and Richardson 1984, Early and Goff 1987, Catts and

Goff 1992). The low number of larvae on the burnt carcass enabled only a small number

to survive as the temperature approached freezing. The temperature rose on days 11 and

12 by 8°C and fresh eggs were present on both carcasses. Formicidae were observed

carrying off Diptera eggs while Staphylinidae preyed upon both Diptera eggs and larvae.

Histerids were observed preying upon both Dermestes and Diptera larvae underneath and

inside the orifices of the head.

As larvae consumed the organs and muscle tissue of each carcass, the skin

covering the abdomen became dry and provided a cover above the feeding larvae. Under

this cover on the burnt carcass on day 11, mating Nicrophorus and Necrodes

surinamensis (Fabricius) (family Coleoptera) were observed walking upon the frothy

maggot mass in the burnt carcass. At the end of the decay stage, larvae of T. lapponicus

and adults of Histeridae and Dermestidae continued to be fairly common on both

carcasses. N. surinamensis adults and larvae were only found on the burnt carcass at this

stage. Staphylinidae and Histeridae continued to prey upon larvae on both carcasses.

Hymenoptera present throughout the decay stage include Vespidae (subfamily Vespinae)

and Formicidae.

Post-Decay Stage (days 14-37)

The appearance of each carcass was modified by the feeding and migration of

larvae. The once fleshy abdomen became devoid of insects with the exception of

Coleoptera and various Hymenoptera. Most skin had fallen away from the thoracic

21

cavity and abdomen revealing the ribs and vertebrae. The skin of the burned carcass was

stiff in areas but brittle in others. A maggot mass was observed in the abdomen of each

carcass and exhibited a bubbly frothy mixture of larvae at the height of larval abundance

and internal temperature readings. Small maggot masses were observed moving in the

mouth, nose, ears, and eyes following the migration of the majority of larvae from the

abdomen. Larvae consumed nearly all muscle tissue along the axial skeleton, shoulder

joints, and pelvis of each carcass (Figures 3 & 4). Coleoptera often remained beneath

each carcass and were only observed when the carcass was lifted.

At the commencement of this stage, Diptera larvae exited the carcasses after the

majority of the flesh had been removed. Migrating larvae of Ph. regina and L. illustris

exited each carcass beginning on day 13 for the control carcass and day 14 for the burnt

carcass, often in full sunlight at midday. Eucalliphora adults were most abundant during

this stage on the burnt carcass and were exceeded in numbers only by adults of Ph.

regina. Larval migration was nearly complete by day 32 and finished by day 37 on each

carcass. Genera present in moderate numbers on the burnt carcass were Eucalliphora and

Fannia (Species 1). Various Sepsidae were collected at each carcass and are listed in

Tables 1 and 2. One mating pair from the genus Nemapoda was captured on July 9th.

The Dipterans, Phaenecia sericata (Meigen), Phaenecia caeruleiviriais

(Macquart), Fannia canicularis (Linnaeus), F. difficilus (Stein), Fannia (species 3),

Piophilidae (species 1), made their first and only appearance during this stage at their

respective carcasses as shown in Tables 1 and 2. Additional Diptera adults attendant for

a brief time are all remaining families listed in Tables 1 and 2.

22

The most abundant adult species during this stage were the Piophilidae species

Proch. brericomis and Proch. xanthostoma (Walker). Mating pairs of Proch.

xanthostoma were captured from day 17 until day 27. Proch. brericomis mating pairs

were captured from day 29 until day 111, in the period directly following Proch.

xanthostoma.

Silphidae larvae of the species T. lapponicus and N. surinamensis increased in

abundance during the post-decay stage. Both species moved through the Diptera maggot

mass but appeared to be most interested in the dried skin of the carcasses. However, after

extended observation, both species were observed preying and fighting over dipteran

larvae. Adults of I lapponicus, Nicrophorus, Dermestidae, Histeridae, and

Chrysomelidae were observed on each carcass with only slight changes in population

numbers since the decay stage.

T. lapponicus were also observed preying upon Dipteran larvae. By day 28, a

copious amount of empty Coleopteran larval skins could be found in and surrounding

both carcasses. Coleopteran families such as Staphylinidae, Coccinellidae, Curculionidae

(subfamily Brachyrhininae), Salpingidae, Scarabaeidae (subfamily Scarabaeinae),

and Elateridae (species 1 and 2), were present in diminished numbers on their respective

carcasses.

Of the numerous Hymenoptera collected, only the families Formicidae, Sphecidae

(subfamily Sphecini), Apidae (subfamily Apinae tribe: Bombini), and Vespidae

(subfamily Vespinae) displayed an active interest in the carcasses. Specidae and

Formicidae were numerous throughout this stage. Formicidae in particular w,ere

observed preying upon migrating larvae more often than upon feeding larvae. At the

23

height of larval migration, virtually every ant visible was carrying either Dipteran or

Coleopteran larvae.

Upon the completion of larval migration, the numbers of ants observed near the

carcasses dwindled. Apidae (subfamily Apinae tribe: Bombini) exhibited aggressive

behavior on various Coleoptera, namely Dermestidae and Histeridae. Araneida

(subfamily Araneae) also predated upon both adult and immature Diptera.

Remains (days 38+)

At the onset of this stage only bone, cartilage, dried skin, and a small amount of

tissue was left on the carcasses (Figures 5 & 6). The rotten smell previously pervading

the site became identifiable only on wet or unusually warm days.

Ph. regina continued to be the primary calliphorid found on the remains, albeit in

small numbers. Prot. terraenovae was found on the burnt carcass in extremely low

numbers and constitutes the only other calliphorid present. Sarcophagidae and Fannia

were now more abundant than Calliphoridae. Four species of Sarcophagidae were found

until the end of September with a few teneral adults still collected on day 60. Members

of various Fannia spp. were also found in relatively moderate numbers throughout the

end of September. Newly emerged flies were collected on day 56 on the burnt carcass.

F. canicularis was found sporadically throughout July and August on both carcasses.

Two species of Piophilidae also were active during this stage. Specimens of

Proch. xanthostoma were found mating in great numbers in mid-July, after which their

numbers tapered off. In early October, their numbers increased a second time, and

thereafter adults were not found again. Specimens of Proch. brericomis were found in

moderate numbers from mid-July until early October. They were observed mating and

24

collected as late as day 105 (early October) on the burnt carcass. Piophilidae larvae were

collected from the control carcass between days 56-83. Unfortunately, they were not

identified to species and it is unclear whether they are Proch. xanthostoma or Proch.

brericomis. Each of these Piophilidae species were vital to the analysis of Diptera

activity during the remains stage as the adults were the principal species observed mating.

The latest seasonal observations of mating were of Heleomyzidae. Mating was

observed to occur from day 116 to day 138 (ie early October to early November) on both

carcasses. Immatures were not observed. Diptera present on both carcasses in low

numbers were Sepsidae (Sepsis spp.) and Tachinidae spp (Tables 1 and 2).

