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The Forensically Important Calliphoridae (Insecta: Diptera) of Pig Carrion in Rural North-Central Florida

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INGEST IEID E20110114_AAAAAG INGEST_TIME 2011-01-14T06:28:15Z PACKAGE UFE0007010_00001
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PAGE 1

THE FORENSICALLY IMPORTANT CALLIP HORIDAE (INSECTA: DIPTERA) OF PIG CARRION IN RURAL NORTH-CENTRAL FLORIDA By SUSAN V. GRUNER A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2004

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Copyright 2004 by Susan V. Gruner

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For Jack, Rosamond, and Michael

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iv ACKNOWLEDGMENTS The successful completion of this thesis would not have been possible without the support, encouragement, understanding, guidanc e, and physical help of many colleagues, friends, and family. My mother, Rosamond, edite d my thesis despite her obvious distaste for anything related to maggots. My husba nd, Michael, cheerfully allowed more than most spouses could bear. And when it got real ly cold outside, he only complained two or three times when I kept multiple containers of stinking liver and writhing maggots on the kitchen counter. He also took almost all of the fantastic photos presented in this thesis and for my presentation. Finall y, Michael had the horrible job of inserting an arrow with twelve temperature probes into pig “E.” I am also indebted to Dan and Jenny Slone, Jon Allen, Debbie Hall, Jane and Buthene Haskell, Dan and Zane Greathouse, owners of Greathouse Butterfly Farm, Aubrey Bailey, and the National Institute of Justice. My professional colleagues, John Capine ra, Marjorie Hoy, and Neal Haskell, guided and encouraged me through my resear ch and writing every step of the way. Although I doubt that John Capinera was great ly interested in maggots, his support and enthusiasm in response to my enthusiasm are very much appreciated. Marjorie Hoy’s beneficial advice was always appreciated and on occasion, she offered a shoulder on which to cry. Neal Haskell, as usual, we nt above and beyond the call of duty in every aspect of my time spent as an M.S. graduate student. I thank them all for their support. Finally, I thank the pigs whic h were sacrificed in the na me of forensic science.

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v TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES............................................................................................................vii LIST OF FIGURES.........................................................................................................viii ABSTRACT....................................................................................................................... xi CHAPTER 1 INTRODUCTION AND LITERATURE REVIEW....................................................1 Definition of and Scope of Forensic Entomology........................................................1 History of Forensic Entomology..................................................................................2 Current Status of Forensic Entomology.......................................................................6 Biology of Calliphoridae..............................................................................................8 Insect Succession on Carrion......................................................................................10 Factors that Affect Blow Fly Succession on Carrion.................................................12 Biology of Human Decomposition.............................................................................14 Further Areas for Study..............................................................................................15 2 MATERIALS AND METHODS...............................................................................17 Study Site....................................................................................................................1 7 Data Collection....................................................................................................18 Protocol for Day 1...............................................................................................18 Protocol for Day 2 Onward.........................................................................................19 Meteorological Measurements............................................................................19 Rearing Procedures..............................................................................................20 Pupation Substrate...............................................................................................21 3 RESULTS...................................................................................................................29 Species of Calliphoridae Co llected on Pig Carrion....................................................29 Seasonal Distribution and Succession of Calliphoridae Species.........................30 Collection 1, November 16-21, 2001..................................................................31 Collection 2, December 29, 2001-January 11, 2002...........................................31

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vi Collection 3, February 5-9, 2002.........................................................................31 Collection 4, March 15-19, 2002.........................................................................32 Collection 5, April 29-May 1, 2002....................................................................32 Collection 6, May 20-23, 2002............................................................................32 Collection 7, July 22-25, 2002............................................................................32 Collection 8, August 19-23, 2002........................................................................33 Collection 9, September 23-27, 2002..................................................................33 Collection 10, October 26-28, 2002....................................................................34 Collection 11, November 30-December 14, 2002...............................................34 Collection 12, December 30, 2002-January 11, 2003.........................................35 Collection 13, March 2-8, 2003...........................................................................35 Collection 14, April 1-6, 2003.............................................................................35 Collection 15, April 26-May 1, 2003..................................................................36 Collection 16, June 12-15, 2003..........................................................................36 Collection 17, December 8-20, 2003...................................................................36 Collection 18, January 23-31, 2004.....................................................................37 Collection 19, March 5-14, 2004.........................................................................37 4 DISCUSSION.............................................................................................................68 APPENDIX A RAW DATA: FLIES COLLECTED AS ADULTS OR REARED FROM LARVAE....................................................................................................................75 B RAW DATA: PRESERVED SPECIMENS...............................................................83 LIST OF REFERENCES...................................................................................................88 BIOGRAPHICAL SKETCH.............................................................................................98

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vii LIST OF TABLES Table page 2-1 Dates of pig deposition, approximate pig weight, and time each pig was placed on site for sampling dates from November, 2001 to October, 2002........................27 2-2 Mean temperatures ( SD) during the study at the Earleton, Florida site................28

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viii LIST OF FIGURES Figure page 2-1 A caged pig carcass is sh own at Greathouse Butterfly Farm property in Earleton, Florida......................................................................................................................23 2-2 Wire cage with bungee cords and tent stakes hammered into the ground...............24 2-3 A HOBO temperature da ta logger is sealed insi de a Gladware container...........25 2-4 The data logger was hung from bush at left and temperature probe was placed on the ground (circled) to record ground temperature near the pig carcass in Earleton, Florida.......................................................................................................26 3-1 Relative abundance of callipho rid adults aerially collected.....................................38 3-2 A calliphorid first-instar larva hatching from its egg...............................................39 3-3 The two sets of spiracles of a first-second transitional larva...................................40 3-4 The two inner slits in the spiracles of a second-instar calliphorid larva..................41 3-5 The two sets of spiracles of a s econd-third transitional calliphorid larva................42 3-6 The three inner splits in the spiracles of a thirdinstar calliphorid larva..................43 3-7 Chrysomya rufifaces third-instar larvae.................................................................44 3-8 Relative abundance of th ird-instar calliphor id larvae preserved from pig carrion during the study (N=8253) in Earleton, Florida.......................................................45 3-9 Calliphorid activity fo r year 1 of the study..............................................................46 3-10 Calliphorid activity fo r year 2 of the study..............................................................47 3-11 Mean daily and mean low temperatur es (SD) for year 1, November 16, 2001 to October 26, 2002 (Part A) and year 2, November 30, 2002 to March 14, 2004 (Part B) of study.......................................................................................................48 3-12 Reared adults, N=232, (Part A), and preserved larvae, N=30, (Part B), from collection 1, November 16 to November 21, 2001..................................................49

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ix 3-13 Reared adults, N=362, (Part A), and preserved larvae, N=568, (Part B), from collection 2, December 29, 2001 to January 11, 2002.............................................50 3-14 Adults aerially collected N=39, (Part A), reared adults, N=305, (Part B), and preserved larvae, N=151, (Part C), from collection 3, February 1-9, 2002.............51 3-15 Adults collected aerially, N=71, (Part A), reared adults, N=326, (Par B), and preserved larvae, N=134, (Part C), from collection 4, March 15-19, 2002.............52 3-16 Adults aerially collect ed, N=107, (Part A), reared adults, N=332, (Part B), and preserved larvae, N=196, (Part C), from collection 5, April 29 to May 1, 2002.....53 3-17 Adults aerially collected N=51, (Part A), reared adults, N=67, (Part B), and preserved larvae, N=175 (Part C), from collection 6, May 20 to May 23, 2002.....54 3-18 Adults aerially collected N=99, (Part A), reared adults, N=223, (Part B), and preserved larvae, N=700, (Part C), from collection 7, July 22 to July 24, 2002......55 3-19 Adults aerially collected N=76, (Part A), reared adults, N=248, (Part B), and preserved larvae, N=499, (Part C) from collection 8, August 19-23, 2002............56 3-20 Adults aerially collected N=76, (Part A), reared adults, N=323, (Part B), and preserved larvae, N=499, (Part C), fro m collection 9, September 23-27, 2002.......57 3-21 Adults aerially collected N=87, (Part A), reared adults, N=131, (Part B), and preserved larvae, N=294, (Part C), from collection 10, October 26-28. 2002.........58 3-22 Adults aerially collected N=60, (Part A), reared adults, N= 384, (Part B), and preserved larvae, N=1248, (Part C), from collection 11, November 30December 8, 2002....................................................................................................59 3-23 Adults aerially collected N=56, (Part A), reared adults, N= 371, (Part B), and preserved larvae, N=581, (Part C), from collection 12, December 30, 2002 to January 11, 2003......................................................................................................60 3-24 Adults aerially collected N=66, (Part A), reared adults, N=166, (Part B), and preserved larvae, N=340, (Part C) from collection 13, March 1-8, 2003...............61 3-25 Adults aerially collect ed, N=247, (Part A), reared adults, N=229, (Part B), and preserved larvae, N=301, (Part C), from collection 14, April 1-6, 2003.................62 3-26 Adults aerially collect ed, N=185, (Part A), reared adults, N=359, (Part B), and preserved larvae, N=651, (Part C), fro m collection 15, April 26-May 1, 2003.......63 3-27 Adults aerially collect ed, N=221, (Part A), reared adults, N=150, (Part B), and preserved larvae, N=388, (Part C), speci mens from collection 16, June 12-15, 2003..........................................................................................................................6 4

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x 3-28 Adults aerially collect ed, N=186, (Part A), reared adults, N=266, (Part B), and preserved larvae, N=648, (Part C), from collection 17, December 8-20, 2003.......65 3-29 Adults aerially collect ed, N=220, (Part A), reared adults, N=294, (Part B), and preserved larvae, N=347, (Part C), from collection 18, January 23-31, 2004.........66 3-30 Adults aerially collect ed, N=316, (Part A), reared adults, N=639, (Part B), and preserved larvae, N=675, (Part C) from collection 19, March 5-14, 2004.............67

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xi Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science THE FORENSICALLY IMPORTANT CALLIP HORIDAE (INSECTA: DIPTERA) OF PIG CARRION IN RURAL NORTH-CENTRAL FLORIDA By Susan V. Gruner August 2004 Chair: John Capinera Major Department: Entomology and Nematology The use of insect life stages in the determination of postmortem intervals in crime scene investigations is an important forens ic science tool used by coroners, medical examiners, and police investigators. For estimation of postmortem interval, basic distribution data for the major i ndicator species of insects are required. It is apparent that the seasonality and species assemblage va ry in different geographical areas. A study to determine possible indicator sp ecies of Calliphoridae present in rural north-central Florida was conducted using pig carrion as models representing human bodies. A wooded habitat was used as the s ite for placement of the pigs. The study involved 19 batches of pigs placed in a woode d site over a period of time including spring, summer, fall, and winter collections from November 16, 2001, to March 2004 (approximately monthly). Larval and adult ca lliphorid flies were collected, as were meteorological data rela ting to the study site.

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xii Seven species of Calliphoridae were co llected from the pig carrion. Relative abundance of each species as a percentage of the total adult Calliphoridae assemblage (% aerially collected/% r eared) for the study was Phaenicia coeruleiviridis 68.1 vs. 77.9%; Cochliomyia macellaria 16.0 vs. 8.5%; Chrysomya rufifaces 7.0 vs. 8.0 %; Phormia regina 8.2 vs. 3.9%; Chrysomya megacephala 0.3 vs. 1.6 %; Calliphora livida 0.4 vs. 0.1%; and Calliphora vicina 0.0 vs. 0.02%. There were obvious seasonal and successional variations of the species assemblage. Phaenicia coeruleiviridis (Macquart) was the predominant species year-round but was lower in abundance during the summer, mid-June to mid September. Only a few specimens of C. vicina Robineau –Desvoidy (= C. erythrocephala Meigen) and C. livida Hall were found during the coldest months, November to February, while C. megacephala (Fabricus) was collected during the hottest months, June to September. Cochliomyia macellaria (Fabricus) was found during the warm months, April to June, when the te mperature did not rise above 30.0 C. Chrysomya rufifaces (Macquart) was found in all but the coldes t months of the year, mid December to mid March. Phormia regina (Meigen) was not found during the winter. Different species of calliphorid flies arrived at the pig ca rrion at different stages during the decomposition process. Within minu tes of placing the pi g carcass on the ground, P. coeruleiviridis usually began to arrive. Cochliomyia macellaria C. rufifaces P. regina, C. vicina and C. livida arrived at the carcass afte r a delay of about 24 hours.

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1 CHAPTER 1 INTRODUCTION AND LITERATURE REVIEW Definition of and Scope of Forensic Entomology Forensic entomology is an extensive di scipline where arthropod science and the judicial system interact (Hall 2001). The fiel d of forensic entomology has been divided into three areas: medicocriminal entomo logy (also referred to as medicolegal entomology), urban entomology and stored product entomology. Information gained from medicolegal entomology typically is used to determine time of death, place of death and other issues of medical or legal importance (Gordh and Headrick 2001). Urban entomology concentrates mainly on controve rsies involving termites, cockroaches, and other insect problems accrui ng to the human environmen t, whereas stored product entomology involves disputes over arthropods and arthropod parts in food and other products (Hall 2001). When human remains are found, the most important questions usually are how, when, where and why the person died. Histor ically, determination of the postmortem interval (PMI) has been estimated through observation and measurement of body conditions such as core body temperature (Nelson 1999), muscular flaccidity, rigor mortis, lividity, pallor of the skin and others (Smith 1986, Bass 2001, Byrd and Castner 2001a). Entomological specimens in medicolega l death investigations can be reliable indicators for estimating the PMI in both early and advanced stages of cadaver decomposition (Nuorteva 1977, Smith 1986, Goff et al. 1988, Kashyap and Pillay 1989, Greenberg 1991, Byrd 1998).

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2 Insects and other invertebra tes feeding on carrion form a distinct faunal succession associated with the various stages of d ecay (Smith 1986). Recognition of the different immature stages of each species involved, t ogether with the knowledge of their rates of development, can give an indication of the PMI (Smith 1986). A forensic (= medicolegal) entomologist can also determine the age of immature insects, based upon knowledge of the variables regarding insect invasion of human remains. Evaluation and interpretation of entomological evidence at a crime scen e can address other complicated issues including season of death, ge ographic location of death, movement or storage of the remains following death, location of speci fic sites of trauma on the body, sexual molestation and use of drugs (Haskell et al. 1997). In case studies conducted in varying temp erate and tropical climates where human remains were exposed to the environment fo r 2.5 months or less, entomology-based PMI estimates differed by 48 hours when comp ared with the inte rvals determined by independent corroboration such as confe ssions and eyewitness testimony (Greenberg 1985, Goff et al. 1988, Lord 1990, Byrd 1998). Ento mological evidence is statistically the most reliable scientific means of estimati ng PMI when compared to other methods such as police reports and autopsy results (Kas hyap and Pillay 1989, Catts and Haskell 1990, Anderson 2001). History of Forensic Entomology The first documented forensic entomology cas e is from thirteenth century China in a book entitled “Hsi yan chi lu” which can be translated as “The Washing Away of Wrongs.” The author, Sung Tz’u, was an educ ated man. He was a doctor, a sheriff and eventually a Judicial Intendant. The book describes appli cations of forensic entomology used in criminal cases during that pe riod. A man was murdered by the roadside

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3 apparently by an assailant with a sickle. S ung Tz’u made a proclamation that the nearest neighbors were to bring all their sickles to him for examination (McKnight 1981). At inquest time, the weather was hot and blow f lies were attracted to one sickle only, even though it had no discernable traces of blood. The owner of the sickle confessed to the murder. In addition to medical and legal experts, sc ulptors, painters and poets have closely observed the decomposition of human bodies, noti ng, in particular, the effects of feeding maggots. Artwork from the Middle Ages accurately depicts the insect-mediated pattern of body mass reduction, particularly the early skel etonization of the skul l and the reduction of internal organs, with large parts of the skin left intact (Benecke 2001). In May 2004, a new painting of Prince Philip entitled “Portrai t of a Prince” was released by artist Stuart Pearson Wright. The painting shows Prince Phil ip with a bluebottle fly sitting on his left shoulder, which represents a memento mori; th e prince’s mortality (The Associated Press 2004). In 1855, Dr. Bergeret, a French physician, used insect succession as a tool (incorrectly) to solve a ca se (Benecke 2001). In the mid-1880s, J.P. Mgnin, also in France, published La Faune des Cadavres: A pplication de Entomologie la Medicin Legale. The recognition by Mgnin of a seque nce and progression of decomposition of a corpse was recorded in this work and in association with this decomposition progression, he observed changes in the insect assembla ges as the corpse aged (Haskell et al. 1997, Benecke 1998). This early interest in insects and deco mposition led to a study on insect succession on human corpses in Quebec, Canada, in 1897 by Wyatt Johnston and Geoffrey

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4 Villeneuve (Anderson 2001, Benecke 2001). At the same time in the United States, Murray Motter systematically tabulated the insect fauna from 150 exhumed corpses from the Washington, D.C. area (Haske ll et al. 1997, Benecke 2001). Species identification of th e most important fly groups, Calliphoridae (blow flies) and Sarcophagidae (flesh flies), used in fore nsic cases could not have been accomplished had it not been for Aldrich’ s (1916) monograph on the Sarco phagidae which illustrated the distinctive male genitalia of adult flie s. Knipling (1936) initiat ed taxonomic work on the larvae of sarcophagids and calliphorids. Hall’s 1948 book, The Blowflies of North America made it possible to identify the mature larvae of most species of calliphorids. In northern Europe, the blow fly Phaenicia sericata (Meigen) is the most economically important ectoparasite of do mesticated sheep. Sheep myiasis is a widespread disease and can cause high levels of mortality. The desire to develop control methods against sheep myiasis led to studi es of calliphorid attractants (Wardle 1921, Cragg and Thurston 1949, Hammack and Ho lt 1983, Ashworth and Wall 1994, Wall and Warnes 1994, Morris et al. 1998). The attractant studies prompted additional studies on blow fly distribution and ecology (Par ish and Cushing 1938, James 1947, Green 1951, Wolff et al. 2001) and were fo llowed by studies that addresse d effects of temperature on developmental time of blow fly life cycl es (Davidson 1944, Kamal 1958, Nuorteva 1977, Greenberg 1991, Byrd and Butler 1996, 1997, 1998). Regional successional studies of Calliphor idae in the United States have been conducted in California (James 1955), Hawaii (Goff et al. 1986, Goff et al. 1988, Goff 1991), Mississippi (Goddard and Lago 1985), Mi ssouri (Hall and Doisy 1993), Virginia (Hall and Townsend 1977), Indiana (Haskell 198 9), Illinois (Baumgartner 1988), Arizona

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5 (Deonier 1942, Baumgartner 1986, Gallowa y et al. 1989), Colorado (Adair 1999), Maryland (Introna et al. 1991), West Virginia (J oy et al. 2002), Louisiana (Tessmer et al. 1995, Watson and Carlton 2003), and South Caro lina (Tomberlin and Adler 1998). Four species--two of which are now found in Flor ida--of Old World blow flies have been confirmed from South or North Amer ica (Baumgartner and Greenberg 1984, Baumgartner 1986, Greenberg 1988, Tantawi an d Greenberg 1993, Martin et al. 1996): Chrysomya rufifaces (Maquart), C. albiceps (Wiedemann), C. megacephala (F.) and C. putoria (Wiedmann). Studies by Byrd (1998) and Peters (2003) were conducted in the Gainesville, FL (= north-central Florida) ar ea. This study will add rural north-central Florida to the list. The study of insects important to forens ic entomology has been conducted mainly through the use of non-human animal models Decomposition studies worldwide have used a variety of different carcass types and sizes, including dogs (Jiron and Cartin 1981, Early and Goff 1986, Richards and Goff 1997), cats (Early and Goff 1986), alligators (Watson and Carlton 2003), voles (Lane 1975), rats (Greenberg 1990, Tomberlin and Adler 1998, Faucherre et al. 1999, Kocarek 2001), squirrels (Johnson 1975), deer (Watson and Carlton 2003), foxes (East on and Smith 1970, Smith 1975), harbor seals (Lord and Burger 1984b), herring gulls (Lord and Burger 1984a), guinea pigs (Bornemissza 1957), mice (Putnam 1978, Blackith and Blackith 1989), lizards and toads (Cornaby 1974), raccoons (Joy et al. 2002), tu rtles (Abell et al. 1982), poultry (Hall and Doisy 1993, Tessmer et al. 1995), sheep (De onier 1940), rabbits (Denno and Cothran 1975, Tantawi et al. 1996, Bourel et al. 1999) elephants (Coe 1978), opossums (Goddard and Lago 1985), black bears (Anderson 1998, Peters 2003, Watson and Carlton 2003),

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6 impala (Braack 1981), and pigs (Payne 1965, Tullis and Goff 1987, Haskell 1989, Anderson and VanLaerhoven 1996, Tessmer a nd Meek 1996, Richards and Goff 1997, Byrd 1998, deCarvalho et al. 1999, Shahid et al. 1999, Davis and Goff 2000, deCarvalho and Linhares 2001, Wolff et al. 2001, Tenorio et al. 2003, Watson and Carlton 2003). The only faunal succession research on hu man remains was conducted in Tennessee (Rodriguez and Bass 1983, Catts and Haskell 1990). Human cadavers are not easily obtainable fo r detailed decomposition studies. Pigs, Sus scrofa are omnivorous, have similar gut fauna, are relatively hair less and have skin that is very similar to that of humans (Anderson and VanLaerhoven 1996). The putrefaction of pigs proceeds a pproximately at the same rate as for human bodies that are of the same weight (Campobasso et al 2001). Haskell’s 1989 study in Tennessee (Schoenly and Haskell 2000) compared the insect commun ity structure and decomposition rates between adult and infant human remains to a pig model and found no significant difference in the composition of the insect communities in human and pig carcasses (Campobasso et al. 2001). Therefor e, twenty-two kg pigs have been recommended as suitable human models for a dult decomposition (Catts and Goff 1992). Current Status of Forensic Entomology The popularity of television shows such as C.S.I. (Crime Scene Investigation), Forensic Files, and Court TV have created a recent surge of interest in the forensic sciences. Several colleges report long waiti ng lists for forensic science courses, and dozens of others are developing courses or entire programs in th e science of crime fighting (Lewerenz 2003). Purdue University ins tituted its first forensic science course in the fall of 2003, formatted by Neal Haske ll and Ralph Williams. Assuming that the course would not generate much enthusiasm a 25-student capacity room was assigned

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7 for the class. Once fall registration was comp leted, the room had to be changed to a lecture hall to accommodate the 425 students who registered for the class (Haskell 2003, personal communication). Lord and Stevenson’s 1986 directory (the only one ever published) of forensic entomologists listed only 62 scientists involved in this field of study; of the 62, only about a third were linked solely with the “m edicolegal” subdiscipline (Catts and Haskell 1990). In 1996, the American Board of Forensic Entomologists was created. Currently, there are only 8 members. However, forensic entomologists have no special group within the American Academy of Forensic Science or the Entomological Society of America. The European Association for Forensic Entomology was created in May, 2002, at the First European Forensic Entomology Se minar, held at the headquarters of the National Gendarmerie in Rosny sous Bois, France. This association was created to promote forensic entomology in Europe (H all 2003). In August, 2003, the first annual meeting of the American Association of Fore nsic Entomologists was held in Las Vegas, NV; approximately 45 people attended th e meeting. The second annual meeting is scheduled for July 24-27, 2004 and will be conducted at the University of CaliforniaDavis. The highly specialized field of forensic entomology has never had a large following. Several reasons may explain this lack of in terest, including: It involves having a close relationship with the larval stages of flies, commonly known as maggots. Most people think that these creatures, al ong with insects in general, are disgusting. Only a small number of colleges or universities offer a course in the specific field of forensic entomology, and none offer majors or minors.