Coleopteran larvae of T. lapponicus and N. surinamensis were essentially absent

during this stage. The final larval stages for these two species were collected in early

August at the same time Dermestes larvae became numerous. Larvae of T. lapponicus

were present on each carcass for approximately 35 days. Anderson (1982) has

commented that the completion of larval stages of T. lapponicus lasts between 25-29

days. The departure of T. lapponicus in the beginning of August is not unexpected as

Anderson and Peck (1985) have remarked that this species decreases in numbers later in

the year and overwinters.

By day 46, Dermestes larvae were abundant underneath the carcasses and inside

the body cavity where the skin had dried. One such area consisted of the carcasses' dried

ears, which had been cleared of flesh, and some cartilage, leaving only dried skin husks.

Dermestes larvae were collected until day 67 for the control and day 70 for-the burnt

carcass. Thereafter, in reduced numbers, only adult Dermestes were found until day 97.

25

The greatest number of species were found in the remains stage for each carcass.

A possible reason for this diversity is the long duration of the stage, which can range

from months to years, depending upon the environment. The rate at which the carcass

decays is determined more by vertebrate scavengers, weather, etc. than by insects, and

therefore the successional value of insects during this stage is limited. Adults of

Dermestes, T. lapponicus, Staphylinidae, and Histeridae were still present. Additional

Coleoptera associated with the remains stage for the control carcass but not for the burnt

carcass are: Scarabaeidae (subfamily Troginae) and Curculionidae (subfamily

Brachyrhininae). The remaining Coleoptera found were present throughout the study and

are listed in Tables 1 and 2.

Archaeology

All insect pupal evidence was found in the topsoil 10-30 centimeters below the

surface. Pupal cases were collected from only two 50 cm test units. Three pupal casings

were found farther than 3.3 meters from the carcasses. The farthest pupal casing was

found 17 meters from the datum point. No puparia were found under rocks overturned at

random in the selected one meter squares. In contrast, dozens of pupal casings were.•j

recovered from the one m excavation unit placed under the cage of the burned pig.

Vertebrate Carnivores and Scavengers

Damage to the carcasses was minimal on occasions when vertebrate carnivores

and scavengers attempted access to the pigs due to the strength of the rebar cages. On

day 2, bear droppings were found in the grass between the two cages and several pieces

of rebar on the control cage were bent outwards. The pig inside was cut open slightly in

the pelvic region, which enlarged a split in the skin already present due to the bloating

26

process. On day 6, more bear activity occurred on the west-facing side of the control

cage. Rebar was peeled back in several places but the integrity of the cage was intact and

the pig was not damaged.

Bear activity was absent throughout the decay and early post-decay stages but on

day 105, during the remains stage, damage occurred to both cages. The door of the cage

containing the control carcass was battered and the east-facing side of the burnt pig's cage

had rebar peeled back in one spot. Damage to the carcasses themselves was insignificant

and the study continued uninterrupted. The primary scavenger observed until the end of

the decay stage was the turkey vulture (Cathartes aura). Fortunately, the rebar was

welded in a close pattern and turkey vulture damage was kept to a minimum.

Larval Development

Dipteran Larvae

Ambient temperature averaged 22°C as a high and 7°C as the low on the day

when larvae were first observed. Thereafter maximum temperatures averaged 20.3°C and

minimum temperatures averaged 7°C until the first larval migration. The average

maximum temperature during the thirteen day migration period was 26°C and the

minimum low averaged 9°C. If the maggot mass is large enough and produces adequate

heat for development only extreme variances in ambient temperature should negatively

affect the larvae. The high ambient temperature during this two week post-peak internal

temperature migration period averaged 18.5 °C with an average low ambient temperature

of 6.5°C. These ambient temperatures were within a range that permitted an adequate

environment for larval growth (Kamal 1958). Rainfall recorded during this period at the

27

site was 2.81 inches. Dew was present every morning and the majority of the daytime

hours were partly cloudy.

When larvae first appeared on day 5 the average humidity was 86%. Days 6-13

had an average humidity of 74%. These are the eight days in which larvae developed

through the first and second instar stages. Day 13 was the first time larvae were observed

migrating from the control. Days 14-28 have an averge humidity of 72%. These are the

fourteen days during which larvae concluded their migration. During times of humidity

extremes, such as 96% and 20%, and temperature extremes, such as 3°C and 34°C, larvae

appeared to become scattered and disorganized. The low ambient temperature had less of

an effect at times when internal temperatures of the pigs were at a peak.

Control S. scrofa

By day 5, approximately one dozen 3rd instar and two dozen 1st and 2nd instar Ph.

Regina larvae were collected. Numbers increased through days 9 and 10. On days 13 and

14 all three larval instars were found on the carcass but large masses of 3rd instar Ph.

regina were predominant. On day 13, the first larval migration was observed, 12 days

after the first eggs were laid. Larvae could be found exiting the carcass as late as day 24.

By day 32, newly eclosed adults were found.

Internal temperatures peaked at 30°C on day 13, the day of the first larval

migration. Larvae continued to migrate as the internal temperature stabilized at an

average of 25°C over the next seven days. On day 20, the internal temperature peaked

again at 35°C conincident with a 3rd instar larval migration. A few scattered larvae

continued to migrate until day 32. Larval migrations from the control pig correspond

roughly with those from the burnt carcass. Migration of larvae from the control pig

28

began approximately a week before peak internal temperatures and continued for almost

two weeks. These differences are due to the higher internal peak temperature and greater

abundance of larvae in the burnt carcass

Burnt S. scrofa

By day 5, dozens of larvae were observed. Day 14 saw the first 3rd instar larval

migration, 13 days after the first eggs were laid. Another larval migration occurred on

day 17. The number of larvae increased steadily until day 20, when large masses of 1st,

2nd, and 3rd instar larvae were found. On days 22-24, additional 3rd instar larvae were

found migrating. However, 3rd instar larvae were still found on, and leaving the carcass,

on days 25 and 26. On day 28, there were virtually no dipteran larvae remaining on the

carcass. By day 32, newly eclosed adults were observed, indicating a minimum

immature development time of 31 days for Ph. regina in this environment. 3rd instar

larvae of L. illustris are found on July 4th and were observed migrating between July 9th

and 11th from both burnt and control carcasses. Larvae of Diptera other than L. illustris

and Ph. regina were not detected.

Internal temperatures for the burnt specimen began at 7.8°C and peaked on day 11

at 26.7°C. The internal temperature peaked again on day 21 at 31.1°C. These peak

internal readings only differed from the ambient temperature by 1-6°C. This peak

temperature corresponds with the migration occurring on days 14 to 17 and on days 22 to

26, Each migration lasted approximately four days. Dillon (1997) has stated that

calliphorid larval migration usually occurrs before the peak internal temperature has been

reached. In this study, migration, began approximately a week before the highest peak

internal temperature and continued for four days following the peak.