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8 Historically, there has been little opportuni ty for full-time employment in this field. Biology of Calliphoridae Two major groups of insects are predictabl y attracted to cadav ers and provide the majority of information in forensic investiga tion; the flies and the beetles (Castner 2001). This research focused on the Family Calli phoridae, commonly cal led the blow flies (blowflies if you live outside th e USA), which are the first in sects to find and colonize human corpses. Experimental studies indicate th at these flies arrive at carcasses within minutes of their exposure (Byrd and Castner 2001b, Watson and Carlton 2003). There are more than 1000 species of bl ow flies throughout the world; about 90 species exist in North America (Haskell 2003, personal communication). This family includes the green bottle flies (genus Phaenicia ), blue bottle flies (genus Calliphora ), the screwworm flies (genus Cochliomyia ) and the black blow flies (tribe Phormiini). According to (Hall 1948), “to blow” is an ancient term that refers to depositing of eggs. The family name means ‘beauty bearer’ in Greek (Greenberg and Kunich 2002). Common blow flies carry at least 2,000,000 bacteria per specimen externally. Internally, each individual fly can carry from eight to ten times as many (Hall 1948). They can carry typhoid, chol era, the plague, anthrax, tuberculosis, tularemia, trypanosomes, leishmanias and can cause pr imary, secondary and tertiary myiasis (infestations of human skin). Some species of blow flies, such as Phaenicia sericata are used for cleaning of non-healing wounds. This healing method is call ed “maggot therapy” and has been in use for hundreds of years (Sherman and Pechter 1988). Sterile maggots debride the wounds by eating necrotic tissue. Urea, ammonium car bonate and allantoin se creted by the larvae disinfect deep tissue wounds and healing is stimulated. Maggot thera py is appropriate for

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9 cases where antibiotics are ineffective and su rgery is impracticable (Sherman and Pechter 1988). Calliphorid flies have highly specialized sense organs on their antennae that are stimulated by putrefaction odors and gase s which are released during post-mortem decomposition of organic matter. Some species of Phaenicia are attracted to various organic sulphur compounds, either alone or in combination with hydrogen sulphide, and also by ammonia (Cragg 1956, Cragg and Co le 1956, Ashworth and Wall 1994, Wall and Warnes 1994). Flight traps baited with dime thyl trisulphide are strong attractants for some calliphorids (Nilssen et al. 1996). Odors from Proteus mirabilis Hauser, a bacterium that causes infections in the fleece of sheep, are attractants to some calliphorid flies (Morris et al. 1998). Some plants have developed a pollination st rategy that targets calliphorid flies. The dead-horse arum ( Helicodiceros muscivorus L. fil.) floret emits an odor that smells like a dead animal. Blow flies are deceived into pollinating the plant. The volatile compounds of the dead-horse arum and of a carcass we re identified by gas chromatography as three structurally similar oligosulphides: dimethyl mono-, diand trisulphide. When calliphorid flies were exposed to each of the odors, iden tical antennal response pa tterns were elicited (Stensmyr et al. 2002). Landing behavior of calliphorids may be aff ected by visual cues such as white and yellow colors (Wall et al. 1992, Hall et al. 1995 ). Oviposition is elicited primarily by the presence of ammonia-rich compounds, mo isture, pheromones, and tactile stimuli (Ashworth and Wall 1994) yet is rarely stimulated by chemicals alone (Cragg 1956).

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10 Unfortunately, the complex interaction of semiochemical and visual cues used for resource location remains little studied in calliphorids (Wall and Fisher 2001). Blow flies are heliotropic and usually rest at night. Eggs are not usually laid at night although clearly ther e are exceptions. Green (1951, page 484) observed that Calliphora deposited eggs at night unde r artificial light in slaught er houses. He wrote that “under laboratory conditions it has been found that Calliphora erythrocephala (now called C. vicina ), Lucilia sericata and Phormia terrae-novae will all oviposit in total darkness, although Wardle (1921) asserts that blowflies do not oviposit in the complete absence of light. Greenberg (1990) observed that Phaenicia sericata Phormia regina (Meigen) and Calliphora vicina (Robineau-Desvoidy) oviposite d a very small number of eggs on rat carrion at night. Singh (2001) pointed out that the flies in Greenberg’s experiment probably were resting on a nearby bush and literally cr awled over to oviposit on the rat carrion, thus indicati ng that blow flies were not actively searching for an oviposition site. Nocturnal ovipos ition has not been observed in large-scale studies in other areas (Greenberg 1990, Byrd a nd Butler 1997, Haskell et al. 1997). Other factors that affect blow fly activity are temperature, size of the carcass, geographical location, humidity, light and shade, seasonal and daily periodicity, availability of food and competition, maggot mass temperature and manner of death (Rodriguez and Bass 1983). Insect Succession on Carrion The first organisms to arrive on a body af ter death are usually the insects. They arrive at predictable times during the decom position process. Each decomposition stage is attractive to a different group of sarcosaprophagous arthropod s. Smith (1986, page 13), in

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11 A Manual of Forensic Entomology defines four ecological categories in the carrion community: 1. Necrophagous species: Feed on the carrion itse lf and constitute the most important category in establishing time of death, e.g., Diptera: Calliphoridae (blow flies); Coleoptera (beetles): Silphidae (in part), Dermestidae. 2. Predators and parasites: Second most im portant forensic cate gory, e.g., Coleoptera: Silphidae (in part), Staphy linidae; Diptera: some carrion feeders become predace ous in later instars, e.g., Chrysomya (Calliphoridae), Ophyra and Hydrotaea (Muscidae) on the necrophagous species. 3. Omnivorous species: Wasps, ants and some beetles feed both on the corpse and its inhabitants. 4. Adventive species Use th e corpse as an extension of their environment, e.g. Collembola (springtails), spiders (which may become incidental predators). Blow flies are attracted by the odors and ga ses which are released during the onset of autolysis and putrefaction, depending on tim e of year and situation of the corpse (Smith 1986). As decomposition proceeds, the odors emanating from the corpse change, making the cadaver more attractive to some spec ies of blow flies and less attractive to others. Once the dry decay stage has been re ached, blow flies are no longer attracted to the corpse (Nuorteva 1977, Anderson 2001). Af ter the invasion of North America by the Chrysomyinae in the 1980’s, the blow fly sequence in North America may be Phaenicia Cynomyopsis Chrysomyinae Calliphora and Cochliomyia (Campobasso et al. 2001). Obviously, species lists will differ by region. Beginning with Mgnin’s (1894) work, eight waves of arthropod invasion on human bodies have been described. Other fore nsic entomologists reduced the number of stages in attempts to define biologica l communities, but ultimately this reduction complicated and lessened the forensic appli cability. Payne (1965) defined the associated

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12 insect community and analyzed the percenta ge abundance of species attracted to the various stages of decay. He condensed eight to six stages of decay: fresh, bloated, active, advanced, dry and remains. Lord and Burger (1984a) and Bornemissza (1957) recognized five stages of carcass decomposition. Accordi ng to Smith (1986, page 17), there exists a broad general agreement in the observati ons of Mgnin, Bornemissza (1957), Reed (1958) and the series of publications by Payne (1965) and (Payne 1965, Payne and King 1970) as follows: 1. Initial decay stage (0-2 days). Carcass appears fresh externally but is decomposing internally due to the activities of bacter ia, protozoa and nematodes present in the animal before death. 2. Black putrefaction stage (1220 days). Flesh of creamy consistency with exposed parts black. Body collapses as gases escape. Odor of decay very strong. 3. Butyric fermentation stage (20-40 days). Ca rcass drying out. Some flesh remains at first and cheesy odor develops. Ventral surface of body moldy from fermentation. 4. Dry decay stage (40-50 days). Carcass almost dry; slow rate of decay. The aforementioned stages of decay ar e not easy to delineate and there is controversy regarding these defi nitions, but they are useful in describing the sequence of decomposition. For example, the number of days varies considerably with temperature. Factors that Affect Blow Fly Succession on Carrion Temperature and access to a body are the tw o most important factors affecting insect succession. Temperature is the most im portant variable influencing the rate of maggot development. High temperatures genera lly reduce development time of Diptera. Large aggregations of dipteran larvae (maggot masses) develop heat due to their frenetic activity and fast metabolism, thus rais ing the microenvironmental temperature (Campobasso et al. 2001). The heat of the ma ggot mass is related to the density of the mass and the size of the carcass (weight and mass). The size of maggot masses and the

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13 degree to which the corpse is either expos ed to, or insulated from, the environment affects the amount of heat abso rbed or dissipated, which in turn has a significant effect upon the rate of larval development and the decomposition of a corpse. Goodbrod and Goff (1990) studied the effects of maggot-gener ated heat during the development cycle in experimental cultures of C. megacephala and C. rufifaces and found an inverse relationship between density and th e duration of larval stage. The insects that colonize corpses vary in species depending on the biogeoclimatic zone in which the remains are found. Each zone has different habita t types, vegetation, soil pH, soil type, flora and fauna, altitude a nd climatic conditions th at affect the species of insects present. Decomposition also is aff ected by the time of year, and the location in which remains are found (Anderson 2001). Many blow fly species vary in abundance depending on season and even time of day. Presence or absence of sunlight or shade can have an effect on which blow fly species will colonize a corpse. Cragg (1956) demonstrated that P. sericata prefer heated surfaces and will not oviposit on carcasses that have surface temperatures below 30 C. Results of a sun-exposed versus shaded pig carrion study indicated that more Lucilia illustris (Meigen) and Phormia regina (Meigen) were observed at the sun-exposed pig whereas Calliphora vomitoria (Linnaeus) were observed in greater numbers at the shaded pig (Shean et al. 1993). Blow flies can be found in both urban a nd rural areas but some species may be found only in wooded areas. Flies primarily asso ciated with human refuse (synanthropy) are usually found in urban areas. Presence of certain species of blow flies found on a body may indicate that the body was moved from an urban to a rural environment or vice

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14 versa (Erzinclioglu 1985, Catts and Haskell 199 0). Blow flies are capable of colonizing corpses inside dwellings and cars, depending upon how well they are sealed. Competition for the food source is the most important factor affecting size and completion of the life history of carrion. Smith (1986, page 34) listed ways in which this occurs: 1. Intraspecific competition may reduce the size of the larvae and can reduce the number and fecundity of individual adults 2. Interspecific competition has similar results as to intraspecific competition, plus a possibility of total elimination 3. Predators and parasites-selective predation or parasitization of one species can be advantageous to a competing species. The early arrivers (such as Calliphora and Phaenicia ) at a corpse may have an advantage. Some female blow flies (Tribe Calliphorini) are capabl e of moving a single egg from one of the ovaries into the vagi na (termed “precocious egg development”) where it is fertilized before an ovipositi on site has been found (Wells and King 2001). Delayed arrivers such as Chrysomya may compensate for this by being viviparous and deposit larvae on the carcass. Some dipteran larvae begin life as carrion feeders and become predators after the second molt. Biology of Human Decomposition Campobasso et al. (2001, page 18) descri be the decomposition of corpses as a mixed process ranging from autolysis of individual cells by internal chemical breakdown, to tissue autolysis from liberated enzymes and from external processes introduced by bacteria and fungi from bot h the intestine and outer environment. The bacterial enzymatic structures caus e putrid liquefaction of tissues by the breakdown of proteins, carbohydrates and lipids into their basic components (amino acids, water and carbon dioxide, fat acids and volatile substances) with gas formation (nitrogen, methane, hydrogen sul phide, ammonia, etc.). As the tissues are digested to a fluid consistency with the production of la rge amounts of foulsmelling gas; they become moist and gasridden, and eventually liquefy down to the skeleton.

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15 Calliphorid larvae hasten tissue decay by di ssemination of bacteria. The larvae also have digestive enzymes in their saliva that liquefy carcass tissues. As the larvae burrow into and consume the carcass, tissues furthe r disintegrate (Easton and Smith 1970). Usually, tissues that conduct the highest ra tes of ATP synthesis, biosynthesis, and membrane transport decompose first. The intestines begin to decompose first, followed by the stomach, accessory organs of digestion, heart, blood and circulation, heart muscle, air passages and lungs, kidneys and bladder, brain and ner vous tissues, skeletal muscles and finally, connective tissues a nd integument (Gill -King 1997). Over a seven-year period, researcher s at the Anthropology Research Facility (ARF—also known as The Body Farm) in east Tennessee studied 150 bodies in various stages of decay, including homicide victims, bodies donated to scie nce, and unidentified persons. They found that variables affecting d ecay rates of human bodies are (in order of importance): temperature, access to the body by insects, burial and depth, carnivore and rodent activity, trauma (penetrating/crushing), humidity, rainfall, size and weight of the body, embalming, clothing, type of surface body was placed on and the soil pH (Mann et al. 1990). Further Areas for Study Entomological studies of Calliphoridae a ssociated with carrion should include investigations of seasonal changes in th e fauna and successional events which are influenced by weather factors. Data ar e needed from different geographic regions including species compositi on of sarcosaprophagous ar thropods and the weather parameters associated with those fauna (i ncluding their maximum/minimum threshold

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16 temperature data). Reliable taxonomic keys are needed for identifica tion of early larval instars and of puparia. The assessment of species composition within rural north-central Florida is addressed in the present study by determini ng the calliphorid species associated with carrion in north-central Florida and the influence of seasons on the calliphorid species assemblage. The objectives of the study were: 1. To identify the species of Calliphoridae asso ciated with pig car rion in rural northcentral Florida. 2. To determine how season (spring, summer, fall and winter) affects the assemblage of calliphorid species associated with pig carrion in rural north-central Florida. 3. To determine the daily succession of cal liphorid species associated with pig carrion during each collection period in rural north-central Florida.

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17 CHAPTER 2 MATERIALS AND METHODS The use of human corpses for field studies is illegal in Florida. As determined in other decomposition studies (Haske ll 1989, Anderson and VanLaerhoven 1996, Campobasso et al. 2001) the rate of pig decom position is very similar to that of humans. Dead pigs, Sus scrofa L., were used as animal models for this study. All pigs were purchased from North Florid a Livestock Market, Lake City, Florida. Prior to purchase, the pigs were killed by a late ral, transverse shot in to the top of the head with a .22 caliber rifle. This method resulted in instant death of the animal. Each dead pig was immediately double-bagged in a heavy-dut y plastic trash bag and was transported from Lake City to the Greathouse Butte rfly Farm in Earleton, Florida. Study Site Greathouse Butterfly Farm is located 19.3 km east of Gainesville, near Earleton, on the southeast corner of N.E. SR 26 and SR 1469. The property consists of 48.6 hectares of north Florida flatwoods (Figure 2-1). Pig car casses were placed in a wooded habitat where sunlight was somewhat restricted. In so me cases, the pigs received direct sunlight during certain parts of the day, while at ot her times they were shaded. The study site mainly consisted of a moderate stand of live oak, Quercus virginiana Mill., and slash pine, Pinus elliottii Engelman with an understory of various saw palmettos and grasses (Florida Chapter 1989). Pigs were placed at least 18.3 meters ap art. Wire cages (86.5 cm long x 50.8 cm wide x 61 cm high) were placed over the pigs to protect them from large vertebrate

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18 scavengers. The cages were cons tructed of wire mesh (5 cm x 5 cm). At least four metal or plastic tent stakes were driven into the ground around th e cage, and bungee cords were attached tightly to the cages and then to tent stakes to minimize disturbance (Figure 2-2). The cages were lifted off and set aside during sampling times. Data Collection Trials were conducted from October 14, 2001 to March 5, 2004; however, data from October 2001 were incomplete and ther efore not included. Th e raw data, starting with the second collection on November, 2001, ar e in appendices A and B. Each trial consisted of 3-4 pigs, with trials conducted throughout the year (Table 2-1). Observations and collections were made daily during the af ternoon, if possible, when flies were most active. The duration of each tria l lasted until the first wave of maggot dispersal occurred (termed “maggot migration” by forensic entomol ogists). After the thirdinstar larvae have reached a certain size, they leave the carcass en mass to pupa riate in the soil. The pigs were not moved or disturbed in any way duri ng the study. Digital pictures of the pigs and of the expanding maggot masses were taken at each collection time. At the beginning of each collection time, a laminated identificati on sheet was placed in front of each pig, indicating the date, time and identification number. A meter-stick was placed below the identification sheet to give document size. Sa mpling usually occurred in the middle of the afternoon when the adult f lies were most active. Protocol for Day 1 On day 1, the pigs were removed from bags and placed in a location on the site, and the GPS (global positioning system) was noted. After all the pigs we re in position, data were collected including pig number, time of death (TOD), date, time, sample number (SA#), ambient temperature (AT), gr ound/pig interface temperature (GPI), ground

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19 temperature at 5 cm depth about 3 m from the pig, a brief description of the weather (sunny, cloudy, rain, etc.) and wind velocity in meters per second (M/S). An aerial collection of adult flies was made over each pig with an insect net, with a target of at least 10 adult calliphorid flie s (although this was not always possible to achieve). Flies were placed in vials with 70% isopropyl alcoho l and later pinned and identified. A data logger (F igure 2-3) was affixed to a nearby tree or bush and the temperature probe was placed on the ground 12 m from each pig. Therefore, the ground temperature being recorded was similar to that which the pig was exposed. Protocol for Day 2 Onward Adult flies were sampled with a net (if possible) and paired samples of approximately 50-500 larvae (L = live, P = pres erved) were collected from the growing maggot mass, usually located on the head. Th e location where the sample was taken on the body area (or ground area) was noted on the collection sheet. About half of the sample specimens were boiled in water for about 2 minutes using a camp stove, and then placed in vials with 70% isopropyl alcohol for preservation. The remainder of larvae were placed in containers and reared to the adult stage in order to compare the identity and relative abundance of adult and larval flies. Meteorological Measurements Temperatures determined with a Tayl or 9841 digital thermometer (Forestry Suppliers, Jackson, MS) shielded from direct rays of the sun (w hen necessary) include ambient air, ground/pig interface, external tissue, oral cavity of the pig, soil at 5 cm depth 3 m from the pig, and soil at 5 cm depth under the maggot mass. Maggot mass temperatures were taken (MM T) but will not be addressed in this thesis. Ground temperatures were taken with HOBO (O nset Computer Corp., Bourne, MA) data

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20 loggers. These units were not 100% waterproof so a plastic housing was made for each of the loggers (Figure 2-3). Three hangars we re cut and cemented with marine glue into one end of each container. A hole was drille d at the bottom of the container (for the temperature probe cable) and foam was packed around each logger. The HOBO unit was set to record temperature every 30 mi nutes for 3 weeks. A temperature probe connected to each logger was set on the ground within 1-2 m of each pig (Figure 2-4). Table 2-4 lists the mean temperatures that were taken during th e collection intervals during the months of the proj ect. The mean temperatures were taken with the Taylor thermometer during the collection times. The temperature at 3:56 AM, taken from the data logger’s data, was used as the mean low temperature (Figures 3-5 A and B). There was no solar interference at this time. For collections 2 and 3, the data logger was not used and the mean low temperatures were taken from the NOAA (National Oceanic and Atmospheric Administration) daily data availa ble for Gainesville Re gional Airport. Rearing Procedures Maggots were removed from the carcass with a plastic spoon and placed immediately into rearing pouches constructe d from aluminum foil (Catts and Haskell 1990) and placed into Rubbermaid 28 L or plas tic Gladware containe rs half-filled with substrate for pupation. The container lids e ach had approximately ten 2-mm holes randomly drilled for air circulation. A paper towel was placed under each lid to prevent larvae/adult flies from escaping through lids and also to prevent other flies from getting into the containers. Each foil pouch was filed with 100-200 g of calves’ liver, which served as food. The containers of live maggots were kept outside—except when temperatures were below 4.5C—on shelving placed in a screened enclosure. The maggots were checked

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21 daily. Once the maggots migrated off the liver, the pouch was removed from the container. On occasion, the maggots would not move out of the foil pouch. When this occurred, a small amount of fresh liver was pl aced directly onto th e substrate, and the maggots were gently moved from the foil pouch onto the liver. The liver would be removed a day or two later after the maggots migrated into the substrate to pupate. At adult emergence, the containers were placed into the freezer for 15 minutes to kill the flies. Adults were put in labeled vials of 70% isopropyl alcohol, then later pinned and identified (White et al. 1940, Dodge 1953, Seago 1953, Furman and Catts 1982, Wells et al. 1999). The preserved maggots we re separated into first, first-second transitional, second, second-third transitional a nd third instars. Only third-instar larvae were identified to species because there ar e no reliable taxonomic keys for firstand second-instar calliphorid larval identification. Gary Steck, dipt erist at Florida Division of Plant Industry (DPI, Department of Agricultur e and Consumer Servi ces, Gainesville, FL), verified the identifications of representative samples of the flies. All data were entered into a Microsoft Office XP Excel versi on 2002 spreadsheet (Microsoft Corp.). The percentages of adult and larval specimens were transformed to their square roots to run the Pearson’s correlation analysis. The raw data are presented in Appendices 1 and 2. Voucher specimens are being kept in lab 2209 at the University of Fl orida Department of Entomology and Nematology and will be deposited with the museum at DPI. Pupation Substrate Pupation substrate consisted of about 90% dried laurel oak leaves ( Quercus laurifolia Michx.) and branches, 2 % rye grass ( Lolium perenne Lam.) cuttings, 8 % magnolia leaves ( Magnolia grandiflora L.), and a minute amount of sandy soil. The plant mixture was ground up with a Troy-Bilt M odel 4731-10 HP chipper/shredder with a

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22 mulching blade. The substrate was stored in a 200 L (liter) Rubbermaid trash container with a tight fitting top. On most occasions, larvae that migrated into the substrate successfully produced adults in containers half-filled with substrate. On a few occasions (less than 3), no adults were obtained due to mortality from fungi. No information is available; however, on the proportion of pupae that successfully reached adulthood.

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23 Figure 2-1. A caged pig carcass is shown at Greathouse Butterfly Farm property in Earleton, Florida.

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24 Figure 2-2. Wire cage with bungee cords and te nt stakes hammered into the ground. The cage protected the carcass from scavengers.

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25 Figure 2-3. A HOBO temperatur e data logger is sealed in side a Gladware container. Data loggers (one for each pig) were used to obtain ground temperatures near pig carcasses at Greathouse Butterfly Farm, Earleton, Florida.

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26 Figure 2-4. The data logger was hung from bush at left and temperat ure probe was placed on the ground (circled) to record gro und temperature near the pig carcass in Earleton, Florida.

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27 Table 2-1. Dates of pig deposition, appr oximate pig weight, and time each pig was placed on site for sampling dates from November, 2001 to October, 2002. Date of Pig Deposition Approx. pig weight (kg)Numb er of pigs per dateApprox. deposition time (h) 15-Nov-20012031730 23-Dec2031500 1-Feb-20022541530 15-Mar2441545 29-Apr3041445 20-May3241630 22-Jul2041550 19-Aug3041445 23-Sep2041450 24-Oct2041730 27-Nov2341500 30-Dec2531530 28-Feb-20032131545 31-Mar1941445 25-Apr2831630 12-Jun2231550 8-Dec2831440 23-Jan-20041531545 5-Mar2821530

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28 Table 2-2. Mean temperatures ( SD) during the study at the Earlet on, Florida site. Mean temperatures were taken during collection times with the Taylor digital thermometer while facing away from s un. Mean low temperatures were taken from the HOBO data at the 3:56 AM mark. November 2001 through January 2002 temperature data were obtained from National Oceanic and Atmospheric Administration. Mean Temp. CSDMean Low Temp. C SD Nov. 16-21, 200123.61.15.22.8 Dec. 29, 2001-Jan.11, 200220.23.20.95.8 February 5-9, 200219.94.18.14.5 March 15-19, 200230.41.216.53.0 April 29 -May 1, 200232.81.119.41.5 May 20-23, 200225.90.917.43.2 July 22-25, 2002 33.21.322.40.7 Aug. 19-23, 200230.52.923.91.3 September 23-27, 200229.33.223.71.7 October 26-28, 200228.42.219.10.5 Nov. 30-December 14, 200216.84.09.74.3 December 30, 2002-January 11, 200317.03.35.75.2 March 2-8, 200322.16.216.93.5 April 1-6, 200325.05.610.85.9 April 26-May 1, 200327.43.918.41.1 June 12-15, 200330.31.323.01.1 Dec. 8-20, 200317.43.49.44.0 January 23-31, 200418.94.58.95.6 March 5-14, 200424.54.613.35.4

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29 CHAPTER 3 RESULTS Species of Calliphoridae Co llected on Pig Carrion The seven species of Calliphoridae colle cted on pig carrion between November 16, 2001 and March 14, 2004 in rural north-central Florida were Phaenicia coeruleiviridis (Macquart), Cochliomyia macellaria (Fabricus), Chrysomya rufifaces (Macquart), Phormia regina (Meigen), Chrysomya megacephala (Fabricus), Calliphora livida Hall, and Calliphora vicina Robineau–Desvoidy (= C. erythrocephala Meigen) (Figure 3-1). Assessment of the fly assemblage by three collection methods (a erial collection of adults, adults reared from larvae, and preser ved larvae) produced similar results (Figure 3-1). Pearson’s correlation anal ysis of aerial collections made when adult flies were most active (usually during the early afternoon) indicated a high degree of correlation with maggot abundance for the same time period. The analysis of aerial a nd reared specimens indicated a high degree of correlation (r = 0.9512, 95% confidence interval = 0.6981 to 0.9930, two-tailed P= 0.0010). The analysis of aerial and preserved specimens (r = 0.9744, 95 % confidence interval 0.8315 to 0.9964, two-tailed P = 0.0002) also indicated a high degree of correlation. Finally, the analys is of reared and preserved specimens (r = 0.9783, 95% confidence interval 0.8557 to 0.9969, two-tailed P = 0.0001) indicated a high degree of correlation. It is mu ch easier to catch adult flies than it is to rear larvae, so an aerial sample taken during the middle of the day represents a good estimation of species present, although, in practice, both adults and la rvae should continue to be collected for legal reasons.