29

Coleopteran Larvae

Burnt and Control S. scrofa

Adults of the genus T. lapponicus were observed mating on days 6 and 8. Adults

of Nicrophorus and N. surinamensis were first observed mating on day 11. Coleopteran

larvae were also observed on this day but were not collected as the few present were

being preyed upon by Histeridae and Formicidae. N. surinamensis larvae of all sizes

were found from days 11 -48 with a few larvae up to day 63. Small larvae of T.

lapponicus first appeared between days 7-26. Medium and large larvae of T. lapponicus

were found between days 26-49. By day 28 in the burnt carcass and day 26 in the control

carcass hundreds of cast off Coleopterous larval skins were found inside, under, and

surrounding the carcasses.

Adults of the genus Dermestes were found from days 2-91. Immatures were

found from days 46-70. Dermestes immatures were found two days before immatures of

N. surinamensis and T. lapponicus made their last appearance. This abrupt change from

the high abundance of immatures of the latter two species to the high abundance of

Dermestes could be explained by the habits of each species. The immature

developmental stages for Dermestes are not described here as individuals in this genus

were not identified to species.

Temperatures under 10°C and over 35°C appeared to have the effect of lowering

activity for coleopteran larvae as it did for dipteran larvae. In both carcasses, many

coleopteran larvae were not only predaceous upon dipteran larvae, but upon newly

eclosed flies as well.

30

Ratcliffe (1972) studied N. surinamensis in Nebraska. Adults were nocturnal.

Eggs were laid around large carcasses and hatched and began to feed 2-4 days later.

Anderson and Peck (1985) observed a total immature time of approximately 42-43 days.

These statistics correspond well with the data collected for the present study where the

relatively few adults were collected in the early morning and evening and the maximum

period of larval development was approximately 37 days.

The immature stage of T. lapponicus has been reported by Anderson (1982) as

having two generations a year in Ontario, the first generation beginning in April-May and

the second from early June-July. Complete development time was estimated at 32-33

days for each generation. The rate of development for T. lapponicus in the present study

was approximately 23 days. Anderson (1982) estimated the egg stage to last 5-6 days.

Eggs were not identified, and therefore, were not included in the 23 day estimate. The

immature stage for this insect in the present study may have lasted between 28-29 days.

The results found during this study are similar to the habits and natural history

recorded for these T. lapponicus and N. surinamensis in the literature. Anderson and

Peck (1985) state that T. lapponicus is a cold-weather species that occurs at high

mountain elevations in the west. The decrease in adults as the fall season progressed also

concurs with the data of Anderson and Peck (1985) for the overwintering habits of this

species.

The life cycle of flies is most commonly used to determine the Post Mortem

Interval (PMI) (Nuorteva 1977). However, with information on the immature stages of

coleopteran species, it may be possible to use Coleoptera in conjunction with dipteran

species for stronger PMI estimates.

31

Dillon (1997) found adults of both N. surinamensis and T. lapponicus. However,

no immatures were collected. This may be explained by 2 factors: (1) breeding

conditions may not have been optimal, and therefore, either breeding was not possible or

the immatures present were preyed upon or went unnoticed, and (2) these species could

have been outcompeted by D. talpinus, present in Dillon's study. In Dillon's sun habitat,

D. talpinus immatures and adults were found in both the decay and post-decay stages.

However, D. talpinus immatures were found only in the bloat stage for the shade habitat.

Therefore, competition between these Coleopteran species may only provide a partial

answer why differences in decomposition time was observed. The importance of season,

temperature, humidity, and the amount of food resources available, may be significant

factors in the ability of some immatures to survive to adults.

32

Discussion: Chapter 4

Kamal (1958) found Ph. regina have a pupal stage duration of 4 to 9 days and a

complete immature stage (egg to adult) of 10 to 12 days under laboratory conditions of

26+l°C and 50+2% relative humidity. Greenberg's (1991) study, conducted at 22°C,

found a pupal stage length of 5 days with approximately 13 days for total development

from egg to adult. The developmental averages in days presented by Kamal (1958) and

Greenberg (1991) are significantly less than those determined for this study. The

temperature range provided by Greenberg (1991) is close to that of the present study.

The 50+2% relative humidity level used by Kamal (1958) is the optimal humidity found

for Ph. regina larval development.

The early decay stages of fresh, bloat, decay, and post-decay are determined by

the activity and rate of development of Diptera larvae (Early and Goff 1986). This

provides an excellent opportunity for comparison of data in the present study and data

collected by Dillon (1997). In the present study the burnt and control carcasses had

nearly identical decay stage lengths, i.e., the onset of each decay stage varied only by

approximately one day. Decay stage length was dependent upon factors such as insect

predators and the lack of maggot mass cohesion on day 9 for the burnt carcass. The lack

of cohesion may be related to the low ambient temperature and insufficient numbers of

larvae present for creating optimal internal temperatures, thus, larval development was

delayed. Not only did the carcasses in this study have nearly identical decay rates (+ 1

day) but they also mirrored the decay rates found by Dillon (1997).

When broken down numerically Dillon's (1997) table of decay stages maybe

evaluated and approximately compared with the present studies numbers (Table 3).

33

While Dillon's experiment contrasted succession in pigs in sun and shade, the present

study contrasted burned and unbumed pigs in a partially shady site. However, these

differences are irrelevant because Dillon found no difference between the types of blow

fly species in shade vs. sun. Therefore, in order to compare her results with those of the

present study the decay stage length in days for Dillon's sun/shade habitats were averaged

in Table 3 with the results of the present study.

Some of the carcasses used in Dillon's study (1997) were clothed in various

human apparel (shirts, etc.). Following the assumption that clothed carcasses retained

fluids longer and thus lengthened the stage in question (Dillon 1997). However, once the

post-decay stage is reached, it is extremely difficult to determine as to when it concludes.

Therefore, this comparison of the length of the post-decay stage is only an approximation

and rate of development of the insects involved should be used as the actual determinant

of the length of a decay stage. With this in mind, the most abundant colonizing Diptera

should be used to model insect succession.

The observed developmental rate of 31 days for Ph. regina does not take into

account possible effects of competition which may have had a negative effect. A

predatory environment combined with low nightly temperatures and the possibility of

inadequate moisture all may have contributed to the lengthened larval developmental

period. There was an abundance of fly egg predators present during the bloat and decay

stages during which unidentified eggs were deposited. These included staphilinids,

silphids, histerids, and Hymenoptera. Putman (1978) found that predation rates of

immature insects of up to 66% may be reached in some instances. Nuorteva (1970) has

also commented on the near extermination of Diptera eggs and larvae by histerid beetles.

34

Previous experiments have found that high numbers of predators and low average

temperatures can lengthen the decomposition progress (Johnson 1975, Nuorteva 1977,

Payne 1965 and Reed 1958).

Temperature and humidity were the principal factors affecting larval

development. Cloud cover also significantly lowered adult Diptera activity. The use of

the interior Douglas-fir/ponderosa pine mixed habitat provided a good location to

contrast with that of Dillon (1997). While Dillon (1997) found eggs on day 1, in the

present study the lack of Calliphoridae eggs on day 1 was probably the result of heavy

rains. As in the present study, Dillon (1997) also found Diptera eggs in the fresh and

bloated stages but not in the decay stage. Maggot masses were found between days 5-12

for both the burnt and control carcasses in the present study. This agrees with Dillon's

results of 5-10 days for a similar interior Douglas-fir habitat.