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30 Preserved larvae collected dur ing the study were identified and counted by instar. Examined through a microscope, each of the diffe rent instars is easily recognizable. Firstinstar larvae have Y-V shaped spiracular slits (Figures 3-2 A and B). First-second transitional instars have a second set of spir acles behind the Y-V slits (Figure 3-3). The second-instar larvae have two inner spiracu lar slits (Figure 3-4). The second-third transitional instars have two sets of spiracular slits (Figure 3-5), and the third instars have three inner spiracular slits (Figure 3-6). Chrysomya rufifaces larvae are recognizable without the aid of a microscope ; they have rows of conspi cuous tubercles (Figure 3-7). Chrysomya megacephala larvae have a distinguishing acc essory oral sclerite, but none were collected in this study. A total of 23,960 larvae were collected. Of these, 8253 were third-instar larvae and thei r relative abundance was: P. coeruleiviridis 76.5%; C. macellaria 7.3%; C. rufifaces 9.1%; P. regina 6.8%; and 0.3% were unknown (Figures 3-1 and 3-8). Seasonal Distribution and Succession of Calliphoridae Species Adult calliphorids colle cted and reared in year 1 (N=3197) and year 2 (N= 3992) are shown in Figures 3.9 and 3.10, respectivel y. Differences in seasonal phenology are evident (spring is March 20 to June 20; su mmer is June 21 to September 21; fall is September 22 to December 20; winter is December 21 to March 19). The mean temperatures taken during afternoon samples and the mean low temperatures for year 1 and year 2 of the study are in Figures 3-11 A and B. Two species, Calliphora livida and Calliphora vicina were found only during the winter, from mid-December to mid-March (Figs. 3-9 and 3-10). One species, Chrysomya megacephala was found only in the summer, from mid-June to late-S eptember (Figs. 3-9 and 3-10). Cochliomyia macellaria C. rufifaces and P. regina and others were not found duri ng the winter (Figs. 3-9 and 3-

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31 10). Phaenicia coeruleiviridis generally were found in great abundance year round, but were lower in abundance during the summer (Figures 3-9 and 3-10). Different species of calliphorid flie s arrived at the pig carri on at different stages during the decomposition process. Within minu tes of removing the pig from the plastic bag and placing it on the ground P. coeruleiviridis usually began to arrive. Cochliomyia macellaria C. rufifaces P. regina C. livida and C. vicina arrived at the carcass after a delay of about 24 hours. The data below are discussed in terms of total insects co llected, reared, or preserved per collection interval. The figures are shown graphically in an effort to evaluate succession. Collection 1, November 16-21, 2001 No adult calliphorids were ae rially collected. Of the to tal reared adults (N=232), 94% were P. coeruleiviridis, 5.6% C. rufifaces and 0.4% were C. macellaria (Figure 3-12 A). One hundred percent of the preserve d third-instar la rvae (N=30) were P. coeruleiviridis (Figure 3-12 B). Collection 2, December 29, 2001-January 11, 2002 No adult calliphorids were ae rially collected. Of the to tal reared adults (N=362), 99.4 % were P. coeruleiviridis while C. livida and C. vicina each comprised of 0.3% (Figure 3-13 A). Preserved la rvae (N=568) consisted of 79.8% P. coeruleiviridis, 18.0% P. regina and 2.3% were unknown (Figure 3-13 B). Collection 3, February 5-9, 2002 One hundred percent of the a dults aerially co llected (N=39) and adults reared (N=305) were P. coeruleiviridis (Figures 3-14 A and B). Preserved larvae (N=151) consisted of 70.9% P. coeruleiviridis and 29.1% P. regina (Figure 3-14 C).

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32 Collection 4, March 15-19, 2002 Adults aerially collected (N =71) consisted of 97.2% P. coeruleiviridis and 2.8% P. regina (Figure 3-15 A). Of the reared adults (N=326), 100% were P. coeruleiviridis (Figure 3-15 B). Preserved larvae (N=134) consisted of 97.8% P. coeruleiviridis and 2.2% were unknown (Figures 3-15 C). Collection 5, April 29-May 1, 2002 Adults aerially co llected (N=107) co nsisted of 86.0% P. coeruleiviridis 0.9% C. macellaria and 13.1% P. regina (Figure 3-16 A). Of the reared adults (N=332), 90.1% were P. coeruleiviridis 2.7% were C. macellaria and 7.2% were P. regina (Figure 3-16 B). Preserved larvae (N=196) consisted of 100% P. coeruleiviridis (Figure 3-16 C). Collection 6, May 20-23, 2002 Figure 3-17 A illustrates successional data. Four species of calliphorids were captured on the first day. Cochliomyia macellaria was the most abundant species, but apparently they were not depositing eggs on the first day because the larvae collected on day 3 were 100 % P. coeruleiviridis The abundance of P. coeruleiviridis declined (adults and larvae) as decomposition progressed. The abundance of C. rufifaces remained about equal, but P. regina increased in abundance as d ecomposition progressed. Aerially collected specimens (N=51) consisted of 17.6% P. coeruleiviridis 39.2% C. macellaria 21.6% C. rufifaces and 21.6% P. regina (Figure 3-17 A). Of the reared adults (N=67) and preserved larvae (N=175), 100% were P. coeruleiviridis (Figures 3-17 B and C). Collection 7, July 22-25, 2002 Figures 3-18 A-C illustrate successional data. P. coeruleiviridis was the most abundant calliphorid present on day 1 and declined in abundance as decomposition progressed while C. macellaria C. rufifaces and P. regina arrived in fewer numbers on

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33 day 1 but increased in abundance as decomposition progressed. Chrysomya megacephala arrived when the carcass was in the active d ecay state on day 3. Adults aerially collected (N=99) consisted of 58.6% P. coeruleiviridis 5.1% C. macellaria 31.3% C. rufifaces 2.0% P. regina and 3.0% C. megacephala (Figure 3-18 A). Of the reared adults (N=223), 47.1% were P. coeruleiviridis and 52.9% were C. rufifaces (Figure 3-18 B). Preserved larvae (N=700 ) consisted of 31.6% P. coeruleiviridis and 68.4% C. rufifaces (Figure 3-18 C). Collection 8, August 19-23, 2002 Figures 3-19 A-C illustrate successional data. Phaenicia coeruleiviridis was the most abundant species for the first 2 days but decreased in abundance as decomposition progressed. Chrysomya rufifaces and C. macellaria did not arrive at the carcass until day 2 (Figure 3-19 A), but increased in abundance as decomposition progressed. Chrysomya megacephala arrived when the carcass was in the ac tive decay state, after 3 days. Adults aerially collected (N=118) consisted of 67.8% P. coeruleiviridis 6.8% C. macellaria 23.7% C. rufifaces and 1.7% C. megacephala (Figure 3-19 A). Of the reared adults (N=248), 43.1% were P. coeruleiviridis 1.2% C. macellaria 42.3% C. rufifaces and 13.3% C. megacephala (Figure 3-19 B). Preserved larvae (N=499) consisted of 83.0% P. coeruleiviridis 0.4% C. macellaria 13.6% C. rufifaces 2.8% P. regina and 0.2% were unknown (Figure 3-19 C). Collection 9, September 23-27, 2002 Figures 3-20 A-C illustrate the successional data. Phaenicia coeruleiviridis was the only calliphorids species collected on day 1 and abundance declined as decomposition progressed. Cochliomyia macellaria and C. rufifaces arrived at the carcass on day 2, and increased in abundance as decomposition progressed. Chrysomya megacephala adults

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34 were reared from the September 26 larval sa mple, but no adults were aerially collected (Figure 3-20 B). Adults aerially collected (N=76) consisted of 44.7% P. coeruleiviridis 34.2% C. macellaria and 21.1% C. rufifaces (Figure 3-20 A). Of the reared adults (N=323), 54.2% were Phaenicia coeruleiviridis 30.7% C. macellaria 1.2% C. rufifaces and 13.0% C. megacephala (Figure 3-20 B). Preserved la rvae (N=697) consisted of 66.1% P. coeruleiviridis 21.5% C. macellaria 11.3% C. rufifaces and 1.0% P. regina (Figure 3-20 C). No P. regina adults were aerially collected or reared (Figure 3-20 B). Collection 10, October 26-28, 2002 Figures 3-21 A-C illustrate successional data. Phaenicia coeruleiviridis C. macellaria and C. rufifaces were collected on day 1. Phaenicia coeruleiviridis was not the most abundant species of adults on da y 1, but was the most dominant species of larvae. Their abundance declined as decompos ition progressed. Adults aerially collected (N=87) consisted of 12.6% P. coeruleiviridis 35.6% C. macellaria and 51.7% C. rufifaces (Figure 3-21 A) Of the reared adults (N=131), 52.7% were P. coeruleiviridis 7.6% C. macellaria and 39.7% C. rufifaces (Figure 3-15 B). Preserved larvae (N=294) consisted of 59.2% P. coeruleiviridis 8.8% C. macellaria and 32.0% C. rufifaces (Figure 3-21 C). Collection 11, November 30-December 14, 2002 Phaenicia coeruleiviridis was the most dominant calli phorid species during this sampling period. Adults aerially collected specimens (N=60) consisted of 91.7% P. coeruleiviridis 1.7% C. macellaria 5.0% C. rufifaces and 1.7% P. regina (Figure 3-22 A). Reared adults (N=3 51) consisted of 88.0% P. coeruleiviridis and 12.0% P. coeruleiviridis 19.0% C. macellaria (Figure 3-22 B). Preserved specimens (N=1248) consisted of 99.8% P. coeruleiviridis and 0.2% C. macellaria (Figure 3-22 C).

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35 Collection 12, December 30, 2002-January 11, 2003 Phaenicia coeruleiviridis was the most dominant calli phorid species during this sampling period. Adults aerially coll ected (N=56) consisted of 98.2% P. coeruleiviridis and 1.8 % C. livida (Figure 3-23 A). Reared adults (N=371) consisted of 99.2% P. coeruleiviridis and 0.8% C. livida (Figure 3-23 B). Preserve d larvae (N=1248) were comprised of 99.8% P. coeruleiviridis 0.2% C. macellaria and 1.0% were unknown (Figure 3-23 C). Collection 13, March 2-8, 2003 Phaenicia coeruleiviridis was the most dominant calli phorid species during this sampling period. Adults aerially coll ected (N=66) consisted of 87.9% P. coeruleiviridis 9.1% P. regina and 3.0% C. livida (Figure 3-24 A). Reared ad ults (N=166) consisted of 98.2% P. coeruleiviridis and 1.8% P. regina (Figure 3-24 B). Preser ved larvae (N=340) consisted of 71.5% P. coeruleiviridis 28.2% P. regina and 0.3% were unknown (Figure 3-24 C). Collection 14, April 1-6, 2003 Figures 3-25 A-C illustrate successional data. Phaenicia coeruleiviridis was the most dominant calliphorid species for the firs t 4 days of the sample period but decreased in abundance as decomposition progressed. Cochliomyia macellaria C. rufifaces and P. regina arrived after a delay of about 24 hours and increased in abundance as decomposition progressed. Adults aerially collected (N=247) consisted of 75.3% P. coeruleiviridis 19.0% C. macellaria and 5.7% P. regina (Figure 3-25 A) Reared adults (N=229) consisted of 69.0% P. coeruleiviridis 12.7% C. macellaria 0.4% C. rufifaces and 17.9% P. regina (Figure 3-25 B). Preserved larv ae (N=301) consisted of 77.7% P. coeruleiviridis 8.3% C. macellaria and 14.0% P. regina (Figure 3-25 C)

PAGE 48

36 Collection 15, April 26-May 1, 2003 Figures 3-26 A-C illustrate successional data. Phaenicia coeruleiviridis was the most dominant calliphorid species for the firs t 4 days of the sample period but decreased in abundance as decomposition progressed. Cochliomyia macellaria C. rufifaces and P. regina arrived after a delay of about 24 hours and increased in abundance as decomposition progressed. Adults aerially collected (N=185) consisted of 68.1% P. coeruleiviridis 29.2% C. macellaria 0.5% C. rufifaces and 2.2% P. regina (Figure 3-26 A). Reared adults (N=3 59) consisted of 50.7% P. coeruleiviridis 46.8% C. macellaria 2.5% P. regina (Figure 3-26 B). Preserved larvae (N=651) consisted of 50.5% P. coeruleiviridis 47.9% C. macellaria 1.4% P. regina and 0.2% were unknown (Figure 326 C). Collection 16, June 12-15, 2003 Figures 3-27 A-C illustrate successional data. Phaenicia coeruleiviridis was the most abundant species of adults aerially co llected on the first 2 days of the sampling period, but C. macellaria was the most abundant species of calliphorid overall during the sampling period (Figure 3-27 A). Adults aeri ally collected (N=221) consisted of 37.1% P. coeruleiviridis 56.1% C. macellaria 2.3% C. rufifaces and 4.5% P. regina (Figure 327 A). Reared adults (N=150) consisted of 6.0% P. coeruleiviridis 64.7% C. macellaria and 29.3% C. rufifaces (Figure 3-27 B). Preserved larvae (N=388) consisted of 73.7% P. coeruleiviridis 19.3% C. macellaria 5.4% C. rufifaces 1.3% P. regina and 0.3% were unknown (Figure 3-27 C). Collection 17, December 8-20, 2003 Phaenicia coeruleiviridis was the most dominant cal liphorid species during the sampling period (Figures 3-28 A-C). Adults aerially collected (N =186) consisted of

PAGE 49

37 89.8% P. coeruleiviridis 0.5% C. macellaria 2.2% C. rufifaces 6.5% P. regina and 1.1% C. livida (Figure 3-28 A). Reared adults (N=266) consisted of 95.9% P. coeruleiviridis 3.4% C. rufifaces and 0.8% P. regina (Figure 3-28 B). Preserved larvae (N=648) consisted of 90.4% P. coeruleiviridis 1.2% C. rufifaces and 8.3% P. regina (Figure 3-28 C). Collection 18, January 23-31, 2004 Figures 3-29 A-C illustrate the successional data. Phaenicia coeruleiviridis was the most abundant dominant species during the sampling period. Phormia regina C. megacephala and C. livida arrived after a delay of about 24 hours. Adults aerially collected (N=220) consisted of 92.3% P. coeruleiviridis 5.9% P. regina 0.5% C. megacephala 1.4% C. livida (Figure 3-29 A). Reared adults (N=294) consisted of 94.2% P. coeruleiviridis and 5.8% P. regina (Figure 3-29 B) Preserved larvae (N=347) consisted of 82.7% P. coeruleiviridis and 17.3% P. regina (Figure 3-29 C). Collection 19, March 5-14, 2004 Figures 3-30 A-C illustrate the successional data. Phaenicia coeruleiviridis was the most abundant dominant species during the fi rst few days of the sampling period, but decreased in abundance as decomposition progressed. Cochliomyia macellaria and P. regina arrived after a 24 hour delay and in creased in abundance as decomposition progressed. Adults aerially collect ed (N=165) consisted of 45.5% P. coeruleiviridis 6.7% C. macellaria and 47.9% P. regina. (Figure 3-30 A). Reared a dults (N=149) consisted of 33.6% P. coeruleiviridis 0.7% C. macellaria 65.8% P. regina (Figure 3-30 B). Preserved specimens (N=305) were comprised of 54.8% P. coeruleiviridis 3.0% C. macellaria and 42.3% P. regina (Figure 3-30 C).

PAGE 50

38 68.1 16.0 7.0 8.2 0.3 0.4 0.00.0 77.9 8.5 8.0 3.9 1.6 0.1 0.020.00 76.5 7.3 9.1 6.8 0.00.00.0 0.30.0 10.0 20.0 30.0 40.0 50.0 60.0 70.0 80.0Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia reginaChrysomya megacephala Calliphora lividaCalliphora vicina unknownRelative Abundance (%) Percentage aerial Percentage reared Percentage preserved 3rd-instar larvae Figure 3-1. Relative abundance of calliphorid adults aerially collected, adults reared from larvae and preserved third-instar larv ae collected from pig carrion during the entire study (N=15,396) between Nove mber 16, 2001 and March 14, 2004 in Earleton, Florida.

PAGE 51

39 (A) (B) Figure 3-2. A calliphorid first-in star larva hatching from its egg, (A), and the tiny Y-V shaped spiracles of a first-instar larva (B).

PAGE 52

40 Figure 3-3. The two sets of spiracl es of a first-second transitional larva. The first-instar Y-V slits (small, dark) can be seen belo w the two slit spiracles (larger, lightcolored) in this molting larva.

PAGE 53

41 Figure 3-4. The two inner slits in the spirac les of a second-instar calliphorid larva.

PAGE 54

42 Figure 3-5. The two sets of sp iracles of a second-third tran sitional calliphorid larva. The second-instar spiracles (sma ll, dark) have two slits, while the third-instar spiracles have three (larger, light-c olored) in this molting larva.

PAGE 55

43 Figure 3-6. The three inner splits in the spir acles of a third-instar calliphorid larva.

PAGE 56

44 Figure 3-7. Chrysomya rufifaces third-instar larvae. The obvious fleshy protuberances found on the secondand third-instar larv ae of this species are easy to see without a microscope.

PAGE 57

45 76.5 7.3 9.1 6.8 0.30.0 20.0 40.0 60.0 80.0Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifacesPhormia reginaunknownRelative Abundance (%) Figure 3-8. Relative abundance of third-instar ca lliphorid larvae preserved from pig carrion during the study (N=8253) in Earleton, Florida.

PAGE 58

46 0.0 20.0 40.0 60.0 80.0 100.0 Nov. 16-21, 2001 Dec. 29, 2001Jan.11, 2002 February 59, 2002 March 1519, 2002 April 29 May 1, 2002 May 20-23, 2002 July 22-25, 2002 Aug. 19-23, 2002 September 23-27, 2002 October 2628, 2002Relative Abundance of Adults Collected Aerially and Reared (N=3197 ) Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina Chrysomya megacephala Calliphora livida Calliphora vicina Figure 3-9. Calliphorid activity for year 1 of the study.

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47 0.0 20.0 40.0 60.0 80.0 100.0 November 30December 14, 2002 December 30, 2002-January 11, 2003 March 2-8, 2003 April 1-6, 2003 April 26-May 1, 2003 June 12-15, 2003 December 820, 2003 January 2331, 2004 March 5-14, 2004 Relative Abundance of Adults Collected Aerially and reared (N=4099 ) Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina Chrysomya megacephala Calliphora livida Figure 3-10. Calliphorid activity for year 2 of the study.

PAGE 60

48 (A) 23.6 20.2 19.9 30.4 32.8 25.9 33.2 30.5 29.3 28.4 5.2 0.9 8.1 16.5 19.4 17.4 22.4 23.9 23.7 19.10 5 10 15 20 25 30 35 40Nov. 16-21, 2001 Dec. 29, 2001-Jan.11, 2002 February 59, 2002 March 15-19, 2002 April 29 -May 1, 2002 May 20-23, 2002 July 22-25, 2002 Aug. 19-23, 2002 September 23-27, 2002 October 2628, 2002Temperature in Degrees C Mean Temperature Mean Low Temperature (B) 16.8 17.0 22.1 25.0 27.4 30.3 17.4 18.9 24.5 9.7 5.7 16.9 10.8 18.4 23.0 9.4 8.9 13.3 0.0 5.0 10.0 15.0 20.0 25.0 30.0 35.0 Nov. 30December 14, 2002 December 30, 2002-January 11, 2003 March 2-8, 2003 April 1-6, 2003 April 26-May 1, 2003 June 12-15, 2003 Dec. 8-20, 2003 January 2331, 2004 March 5-14, 2004Temperature in Degrees C Mean Temperature Mean Low Temperature Figure 3-11. Mean daily and mean low temp eratures (SD) for year 1, November 16, 2001 to October 26, 2002 (Part A) and year 2, November 30, 2002 to March 14, 2004 (Part B) of study.

PAGE 61

49 (A) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0November 16, 2001November 21, 2001 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces (B) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0November 21, 2001 Phaenicia coeruleiviridis Figure 3-12. Reared adults, N=232, (Part A), and preserved larvae, N=30, (Part B), from collection 1, November 16 to November 21, 2001. Relative Abundance (%) Relative Abundance (%)

PAGE 62

50 (A) Reared Adults 0.30.30.0 20.0 40.0 60.0 80.0 100.0January 11, 2002 Phaenicia coeruleiviridis Calliphora livida Calliphora vicina (B) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0December 29, 2001January 11, 2002 Phaenicia coeruleiviridis Phormia regina unknown Figure 3-13. Reared adults, N=362, (Part A), and preserved larvae, N=568, (Part B), from collection 2, December 29, 2001 to January 11, 2002. Relative Abundance (%) Relative Abundance (%)

PAGE 63

51 (A) Adults Collected Aerially 0.0 20.0 40.0 60.0 80.0 100.0February 1, 2002 (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0February 5, 2002February 9, 2002 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0February 9, 2002 Phaenicia coeruleiviridis Phormia regina Figure 3-14. Adults aerially collected, N=39, (Part A), rear ed adults, N=305, (Part B), and preserved larvae, N=151, (Part C), from collection 3, February 1-9, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 64

52 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0March 15, 2002March 17, 2002 Phaenicia coeruleiviridis Phormia regina (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0March 17, 2002March 18, 2002 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0March 17, 2003March 18, 2003March 19, 2003 unknown Figure 3-15. Adults collected aerially, N=71, (Part A), reared adults, N=326, (Par B), and preserved larvae, N=134, (Part C) from collection 4, March 15-19, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 65

53 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0April 29, 2002May 1, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Phormia regina (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0April 29, 2002May 1, 2002 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0May 1, 2002 Figure 3-16. Adults aerially collected, N=107, (Part A), rear ed adults, N=332, (Part B), and preserved larvae, N=196, (Part C), from collection 5, April 29 to May 1, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 66

54 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0May 20, 2002May 22, 2002May 23, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0May 23, 2002 (C) Preserved larvae 0.0 20.0 40.0 60.0 80.0 100.0May 23, 2002 Figure 3-17. Adults aerially co llected, N=51, (Part A), reared adults, N=67, (Part B), and preserved larvae, N=175 (Part C), from collection 6, May 20 to May 23, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 67

55 (A) Adults Aerially Collected 0.0 18.0 36.0 54.0 72.0 90.0July 22, 2002July 23, 2002July 24, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina Chrysomya megacephala (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0July 24, 2002July 25, 2002 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0July 24, 2002July 25, 2002 Figure 3-18. Adults aerially collected, N=99, (Part A), rear ed adults, N=223, (Part B), and preserved larvae, N=700, (Part C), fr om collection 7, July 22 to July 24, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 68

56 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0August 19, 2002August 20, 2002August 21, 2002August 22, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Chrysomya megacephala (B) 0.0 18.0 36.0 54.0 72.0 90.0August 21, 2002August 22, 2002August 23, 2002 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0August 21, 2002August 22, 2002 unknown Figure 3-19. Adults aerially collected, N=76, (Part A), rear ed adults, N=248, (Part B), and preserved larvae, N=499, (Part C), from collection 8, August 19-23, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 69

57 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0September 23, 2002September 25, 2002September 26, 2002September 27, 2002 (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0September 25, 2002September 26, 2002September 27, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Chrysomya megacephala (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0September 25, 2002September 26, 2002September 27, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina Figure 3-20. Adults aerially collected, N=76, (Part A), rear ed adults, N=323, (Part B), and preserved larvae, N=499, (Part C), from collection 9, September 23-27, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 70

58 (A) Adults Aerially Collected 0.0 14.0 28.0 42.0 56.0 70.0October 26, 2002October 27, 2002October 28, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces (B) Reared Adults 0.0 14.0 28.0 42.0 56.0 70.0October 27, 2002October 28, 2002 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0October 26, 2002October 27, 2002October 28, 2002 Figure 3-21. Adults aerially collected, N=87, (Part A), rear ed adults, N=131, (Part B), and preserved larvae, N=294, (Part C), from collection 10, October 26-28. 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 71

59 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0November 30, 2002December 8, 2002 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0December 7, 2002December 8, 2002December 10, 2002 December 12, 2002 December 14, 2002 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0December 7, 2002 December 8, 2002 December 10, 2002 December 12, 2002 December 13, 2002 December 14, 2002 Figure 3-22. Adults aerially co llected, N=60, (Part A), reared adults, N= 384, (Part B), and preserved larvae, N=1248, (Part C) from collection 11, November 30December 8, 2002. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 72

60 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0December 30, 2002January 2, 2003January 5, 2003January 6, 2003 Phaenicia coeruleiviridis Calliphora livida (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0January 3, 2003 January 4, 2003 January 5, 2003 January 6, 2003 January 8, 2003 January 9, 2003 January 11, 2003 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0January 3, 2003 January 4, 2003 January 5, 2003 January 6, 2003 January 8, 2003 January 9, 2003 January 11, 2003 Phaenicia coeruleiviridis Cochliomyia macellaria unknown Figure 3-23. Adults aerially co llected, N=56, (Part A), reared adults, N= 371, (Part B), and preserved larvae, N=581, (Part C), from collection 12, December 30, 2002 to January 11, 2003. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 73