The rate of development for Ph. regina was nearly identical on each carcass with

the first migration of 3rd instar larvae occurring only a day apart. This pattern is followed

closely by L. illustris. This is in contrast to findings by Dillon (1997), where Ph. regina

and Prot. terraenovae were observed to be co-dominant on carrion where L. illustris was

not. Ph. regina larvae of 1st, 2nd, and 3rd instars are found in the present study during the

bloat, decay, and post-decay stages. In contrast Dillon (1997) found Calliphoridae 1st

instars only in the bloat stage (sun) and fresh and bloat stages (shade), 2nd instars during

bloat and decay stages (sun) and fresh and bloat stages (shade) and 3 rd instars during the

bloat, decay, and post-decay stages in both sun and shade carcasses. Adul ts'of Ph. regina

were present throughout both studies until the remains stage was reached, after which

they were absent except in a few instances.

35

The presence of Ph. regina 1st and 2nd instars during later stages in the present

study indicates that eggs continued to be deposited well into the bloat stage and possibly

the decay stage. Each carcass, therefore, continued to provide a suitable substrate for the

deposition of eggs and their subsequent development into viable larvae. An important

point noted by Kamal (1958) is that Ph. regina is not as susceptible to crowding, low

humidity, etc. as the C. terraenovae (Macquart) and Phaenecia cadaverina that were also

present. This adaptation may have aided Ph. regina to achieve its high level of

abundance over the latter two species.

Diptera

The succession of insects in the two pigs is summarized in Tables 1 and 2.

Species diversity was recorded in a logarithmic table for the burnt and control carcasses

separately (Figures 7-15). While some species were absent in one or the other carcass,

they probably made little contribution to the decomposition process. Among the

Calliphoridae only Ph. regina made a significant contribution to this process. Ph. regina

adults and immatures were most abundant among the order Diptera throughout the study

until the post-decay stage when they shared high levels of abundance with L. illustris. It

was also during the post-decay stage that Proch. brericomis and Proch. xanthostoma of

the family Piophilidae and at least one species of Sepsidae were highly abundant.

Heleomyzidae were found mating in the remains stage.

The most abundant necrophagous species in this study was Ph. regina. The

abundance of this blow fly at the site is not surprising considering its habits. It is a

predominantly Rolarctic and rural species that has a seasonal appearance from the spring

through the fall (Hall 1947, Greenberg 1973 and Anderson 1995) and is common at high

36

altitudes during these seasons. Kamal (1958) found that this blow fly was less sensitive

to crowding in a controlled environment than C. terraenovae and Cy. cadaverina. Each

of these species was present in small numbers in the present study, however, crowding

may have influenced their relative abundance.

Cy. cadaverina is found from northern Labrador to the southern US border,

however, it is most abundant along the Canadian-US border (Greenberg 1973). Although

it has reproductive peaks in early spring and late fall, the conditions discussed above may

have proved too negative, in terms of cold and predators, for this species to proliferate

under the circumstances.

It was surprising to find such low numbers of C. terraenovae during the present

study. This blow fly is most common in the northern United States and southern Canada,

especially the Rocky Mountains. The seasonal activity pattern for this species is from

late July to August (Hall 1947). The reason behind this blow fly's near absence is

unexplained although temperature, overcrowding, and predation could be factors.

Temperature fluctuations and behavior relations for Ph. regina have been noted

previously by Deonier (1940). Deonier found that this blow fly has a minimum activity

temperature of 5°C. Several days in the first month of the study experienced

temperatures near 5°C and Ph. regina larvae and adults were active in low numbers. This

low activity level doubtlessly played a role in the extended developmental time observed

for this blow fly.

L. illustris has been categorized by Hall (1947) and Greenberg (1973) as an open

woodland and meadow blow fly species. Both Anderson (1995) and Greenberg (1973)

agree that it has a Holarctic distribution, from southern Canada to northern Mexico and is

37

both an. urban and rural species. L. illustris are attracted primarily to fresh carcasses and

only rarely to dung (Greenberg 1973). Ironically, in the present study the only adults

collected were, in fact, found on the first day of the study during the fresh stage. No L.

illustris were found again until the first immatures were sampled 17 days later. Due to

the absence of immatures at this time of any other blow fly species.

Cole (1969) has reported Prot. terraenovae in the western US at high altitudes. It

may perhaps be viewed as the counterpart of Ph. regina at the study site used in the

present study. It is predominantly an early spring species that occurs in Utah from

March-June at altitudes o f4,000 to 6,000 feet. It has also been found in Colorado at

7,000+ feet from July to August (Hall 1947). This species therefore has a more northern

distribution than Ph. regina, and so it is not surprising that it was in lower numbers.

Two species of Piophilidae, Proch. brericomis and. Proch. xanthostoma, were

found mating in July and adults were found through early October on both carcasses'.

The presence of these Piophilidae species is not surprising. Cole (1969) reported

specimens of Proch. brericomis from Flathead Lake, MT and Yellowstone Park from

July through August. Cole (1969) found that Proch. xanthostoma occurs from Colorado

to California and north to Alaska. These species were most abundant during the post­

decay stage and may contribute more to PMI calculations during this stage than other

Diptera present.

The primary difference in the succession pattern during the fresh and bloat stages

for Sepsidae on the burnt and control carcasses. Succession during the decay stage was

similar for both carcasses while the post-decay stage brought about the greatest

divergences in species diversity. Calliphoridae species such as Phae. caeruleiviriais,

38

Phae. sericata, Prot. terraenovae, and C. terraenovae each showed a preference for one

of the carcasses (Tables 1 and 2). Species of the families Fannidae, Sarcophagidae and

Sepsidae became numerous on both carcasses.

Coleoptera

Coleoptera were not present during the fresh stage. During the bloated stage the

species Dermestes and T. lapponicus were the most abundant. The major differences

between coleopteran in the two carcasses were the types and population sizes of predators

of dipteran larvae. The only contrast between the insects on the two pigs during the

decay stage was the absence of N. surinamensis and Nicrophorus on the control carcass.

Coleoptera of the post-decay stage were of the same composition for each carcass, with

the major obserable variation being in numbers of Staphylinidae and Histeridae

predators.

Hymenoptera and Other Arthropods

The only contrast in presence of Hymenoptera, non-coleopteran and dipteran

arthropods between carcasses, was in species of Reduvidae (order Hemiptera) and the

order Araneida. Both of these groups are predaceous upon insects and have been

observed in previous forensic entomology studies (Payne et. al. 1968). The majority of

the Hymenoptera collected in the present study were incidental species that did not

interact directly with the carcass or the necrophagous species present. However, some

Hymenoptera collected were predacious upon insect larvae and eggs (Formicidae and

Vespidae wasps) or parasitic insects (Ichneumonidae and Pteromalidae).

39

A single species of Lepidoptera was observed in the site area. Although it did not

appear to interact with the carcasses themselves, Anderson et. al. (1996) has reported that

one Lepidopteran in the family Nymphalidae was collected while feeding on carrion.

Competition and Successional Pattern

Competition occurs between insects when utilizing a limited resource (Birch

1957). Competition among Diptera may have occurred in the pigs. A geometric

distribution pattern can indicate that one or two species are highly abundant, and

comprise roughly two thirds of the total numbers of individuals (Kuusela and Hanski

1982). This pattern was found in the present study and is often found among carrion fly

communities (Nuorteva 1970, Groth and Reissmiiller 1973, Beaver 1977, Hanski and

Kuusela 1977, Palmer 1980).

Examination of the species diversity for each pig indicates that competition may

have played a role in the outcome of the successional pattern. Kneidel (1984) predicted

that large carrion should have high levels of Diptera competition with low species

diversity among decomposition stages. Diptera species diveristy was high during every

stage in the present study. A high species diversity in the presence of a large carcass is to

be expected (Schoenly and Reid 1983). A possible explanation is that each carcass was

large enough to provide adequate niches and resources for a higher number of species.

While the size of the carcass would seem to be the dominant factor in the discussion of

species diversity, one must remember that the variation in environmental conditions, site

diversity, and baseline populations of colonizing insects are also significant factors to

consider.

40

Gilbert and Bass (1967) have reported finding pupae of Callitroga spp. and Ph.

regina in 130-160 year old Arikara burials. With the knowledge that both families of

blow fly appear by late March in the area of Gilbert and Bass's (1967) excavations and

are gone by mid-October the authors could determine when the burials were made.

Using this information, an archaeologist could determine the seasonal use of a site.

Burial practices may also be ascertained from this evidence. If a body was laid out for

defleshing or set out for a time until burial could take place, blow flies may have time to

lay eggs, complete a number of instars, or even reach the pupal stage.

Greenberg (1990) found Ph. regina postfeeding larvae pupate close to the carcass

and do not migrate farther than 3.3 m from the food site. In the present study, a puparium

was found 17 m from the datum point and it could be speculated that this pupal case was

from a different larval migration and was not associated with the study pigs. Dozens of

pupal casings were found in the immediate vicinity of the carcasses. The majority were

located underneath the carcasses in the topsoil. Unfortunately, the species to which the

pupal casings belong is undetermined. But with the predominance of Ph. regina larvae,

and the postfeeding behavior typical of the species, it is most likely that the majority are

from this blow fly.

41

Conclusions-Chapter 5

The first hypothesis tested in this study was that the successional fauna in pig

carcasses present in western Montana will be similar to that of pigs used by Dillon

(1997), who conducted a forensic entomological study in a similar biogeoclimatic zone,

to that used in the present study, located in British Columbia, Canada. The present case

study was conducted in a similar biogeoclimatic zone of interior Douglas-fir with a

mixture of ponderosa pine.

There were two principal factors to compare between the two studies. The first

comparison is the type of Diptera species present. Ph. regina and Prot. terraenovae were

the two most abundant species in Dillon's study. The present study had Ph. regina

(Meigen) and L. illustris as the two most abundant Calliphoridae with Prot. terraenovae

being less abundant. Ph. regina is a urban and rural cold-weather species, common to

high altitudes from the spring into the fall seasons. P. terraenovae has habits similar to

Ph. regina but is more common to northern regions of North America and is considered

Ph. regina's northern counterpart by Hall (1947). Ph. regina 1st and 2nd instar larvae

were also found into the post-decay stage in the present study, much later than in Dillon's

study. L. illustris is also urban and rural, but prefers habitats which are more open.

Reasons why L. illustris was not as abundant on Dillon's test subjects is unknown but

may be due to competition between the two species.

Additional Diptera species diversity similarities include the presence of Fanniidae

almost exclusively during the post-decay and remains stages for both studies. The

primary contrast in Diptera activity was the early arrival of Piophilidae larvae oh Dillon's

test subjects between days 21-26. In contrast, Piophilidae larvae were not found on

42

carcasses in the present study until day 56. These arrival times are both earlier than

previously reported 3-6 months for Southern Canada and Europe (Smith 1986).

Decay rates were similar for both studies with only a 2-3 day difference between

stages and similar lengths of time for cohesive maggot masses (2 days). Also, no

reduction in insect numbers and diversity was observed in either study as the carcasses

became desiccated in the remains stage. Furthermore, test subjects for both studies were

only colonized by rural blow flies. A final interesting point is that species diversity was

roughly the same for both studies although the pigs in the present study weighed

approximately 82 kilograms and Dillon's pigs weighed about one-third this amount.

Coleoptera families were similar for both studies but species similarity could not be

determined, except in the case of the family Silphidae, as other Coleoptera were not

identified beyond genus for the present study. The Silphidae species N. surinamensis, T.

lapponicus, and genus Nicrophorus were found in both studies. The only contrast was

that Dillon found no immatures of these two species while immatures were prolific in the

present study. This could be due to many factors including predation or adverse

environmental conditions for reproduction.

These parallels in decay rates and species diversity for two studies conducted in an

interior Douglas-fir and Douglas-fir/ponderosa pine Rocky Mountain biogeoclimatic

zone may or may not be coincidence. Although it would be precarious to directly

extrapolate data from one region to the other for forensic purposes, one could possibly

confirm case study data found in one region by comparing it with the baseline data

presented here. Direct extrapolation of data may lead to errors until more information is

compiled for comparison. This tentative confirmation with data collected in a similar

43

biogeoclimatic zone, yet hundreds of miles away, is a positive step for Montana forensic

entomological studies.

The second hypothesis states that the burned and unbumed specimens will have

different successional fauna specific to their physical states. The successional pattern

was very similar between the burnt and unbumt pigs. The rate of decay differed by

approximately one day during the decay stage and the co-dominant species present was

the same for each pig throughout the study. The similarities between the conclusions of

this bum study and the previous bum study by Avila and Goff (1998) are also not

surprising. Unfortunately, species types, successional patterns, and decay rates could not

be compared due to an extreme contrast in biogeoclimatic zones (Hawaii and Montana)

and indigenous species. From Avila and Goff (1998) and the present study it maybe

concluded that carrion Diptera and Coleoptera are attracted to burned carrion as quickly

as to non-bumed carrion. Future studies may determine if differences in accelerant use

create differences in successional patterns of species diversity.

The third hypothesis for this study states that the insect successional pattern and

species diversity on carcasses present in a mixed urban/rural and those present in a rural

area will be distinct (Temenyl997). A comparison was made with results from the

present study and those of Temeny (1997) who conducted a spring study in a mixed

urban/rural environment. The decomposition stages of the Temeny study compared with

those of the present study are presented in Table 4.