61 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0March 1, 2002March 2, 2003March 5, 2003March 6, 2003March 7, 2003 Phaenicia coeruleiviridis Phormia regina Calliphora livida (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0March 5, 2003March 6, 2003March 7, 2003March 8, 2003 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0March 5, 2003March 6, 2003March 7, 2003March 8, 2003 unknown Figure 3-24. Adults aerially collected, N=66, (Part A), rear ed adults, N=166, (Part B), and preserved larvae, N=340, (Part C), from collection 13, March 1-8, 2003. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 74

62 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0April 1, 2003April 2, 2003April 4, 2003April 5, 2003April 6, 2003 (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0April 4, 2003April 5, 2003April 6, 2003 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0April 4, 2003April 5, 2003April 6, 2003 Figure 3-25. Adults aerially collected, N=247, (Part A), rear ed adults, N=229, (Part B), and preserved larvae, N=301, (Part C) from collection 14, April 1-6, 2003. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 75

63 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0April 26, 2003April 27, 2003April 28, 2003April 29, 2003April 30, 2003 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0April 27, 2003April 28, 2003April 29, 2003April 30, 2003 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0April 28, 2003April 29, 2003April 30, 2003May 1, 2003 unknown Figure 3-26. Adults aerially collected, N=185, (Part A), rear ed adults, N=359, (Part B), and preserved larvae, N=651, (Part C), from collection 15, April 26-May 1, 2003. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 76

64 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0June 12, 2003June 13, 2003June 14, 2003June 15, 2003 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0June 13, 2003June 14, 2003June 15, 2003 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0June 14, 2003June 15, 2003 unknown Figure 3-27. Adults aerially collected, N=221, (Part A), rear ed adults, N=150, (Part B), and preserved larvae, N=388, (Part C), specimens from collection 16, June 1215, 2003. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 77

65 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0 December 8, 2003 December 10, 2003 December 11, 2003 December 12, 2003 December 13, 2003 December 14, 2003 December 16, 2003 Phaenicia coeruleiviridis Cochliomyia macellaria Chrysomya rufifaces Phormia regina Calliphora livida (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0December 12, 2003 December 13, 2003 December 14, 2003 December 16, 2003 December 17, 2003 December 20, 2003 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0December 13, 2003 December 14, 2003 December 16, 2003 December 17, 2003 December 20, 2003 Figure 3-28. Adults aerially collected, N=186, (Part A), rear ed adults, N=266, (Part B), and preserved larvae, N=648, (Part C), from collection 17, December 8-20, 2003. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 78

66 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0January 23, 2004 January 24, 2004 January 25, 2004 January 26, 2004 January 27, 2004 January 28, 2004 January 29, 2004 January 30, 2004 January 31, 2004 Phaenicia coeruleiviridis Phormia regina Chrysomya megacephala Calliphora livida (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0January 27, 2004January 28, 2004January 29, 2004 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0January 28, 2004January 29, 2004January 30, 2004January 31, 2004 Figure 3-29. Adults aerially collected, N=220, (Part A), rear ed adults, N=294, (Part B), and preserved larvae, N=347, (Part C), from collection 18, January 23-31, 2004. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

PAGE 79

67 (A) Adults Aerially Collected 0.0 20.0 40.0 60.0 80.0 100.0March 5, 2004 March 6, 2004 March 7, 2004 March 8, 2004 March 9, 2004 March 13, 2004 March 14, 2004 Phaenicia coeruleiviridis Cochliomyia macellaria Phormia regina (B) Reared Adults 0.0 20.0 40.0 60.0 80.0 100.0March 8, 2004March 9, 2004March 10, 2004March 14, 2004 (C) Preserved Larvae 0.0 20.0 40.0 60.0 80.0 100.0March 8, 2004March 9, 2004March 10, 2004March 13, 2004March 14, 2004 Figure 3-30. Adults aerially collected, N=316, (Part A), rear ed adults, N=639, (Part B), and preserved larvae, N=675, (Part C), from collection 19, March 5-14, 2004. Relative Abundance (%) Relative Abundance (%) Relative Abundance (%)

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68 CHAPTER 4 DISCUSSION The most common calliphorid species coll ected on pig carrion during the study aerially or by rearing was Phaenicia coeruleiviridis Of the total adult specimens (N=6971) identified duri ng the study, 75.0% were P. coeruleiviridis Similarly, of the identifiable third-instar larvae (N=8253), 76.5% were P. coeruleiviridis This species was most abundant from late October to th e end of May (Figures 3-9 and 3-10). Chrysomya rufifaces and C. macellaria were also collected during thos e months, but in much smaller numbers. Phaenicia coeruleiviridis was less abundant during the summer months-when the temperatures were over 25.0 C. Cochliomyia macellaria was not present during the winter and was the most abunda nt species collected in J une 2003 (Figures 3-9 and 3-10) Phormia regina was present from mid-November 2001 until the end of July 2002 (Figures 3-9 and 3-10). No specimens were co llected in August or September in 2002. It was the most abundant species collected in March 2004, but that was unusual compared to the rest of the study because P. regina was quite low in abundance compared to the other calliphorids. Hall (1948) found that P. regina is abundant in the spring months in the southern states (location not defined by Hall) but apparently this was not the case during my study in rural nor th-central Florida. Of the seven species of calliphorids collect ed during the study, all were consistent with published distribution records, seasona lity, and successional patterns (Campobasso et al. 2001). However, Byrd (1998) collected Phaenica cuprina (Wiedemann) (= Phaenicia pallescens Shannon) from pig carcasses in June and September 1996, and also

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69 in September 1997, but this species was not found during this study. Byrd’s study was conducted at an undesignated lo cation in the Gainesville area a “woodland habitat” (page 28 of dissertation), which consisted of a Live Oak hammock with no understory or roosting sites (for female blow flies) within close proximity to the pigs. Phaenicia cuprina is an urban fly that prefers excrem ent to carrion (Byrd 2001), which could explain why it was not found on pig carrion in rural Earleton. During this study, the species and relativ e abundances of calliphorids found during adult aerial, reared and larval collections made during December 2002 and 2003 (Figures 3-23 A and B and 3-28 A-C) were similar, as were the collections made during November 2001 and November 2002 (Figures 3-12 A and B, 3-22 A-C). Finally, data obtained for three years (2002, 2003, and 2004) in March (Figures 3-15 A-C, 3-24 A-C, and 3-30 A-C) also yielded similar species and relative abundances of species. The congruence between these samples suggests that there is a consistent pattern in species composition and relative abundance through time in this sample site. Common calliphorid species such as Phaenicia sericata and Phaenicia eximia apparently are not normally found in rural no rth-central Florida, but were found in the urban Gainesville area by Byrd (1998) and Peters (2003). Different calliphorids are associated with different hab itats. An urban area, with odor s from human refuse, cooking, garbage dumps, and improper sanitation will have different species assemblages than rural, wooded areas or arid regions. Some ca lliphorid species were not expected to be found because they are not associated with car rion. For example, Hall (1948) stated that the calliphorid Pollenia rudis (Fabricus) has been collected in northern Florida, but this species is exclusively a pa rasite of earthworms.

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70 Calliphorid species (both adults and larv ae) found on human remains (in cases from 1991) in Palm Beach and Lake Counties, Florida were identified as C. rufifaces and C. macellaria (Haskell 2003, personal communication). The evidentiary specimens were identified by Neal Haskell during the court cases. These two species were collected on pig carrion in rural north-central Florida during my study, on pig carrion in Gainesville by Byrd (1998) and on bear carrion in Gainesville by Peters (2003). In northwest Indiana, Haskell (1989) found P. regina to be the most abundant species of Calliphoridae from late spring to early autumn. Phaenicia coeruleiviridis was dominant during the spring and au tumn. Haskell also found a few C. vicina and C. livida specimens during the spring and autumn. In West Virginia, Joy et al. (2002) found P. regina to be the dominant calliphorid in May 2002 on raccoon carrion. They found only a few Phaenicia species on these raccoons, which were killed by cars so th at the time of death was unknown. The dead raccoons were frozen and transported to the site in garbage bags. In Menard County Texas, Cushing and Parish (1938) captured Cochliomyia and Phaenicia species from April to November 1933, but P. regina was far more abundant. They caught hundreds of thousands of flies with pit-fall traps and traps baited with beef. No P. coeruleiviridis were found. Calliphora vicina C. livida P. coeruleiviridis C. macellaria C. rufifaces P. regina and C. megacephala were collected at both rural and urban sites during a recent pig carrion study in central Te xas (Tenorio et al. 2003). Phaenicia coeruleiviridis was found from March until May while C. livida and C. vicina were found from December to May. Cochliomyia macellaria were found all year and C. rufifaces was found from

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71 March until November. Chrysomya megacephala was found from September to November, and P. regina was not found from June through August. In northern Mississippi, P. coeruleiviridis P. regina and C. macellaria were the dominant species from April to September (Goddard and Lago 1985). Phormia regina was dominant from October to March on carcasses of rabbits, opossum, and fish carcasses. Deonier (1937, 1938) collected hundreds of thousands of calliphorids with beefbaited traps in Arizona. Phormia regina, C. macellaria and Phaenicia species were found. Cochliomyia macellaria were captured in enormous numbers all year. A single P. regina specimen was collected in August and again in September of 1937, while almost 293,000 C. macellaria and 1000 Phaenicia species were collected at the same time (Deonier 1942). Denno and Cothran (1975) found P. sericata and P. regina to be the dominant calliphorids on rabbit carrion in Davis, CA dur ing June to September. Hall and Doisy (1993) found C. macellaria P. regina and P coeruleiviridis during their field studies in Missouri from mid-June to late August 1992. Reed (1958) found C. livida C. vicina C. macellaria P. coeruleiviridis and P. regina during his dog carcass study in Tennessee. Phormia regina was the most abundant calliphorid species on the dog carcasses (Reed 1958). Watson and Carlton (2003) c onducted research in Louisiana from April 1 to July 1999 using bear, deer, alligator, and pig carca sses and found similar species to those of my study. The most common species on all four carcasses (in order of occurrence) were P. regina P. coeruleiviridis and P. sericata Of note was that P. coeruleiviridis landed

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72 on carcasses within minutes of deposition. Cochliomyia macellaria and C. rufifaces were also found on the pig carcasses, but in smaller numbers. Peters (2003, page 78) conducted a ca rrion study using three dead bears ( Ursus americanus floridanus ) in the Gainesville area and c oncluded that the most abundant blow fly found in north central Fl orida is the Hair y Maggot Blow Fly, Chrysomya rufifaces . Only 8 P. coeruleiviridis adults were among 94 calliphorid specimens collected by Peters (2003). Th is is contrary to my resu lts, which indicate that P. coeruleiviridis is most abundant on pig carrion in this area while C. rufifaces consisted of less than 10 % of the total sp ecimens collected during the entire study. The differences in our data are even greater wh en comparing abundances of C. megacephala which consisted of less than 2.0 % of the total f lies during my study wh ile Peters found this species to comprise 44.7 % of adult calliphorids collected in her study. The differences in our findings may be attributed to the following: The actual time of death of the bears was approximated; time O was not the actual time of death. Once retrieved by the w ildlife officer, the dead bears were not protected in plastic or placed in containers while being transferred to the study site. At least 48 hours passed before any samp le collections were taken on Bear 1. Therefore, each bear had a collection delay of at least 48 hours. These differences in the method of procuring and placeme nt of carcasses may have resulted in completely missing the first wave of carri on insects attracted to fresh carcasses, specifically P. coeruleiviridis. Apparently, P. coeruleiviridis larvae were collected on all three bears, which clearly indicate that the adults had been present. Bears may decompose differently than do pi gs or humans; bears have thick fur, thick skin, and layers of ad ipose tissue that are unique to bears (Watson and Carlton 2003). These differences may make b ear carcasses more attractive to C. rufifaces and C. megacephala or less attractive to P. coeruleiviridis Watson and Carlton (2003) noticed that P. coeruleiviridis arrived earlier on the carcasses of deer, alligator and pigs than on the carcass of the bear. The number of larval specimens collected or reared (if any) is not disclosed by Peters (2003). Apparently, only 94 adu lt calliphorid specim ens were aerially collected, of which almost half were C. megacephala and 19 were C. rufifaces

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73 Additionally, aerial collections were made late in the day (between 5-6 PM) when some calliphorid flies are less active. The three bears were each placed at th e site by Peters (2003) during different months of the year. As a result, there could be variability due to a lack of replications or di fferent seasons. In my study, P. coeruleiviridis was the most abundant species of calliphorid found on pig carrion in rural north-central Fl orida between 2001and 2004 during most of the year. This species was always the first to ar rive at fresh pig carri on, the first to deposit eggs, the first to complete development and the first to migrate off the carcass to pupariate in the soil. In the spring and summer, these events took 5 days, or fewer, to complete. Depending on the season, and almost always after an approximate 24-h delay, C. macellaria C. rufifaces P. regina and C. megacephala arrived at the pig carcasses Calliphora livida and C. vicina arrived at the carcasses also after a delay, but only a few specimens of each were collect ed (Figures 3-3 and 3-4). Despite the fact that P. coeruleiviridis is commonly found in many other states, there are no developmental rate data for this species because, until now, no one has been able to successfully rear the larvae to adulthood (Has kell 1989, Hall and Doisy 1993). During this study, I discovered an organic pupa tion substrate that results in successful rearing of P. coeruleiviridis larvae to adulthood. Now that we can rear P. coeruleivirdis we need detailed developmental rate data for this species so a fore nsic entomologist can determine accumulated degree hours (ADH) or days (ADD) to determine the postmortem interval. The fact that we do not have developmental data for P. coeruleiviridis is very unfortunate because it is an important indicat or species for determining PMI, especially in rural north-central Florida. Standa rdized rearing data are needed for P. coeruleiviridis

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74 as are rearing data for other calliphorids that are attracted to human remains including Calliphora vicina, C. vomitoria, Eucalliphora sp ., Cynomyopsis cadaverina, Phaenicia sericata, Lucilia Illustris, Phor mia regina, Cochliomyia macellaria and Paralucilia wheeleri.