Temeny (1997) defines the end of the decay stage as when the greater part of the

flesh has been removed. In Temeny's study maggots were visible by day 11 compared

with day 5 in the present study. This developmental delay as compared with

44

developmental time in the present study may be due to cooler temperatures in the spring,

although no climatological data is presented in Temeny's study, making comparisons

difficult.

No specific statements can be made concerning a comparison of species diversity for

spring/summer habitats and urban/rural locations using Temeny's Montana study.

Because insects were not identified beyond the family level. All of the families collected

in Temeny's study were also found in the present study but nothing more may be

generalized with the data provided.

The present case study has provided new information to be included in the forensic

entomology database for the state of Montana. The study has revealed numerous

similarities in insect succession with an experiment performed in a similar biogeoclimatic

zone located in British Columbia, Canada. While this information should not be directly

extrapolated to forensic cases in Montana it does provide investigators with an additional

database for use in forensic entomology cases in a similar environment. The results of

the present bum study parallel those found by Avila and Goff (1998) and strengthen their

decision that although a slight time difference occurs among the early decompositional

stages, the fauna for both burned and unbumed carcasses remains similar. Case studies

from both urban and rural forensic cases obtained from the Montana Forensic Science

Division have been, and will continue to be, analyzed for an ameliorated view of forensic

entomology in the state of Montana.

45

Table 1Insect Succession for the burnt Sus scrofa:

Stage and time since death

Fresh (day 1)

Bloat (days 2-8)

Order Familv Genus and species

Diptera Calliphoridae Phormia reginaEucalliphoraProtophormia terraenovaeLucilia illustris

Sarcophagidae Sarcophaga sp. 1Sarcophaga sp. 2Sarcophaga sp. 3Sarcophaga sp. 4

Fannidae Fannia sp. 1Tachinidae sp. 1

sp. 4Phoridae sp. uniden.

Hymenoptera Megachilidae not collectedVespidae not collected

Lepidoptera spe. 1

Diptera Calliphoridae Phormia regiriaEucalliphoraProtophormia terraenovaeCalliphora terraenovae

Sarcophagidae Sarcophaga sp. 1Sarcophaga sp. 2

Fanniidae Fannia sp. 4Piophilidae Prochyliza brericomisSepsidae Meroplius Stercoronius

Sepsis sp.Coleoptera Silphidae Silpha sp.

Dermestidae Dermestes sp.Scarabaeidae sp. uniden.

sub: AphodiinaeHisteridae sp. uniden.Staphylinidae sp. uniden.Chrysomelidae sp. uniden.Tenebrionidae sp. uniden.Elateridae sp. 4

Hymenoptera Formicidae sp. uniden.Megachilidae sp. uniden.

sub: MegachilinaeVespidae sp. uniden.

sub: Vespinae Acari sp. uniden.

InsectStage

adadadadadadadadadadadad

ad

ad, imadadadadadadadadadadadad

adadadadadadad

ad

46

Decay (days 9-13)

Post-decay (14-37)

Diptera Calliphoridae Phormia regina ad, imProtophormia terraenovae ad

Sarcophagidae Sarcophaga sp. 6 adFanniidae Fannia sp. 1 adPiophilidae sp. 2 ad

Coleoptera Silphidae Nicrophorus adNecrodes surinamensis adThanatophilus lapponicus ad, im

Dermestidae Dermestes ad, imHisteridae sp. uniden. adStaphylinidae sp. uniden. ad

Hymenoptera Formicidae sp. uniden. adVespidae sp. uniden. ad

Acarisub: Vespinae

sp. uniden.

Diptera

Coleoptera

Calliphoridae Phormia regina(1st, 2nd, and 3rd instars)

ad, im

Eucalliphora adProtophormia terraenovae adLucilia illustris

(3rd instar)ad, im

Calliphora terraenovae adPhaenecia caeruleiviriais ad

Sarcophagidae Sarcophaga spe. 1 adSarcophaga spe. 5 ad

Fanniidae Fannia canicularis adFannia difficilus adFannia sp. 1 ad

Piophilidae Prochyliza brericomis adProchyliza xanthostoma adsp. 1 adsp. 2 ad

Sepsidae Meroplius Stercoronius adSepsidomorph sp. uniden. ad

Tachinidae Dexinae sp. uniden. adsp. 1 ad

Anthomyiidae sp. uniden. adChloropidae sp. uniden. adEmpididae sp. uniden. adRhagionidae sp. uniden. adSilphidae Nicrophorous sp. uniden. ad

Silpha sp. uniden. adThanatophilus lapponicus ad,imNecrodes surinamensis ad,im

Dermestidae Dermestes sp. uniden. adScarabaeidae sp. uniden. ad

sub: ScarabaeinaeHisteridae sp. uniden. ad

47

Hymenoptera

Remains (days 38+)

AraneidaAcari

Diptera

Coleoptera

Staphylinidae sp. uniden. adChrysomelidae sp. uniden. adCoccinellidae sp. uniden. adElateridae sp. 1 ad

sp. 2 adFormicidae sp. uniden. adIchneumonidae sp. uniden. adColletidae sp. uniden. ad

sub: HylaeinaeSphecidae sp. uniden. ad

sub: SpheciniMegachilidae sp. uniden. ad

sub: MegachilinaeApidae sp. uniden. ad

sub: Apinae tribe: BombiniVespidae sp. uniden. ad

sub: Vespinaesub: Araneae sp. 1 adsp. uniden. ad

Calliphoridae Phormia regina adProtophormia terraenovae ad

Sarcophagidae Sarcophaga sp. 1 ad, imSarcophaga sp. 3 adSarcophaga sp. 4 adSarcophaga sp. 5 ad

Fanniidae Fannia canicularis adFannia sp. 1 adFannia sp. 2 adFannia sp. 5 ad

Piophilidae Prochyliza brericomis ad, imProchyliza xanthostoma ad, im

Sepsidae Sepsis sp. uniden. adTachinidae Dexinae sp. uniden. ad

sp. 1 adsp. 2 adsp. 5 ad

Drosophilidae sp. uniden. adTrixoscelididae sp. uniden. adOtitidae sp. uniden. adHeleomyzidae sp. uniden., mating adAnthomyiidae sp. uniden. adEmpididae sp. uniden. adSilphidae Silphus sp. uniden. ad

Thanatophilus lapponicus imNecrodes surinamensis im

Dermestidae Dermestes sp. uniden. ad, imStaphylinidae sp. uniden. adHisteridae sp. uniden. adChrysomelidae sp. uniden. ad

48

Elateridae sp. 3 adNitidulidae sp. uniden. ad

Hymenoptera Formicidae sp. uniden. adIchneumonidae sp. uniden. adSphecidae sp. uniden. ad

sub: Sphecini Apidae sp. uniden. ad

sub: Apinae tribe: Bombini Vespidae sp. uniden. ad

sub: Vespinae Vespidae sp. uniden. ad

sub: Eumeninae Andrendiae sp. uniden. ad

sub: Panurginae Andrenidae sp. uniden. ad

sub: Andreninae Superfamily Chalcidoidea

Pteromalidae sp. uniden. adHemiptera Reduvidae sp. 1 ad

sp.2 adsp.3 adsp.5 ad

Araneida Araneae sp. 1 adsp.5 ad

Acari sp. uniden. ad

49

Table 2Insect Succession for the control Sus scrofa:

Stage and time since death

Fresh (day 1)

Bloat (days 2-8)

Decay (days 9-13)

InsectOrder_______ Family_________ Genus and species__________Stage

Diptera Calliphoridae Phormia regina adEucalliphora adProtophormia terraenovae adLucilia illustris adPhaenecia cadaverina ad

Sarcophagidae Sarcophaga sp. 1 adSarcophaga sp. 2 adSarcophaga sp. 3 adSarcophaga sp. 4 ad

Faniidae Fannia sp. 2 ad: Heleomyzidae sp. uniden. adAnthomyiidae sp. uniden. adPhoridae sp. uniden. ad

Hymenoptera Megachilidae not collectedVespidae not collected

Diptera Calliphoridae Phormia regina ad, imEucalliphora adProtophormia terraenovae adCalliphora terraenovae ad

Sarcophagidae Sarcophaga sp. 1 adSarcophaga sp. 2 ad

Piophilidae Prochyliza brericomis adSepsidae Nemapoda sp. uniden. adHeleomyzidae sp. uniden. ad

Coleoptera Silphidae Silpha spe. uniden. adDermestidae Dermestes sp. uniden. adHisteridae sp. uniden. adStaphylinidae sp. uniden. adChrysomelidae sp. uniden. adScolytidae sp. uniden. ad

Hymenoptera Formicidae sp. uniden. adAcari sp. uniden.

Diptera Calliphoridae Phormia regina ad, imProtophormia terraenovae ad

Sarcophagidae Sarcophaga sp. 6 adPiophilidae sp. 2 ad

Coleoptera Silphidae Thanatophilus lapponicus ad, imDermestidae Dermestes ad, imHisteridae sp. uniden. adStaphylinidae sp. uniden. ad

Hymenoptera Formicidae sp. uniden. ad

50

Post-decay (14-37)

Vespidae sp. uniden. adsub: Vespinae

Acari sp. uniden.

Diptera Calliphoridae Phormia regina ad, im(1st, 2 , and 3rd instars)

Eucalliphora adLucilia illustris ad, im

(3rd instar)Phaenecia sericata ad

Sarcophagidae Sarcophaga sp. 1 adSarcophaga sp. 5 ad

Fanniidae Fannia canicularis adFannia sp. 1 adFannia sp. 3 ad

Piophilidae Prochyliza brericomis adProchyliza xanthostoma ad

Sepsidae Meroplius Stercoronius adNemapoda sp. uniden. ad

(mating)Sepsis sp. uniden. ad

Tachinidae sp. 1 adSyrphidae sp. uniden. adLauxaniidae sp. uniden. adDryomizidae sp. uniden. adTabanidae sp. uniden. ad

sub: TabanusDiapriidae sp. uniden. ad

Coleoptera Silphidae Nicrophorous adsp. uniden. adSilpha sp. uniden. adThanatophilus lapponicus ad,imNecrodes surinamensis ad,im

Dermestidae Dermestes sp. uniden. adScarabaeidae sp. uniden. ad

sub: Scarabaeinae Histeridae sp. uniden. adStaphylinidae sp. uniden. adChrysomelidae sp. uniden. adCurculionidae sp. uniden. ad

sub: Brachyrhininae Salpingidae sp. uniden. adElateridae sp. 1 ad

sp. 2 ad

Hymenoptera Formicidae sp. uniden. adIchneumonidae sp. uniden. adColletidae sp. uniden. ad

sub: Hylaeinae Sphecidae sp. uniden. ad

51

Remains (days 38+)

sub: SpheciniMegachilidae sp. uniden. ad

sub: Megachilinae Apidae sp. uniden. ad

sub: Apinae tribe: Bombini Apidae sp. uniden. ad

sub: Apinae tribe: Apini Vespidae sp. uniden. ad

sub: Vespinae Halictidae sp. uniden. ad

sub:Halictinae Andrendiae sp. uniden. ad

sub: PanurginaeLepidoptera sp. 2 adAcari sp. uniden.

Diptera Calliphoridae Phormia regina adSarcophagidae Sarcophaga sp. 1 ad, im

Sarcophaga sp. 3 adSarcophaga sp. 4 adSarcophaga sp. 5 ad

Fanniidae Fannia sp. 1 adFannia sp. 2 ad

Piophilidae Prochyliza brericomis ad, imProchyliza xanthostoma ad, im

Sepsidae Sepsis sp. uniden. adTachinidae Dexinae sp. uniden. ad

sp. 1 adsp. 2 adsp. 3 adsp. 4 ad

Trixoscelididae sp. uniden. adOtitidae sp. uniden. adHeleomyzidae sp. uniden., mating adSyrphidae sp. uniden. ad

Coleoptera Silphidae Silphus sp. uniden. adThanatophilus lapponicus imNecrodes surinamensis im

Dermestidae Dermestes spec, uniden. ad, imStaphylinidae sp. uniden. adHisteridae sp. uniden. adChrysomelidae sp. uniden. adLygaeidae sp. uniden. adElateridae sp. 3 adNitidulidae sp. uniden. adScarabaeidae sp. uniden. ad

sub: Troginae Curculionidae sp. uniden. ad

sub: Brachyrhininae Hymenoptera Formicidae sp. uniden. ad

52

Ichneumonidae sp. uniden. adColletidae sp. uniden. ad

sub: Hylaeinae Sphecidae sp. uniden. ad

sub: SpheciniMegachilidae sp. uniden. ad

sub: Megachilinae Halictidae sp. uniden. ad

sub:Halictinae Andrendiae sp. uniden. ad

sub: Panurginae Andrenidae sp. uniden. ad

sub: Andreninae Super ChalcidoideaPteromalidae sp. uniden. ad

Hemiptera Reduvidae sp. 1 adsp. 2 adsp. 4 ad

Araneida Araneae sp. 1 adsp. 2 adsp. 3 adsp. 4 adsp. 6 ad

Acari sp. uniden. ad

53

Table 3

Comparison of decay stage lengths from Dillon (1997) and the present study:

Decay Stage______________ Length in days: Dillon . Length in days: Present Study

Fresh 2 1Bloat 5 7Decay 8.5 5Post-Decay 65 77

Table 4

Comparison of decay stage lengths from Temeny (1997) and the present study:

Decay Stage_____________ Length in days: Dillon______ Length in days: Present Study

Fresh 2 1Bloat 7 7Decay 65 5Post-Decay 25 77

54

Figure 1

Species Diversity, FFesh: Burnt1.000

0.001

I. Phonria regina 2.SarcaSp.l3. Protaphorrria terraenovae4. Ludlaillustris5. Eucalliphora6. Saim Sp. 27. Saim Sp. 38. Saim Sp 49. Farniia Sp 110. Tachinidae Sp. 1II. Tachinidae Sp. 4

9 10 11

Types of Spedes

Species Diversity, Fresh: Control* * 1.000

0.0018 12 13 162 4 5 6 7 14 IS1

L Phormia regina 2 Sarca Sp 14. Lutilia Oiusrtris5. Eucalliphora 6S arcoS p2 7. SarcoSp3 &SarcaSp. 4 12Phaeneda cadaverina13. Fannia Sp 2 14 Hdeopmyzidae 15. Anthomyiidae 16 Phoridae

Types of Spedes

55

Figure 2

Species Diversity, Bloated: Burnt1. Phormia regina 2.SarcaSp.l 3. Protophormia5. Eucalliphora6. Sarca Sp. 217. Prochyliza brericomis18. Chlliphora terraenovae19. Fannia Sp. 420. Memphis stercomis21. Sepsis Sp.

1 2 3 5 6 17 18 19 20 21

Types of Spedes

ft 0.100,

0.010 :

Species Diversity, Bloated: Control1.000

g 0.100

3 5 14 17

Types of Species18 22

1. P. regina2. Sarca Sp. 13. Protophormia5. Eucalliphora14.Heleomyzidae17. Prochyliza brericomis18. Calliphora terraenovae 22. NemannHn

56

Figure 3

Species Diversity, Decay: Burnt1.000 ?

1. Phormia regina

Protophormiaterraenovae

0.010 :

0.001

Types of Species

Species Diversity, Decay: Control1.000 q 1. P. regina

Protophormia terraenovae 23. Sarca Sp.S 0.100 :

24. Piophilidae

0.010 :

0.001

Types of Species

57

Figure 4

Spedes Diversity, Post-Decay: Burnt1.000

sa4)OUaa.Vuart-OsaaA<<u

•js<9

cc

0.100

aoio

0.0011 2 3 4 5 9 10 15 17 18 20 25 26 27 28 29 30 31 32 33 34 35 36

Types of Spedes

1. Phormia regina 2.SarcaSpl3. Protophonria4. LucillaOlustris5.Eucafliphna9. Fannia Sp 110. Tachinidae Sp 1 15. Anthontyiidae17. Prochyliza breritaniis18. Calliphora terraenovae 20. Merioplius stericomis25. Phaeneciacaeruldviriais26. Sarax Sp527. Fannia canktdaris28. Fannia difficflus29. Prochyliza xanthostoma30. Sepsidomorph31.Dexinae32. Gdoropidae33.Enpdidae34. Rhagionidae 35Pi(){iiilidaeSp.l'V* PiiwhSliHao An 7

Species Diversity, Post-Decay: Control

« 0.100 :

< 0.010 :

0.0011 2 5 9 10 17 20 22 29 37 38 39 40 41

Ttypes of Spedes

1. P. regina2. Saim Sp 1 5JEucalliph.9. Fannia S p l10. Tachinidae Sp 1 17. Prochyliza brericomis20. Mencius stercoroniiK22.Nemapoda 29. Prochyliza xanthostoma37. Syrphidae38. Lauxaniidae39. Diyorayz.40. Tabanidae

58

Figure 5

I. Fhomia reginaSpedes Diversity, Remains: Burnt 2. Sarcophaga S p l

3. ftotophorrria10. Tachinidae sp. 1II. Tachinidae Sp 214.Hdeomradae15. Anthonwiidae17. Prochyliza brericomis21. Sepsis 27. Fannia canicularis 29. Prochyliza xairthostoma 31.Dexinae42.Drosophilidae43. Irixoscdididae

<' 44. Otitidae1 2 3 10 11 14 15 17 21 27 29 31 42 45 44 45 46 47 45. FanniaSp. 5

Types of l̂ redes 46. Tachinidae Sp 547. Empididae

Species Diversity, Remains: Control1.000

a88Ana> 0.100 « a «

T9a s .o<

I0.010

£0.001

8 9 10 11 13 14 15 17 21 26 29 31 37 43

1. Phormia regina2. Sarcophaga Sp. 17. Sarcophaga Sp. 38. Sarcophaga Sp. 49. Fannia Sp. 110. Tachinidae Sp.l11. Tachinidae Sp. 213. Fannia Sp. 214. Heleomyzidae15. Tachinidae Sp. 3 17. Prochyliza brericornis21. Sepsis26. Sarcophaga Sp. 5 29. Prochyliza xanthostoma 31. Dexinae 37. Syrphidae 43. Trixoscelididae

Types of Species

59

Figure 6

Spedes Diversity, Bloated: Bunt

1. Dermestes 2 Thanatophilus lapponiciB3. Staphyliridae 4 Scarabaeidae sub: Aphodiinae 5. Lfistseridae & Onysomelidae7. Tenebrionidae & Elateridae Sp. 4

1 2 3 4 5 6 7 8

Types of Species

Species Diversity, Bloated: Control1.000

£ 01100:

Types of Species

60

Rela

tive A

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ance

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cent

o

M

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»H

r

Rela

tive A

bund

ance

Per

cent

Figure 7

Figure 8

Spedes Diversity, Post-Decay: BurntI. DenrcstesZ ThanatopKliB bpporiciE 3. Stapfyiimbe5. Ifcteridae6, QrvwrreBdaeII. NcropinroiB 1Z Scarabaeidae sife Scarabaeirae13. Cbcrinellidae14. Qeterkbe Sp 1

1 2 3 5 6 11 12 13 14

Types of Spedes

Spedes Diversity, Post-Decay: ControlI. DernestesZ Thanatoplihs lagporiaB 3. Staphyliiiclae5. Ifcteridae6. ChrysorreliddeII. NcrophoroiE 1Z Scarabaeidae 15. CUrctfioiidae 1& Saifingidae17. Elateridae Sp 2

1 2 3 5 6 11 12 15 16 17

Types of Spedes

62

Figure 9

Species Diversity, Remains: Burnt1. Denmestes ZlhaiatqMiK la(pricus3. Stafiryiini(fae5.Hsteridae6.Girysameli(be18.HatericbeSp319. NMdiicfae 20.0titidae

1 2 3 5 6 18 19 20

'types of Spedes

Spedes Diversity, Remains: Control1.000

< 0.010

0.0012 3 5 6 15 18 8 21 22

Types of Spedes

1. Dermestes2. ThanatophiliR Lapponicus3. StaphylinidaeS. Ifcteridae (■.Chysomelidae15. CknuKoradae18. Elateridae Sp 319. NitxMidae 2L Scarabaeidae adx Ttoginae 22.Ljgaeidae

63

Tem

pera

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in

Celci

us

Tem

pera

ture

in

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ius

Figure 10 Temperature: BSS

S * \

2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Time Since Death in Days—• —Ambient Temp. Low —♦—Ambient Temp. High —A—Internal Temp.

Temperature: CSS

50 i

rVA H- 20

2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

l ime Since Death in Days—• — Ambient Temp. Low —♦ —Ambient Temp. High - A - Internal Temp.

64

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