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75 APPENDIX A RAW DATA: FLIES COLLECTED AS ADU LTS OR REARED FROM LARVAE PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 2A 11/16/2001 R 7A 21 8 0 13 0 0 0 0 2B 11/16/2001 R 8A 29 28 1 0 0 0 0 0 2B 11/21/2001 R 10A 157 157 0 0 0 0 0 0 2C 11/16/2001 R 9A 25 25 0 0 0 0 0 0 3A 1/11/2002 R 13A 145 145 0 0 0 0 0 0 3B 1/11/2002 R 17A 115 115 0 0 0 0 0 0 3C 1/11/2002 R 15A 102 100 0 0 0 0 1 1 4A 2/1/2002 A 23 4 4 0 0 0 0 0 0 4A 2/5/2002 R 24 54 54 0 0 0 0 0 0 4B 2/1/2002 A 22 12 12 0 0 0 0 0 0 4B 2/5/2002 R 25 71 71 0 0 0 0 0 0 4C 2/1/2002 A 21 12 12 0 0 0 0 0 0 4C 2/5/2002 R 26 38 38 0 0 0 0 0 0 4E 2/1/2002 A 20 11 11 0 0 0 0 0 0 4E 2/5/2002 R 27 55 55 0 0 0 0 0 0 4E 2/9/2002 R 28 87 87 0 0 0 0 0 0 5A 3/15/2002 A 30 17 17 0 0 0 0 0 0 5A 3/17/2002 R 35 59 59 0 0 0 0 0 0 5A 3/17/2002 A 36 22 22 0 0 0 0 0 0 5A 3/18/2002 R 47 65 65 0 0 0 0 0 0 5B 3/15/2002 A 31 15 15 0 0 0 0 0 0 5B 3/17/2002 R 40 56 56 0 0 0 0 0 0 5C 3/15/2002 A 32 7 7 0 0 0 0 0 0 5C 3/17/2002 R 38 42 42 0 0 0 0 0 0 5E 3/15/2002 A 33 10 8 0 0 2 0 0 0 5E 3/17/2002 R 42 47 47 0 0 0 0 0 0 5E 3/18/2002 R 44A 57 57 0 0 0 0 0 0 6A 4/29/2002 A 49 11 11 0 0 0 0 0 0 6A 5/1/2002 R 53 58 58 0 0 0 0 0 0 6A 5/1/2002 A 55 13 0 0 0 13 0 0 0 6B 4/29/2002 A 50 17 17 0 0 0 0 0 0 6B 5/1/2002 R 57 33 33 0 0 0 0 0 0 6B 5/1/2002 R 59 42 42 0 0 0 0 0 0 6B 5/1/2002 A 61 6 4 1 0 1 0 0 0 6C 4/29/2002 A 51 44 44 0 0 0 0 0 0 6C 5/1/2002 R 63 56 56 0 0 0 0 0 0 6C 5/1/2002 R 65 38 38 0 0 0 0 0 0 6C 5/1/2002 A 66 0 0 0 0 0 0 0 0 6E 4/29/2002 A 52 16 16 0 0 0 0 0 0 6E 4/29/2002 R 70 69 36 9 0 24 0 0 0

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76 PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 6E 5/1/2002 R 68 36 36 0 0 0 0 0 0 7A 5/20/2002 A 72 11 0 11 0 0 0 0 0 7A 5/22/2002 A 76 18 3 6 3 6 0 0 0 7A 5/23/2002 A 77 8 1 1 2 4 0 0 0 7A 5/23/2002 R 78 67 67 0 0 0 0 0 0 7B 5/20/2002 A 73 12 4 2 6 0 0 0 0 7C 5/20/2002 A 74 2 1 0 0 1 0 0 0 7E 5/20/2002 A 75 0 0 0 0 0 0 0 0 8A 7/22/2002 A 80 0 0 0 0 0 0 0 0 8A 7/23/2002 A 84 11 4 3 3 1 0 0 0 8A 7/24/2002 A 91 11 0 0 9 0 2 0 0 8A 7/24/2002 R 92 21 21 0 0 0 0 0 0 8A 7/24/2002 R 94 25 24 0 1 0 0 0 0 8A 7/25/2002 R 107 57 0 0 57 0 0 0 0 8B 7/22/2002 A 81 9 8 1 0 0 0 0 0 8B 7/23/2002 A 87 13 10 0 2 1 0 0 0 8C 7/22/2002 A 82 11 11 0 0 0 0 0 0 8C 7/23/2002 A 86 7 5 0 2 0 0 0 0 8C 7/24/2002 A 102 5 0 0 5 0 0 0 0 8C 7/24/2002 R 103 42 42 0 0 0 0 0 0 8E 7/22/2002 A 85 8 3 0 5 0 0 0 0 8E 7/24/2002 A 83 17 17 0 0 0 0 0 0 8E 7/24/2002 A 97 7 0 1 5 0 1 0 0 8E 7/24/2002 R 98 24 18 0 6 0 0 0 0 8E 7/25/2002 R 109 54 0 0 54 0 0 0 0 9A 8/19/2002 A 111 14 14 0 0 0 0 0 0 9A 8/20/2002 A 115 18 18 0 0 0 0 0 0 9A 8/21/2002 R 119 5 2 0 1 0 2 0 0 9A 8/21/2002 A 123 10 3 1 6 0 0 0 0 9A 8/22/2002 A 153 2 0 2 0 0 0 0 0 9A 8/22/2002 R 157 11 8 0 1 0 2 0 0 9A 8/23/2002 R 154 27 4 1 20 0 2 0 0 9B 8/19/2002 A 112 2 2 0 0 0 0 0 0 9B 8/20/2002 A 116 10 10 0 0 0 0 0 0 9B 8/21/2002 R 124 43 16 0 7 0 20 0 0 9B 8/21/2002 A 125 10 3 1 6 0 0 0 0 9B 8/22/2002 A 148 4 1 0 2 0 1 0 0 9B 8/22/2002 R 149 14 12 0 1 0 1 0 0 9B 8/23/2002 R 151 38 0 0 38 0 0 0 0 9C 8/19/2002 A 113 1 1 0 0 0 0 0 0 9C 8/20/2002 A 117 12 11 0 1 0 0 0 0 9C 8/20/2002 A 134 0 0 0 0 0 0 0 0 9C 8/22/2002 R 135 13 4 0 7 0 2 0 0 9C 8/22/2002 R 137 16 14 0 0 0 2 0 0 9E 8/20/2002 A 118 11 10 0 1 0 0 0 0 9E 8/21/2002 A 129 8 0 0 7 0 1 0 0 9E 8/21/2002 R 130 27 27 0 0 0 0 0 0 9E 8/22/2002 A 114 11 7 2 2 0 0 0 0 9E 8/22/2002 A 142 5 0 2 3 0 0 0 0

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77 PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 9E 8/22/2002 R 145 13 10 0 2 0 1 0 0 9E 8/23/2002 R 143 41 10 2 28 0 1 0 0 10A 9/23/2002 A 159 0 0 0 0 0 0 0 0 10A 9/25/2002 A 163 10 3 4 3 0 0 0 0 10A 9/25/2002 R 165 32 32 0 0 0 0 0 0 10A 9/26/2002 A 177 13 0 8 5 0 0 0 0 10A 9/26/2002 R 178 22 22 0 0 0 0 0 0 10A 9/26/2002 R 180 22 15 0 0 0 7 0 0 10B 9/23/2002 A 160 6 6 0 0 0 0 0 0 10B 9/25/2002 A 166 10 3 5 2 0 0 0 0 10B 9/25/2002 R 167 32 27 5 0 0 0 0 0 10B 9/26/2002 R 183 16 16 0 0 0 0 0 0 10B 9/26/2002 R 185 55 0 28 0 0 27 0 0 10C 9/23/2002 A 162 10 10 0 0 0 0 0 0 10C 9/25/2002 A 174 9 5 1 3 0 0 0 0 10C 9/25/2002 R 175 26 26 0 0 0 0 0 0 10C 9/27/2002 A 187 6 0 4 2 0 0 0 0 10C 9/27/2002 R 188 42 12 17 4 0 9 0 0 10E 9/23/2002 A 161 7 7 0 0 0 0 0 0 10E 9/25/2002 A 170 5 0 4 1 0 0 0 0 10E 9/25/2002 R 171 25 25 0 0 0 0 0 0 10E 9/27/2002 R 191 51 0 49 0 0 2 0 0 11A 10/26/2002 A 193 8 2 1 5 0 0 0 0 11A 10/27/2002 A 200 3 0 2 1 0 0 0 0 11A 10/27/2002 R 201 43 20 9 14 0 0 0 0 11A 10/27/2002 R 203 31 15 0 16 0 0 0 0 11A 10/28/2002 A 209 8 0 1 7 0 0 0 0 11A 10/28/2002 R 210 14 14 0 0 0 0 0 0 11A 10/28/2002 R 212 28 20 1 7 0 0 0 0 11B 10/26/2002 A 195 17 3 5 9 0 0 0 0 11B 10/26/2002 R 196 0 0 0 0 0 0 0 0 11C 10/26/2002 A 199 19 2 8 9 0 0 0 0 11C 10/28/2002 A 205 6 1 1 4 0 0 0 0 11C 10/28/2002 R 206 15 0 0 15 0 0 0 0 11E 10/26/2002 A 198 17 3 7 7 0 0 0 0 11E 10/28/2002 A 208 9 0 6 3 0 0 0 0 12A 11/30/2002 A 214 10 10 0 0 0 0 0 0 12A 12/7/2002 R 221 33 33 0 0 0 0 0 0 12A 12/8/2002 R 228 34 34 0 0 0 0 0 0 12A 12/8/2002 A 230 19 14 1 3 1 0 0 0 12A 12/12/2002 R 253 29 29 0 0 0 0 0 0 12A 12/13/2002 A 255 0 0 0 0 0 0 0 0 12A 12/14/2002 R 258 21 21 0 0 0 0 0 0 12B 11/30/2002 A 215 6 6 0 0 0 0 0 0 12B 11/30/2002 R 218 0 0 0 0 0 0 0 0 12B 12/7/2002 R 222 33 33 0 0 0 0 0 0 12B 12/8/2002 R 231 25 25 0 0 0 0 0 0 12B 12/8/2002 A 233 0 0 0 0 0 0 0 0 12B 12/12/2002 R 249 9 9 0 0 0 0 0 0

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78 PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 12B 12/12/2002 R 251 2 2 0 0 0 0 0 0 12C 11/30/2002 A 216 10 10 0 0 0 0 0 0 12C 12/7/2002 R 226 24 24 0 0 0 0 0 0 12C 12/10/2002 R 235 27 27 0 0 0 0 0 0 12C 12/10/2002 R 239 46 0 0 46 0 0 0 0 12C 12/12/2002 R 245 9 9 0 0 0 0 0 0 12E 11/30/2002 A 217 7 7 0 0 0 0 0 0 12E 12/7/2002 R 224 38 38 0 0 0 0 0 0 12E 12/8/2002 A 234 8 8 0 0 0 0 0 0 12E 12/10/2002 R 240 36 36 0 0 0 0 0 0 12E 12/12/2002 R 247 18 18 0 0 0 0 0 0 13B 12/30/2002 A 262 2 2 0 0 0 0 0 0 13B 1/2/2003 A 263 17 17 0 0 0 0 0 0 13B 1/3/2003 R 266 27 27 0 0 0 0 0 0 13B 1/4/2003 R 276 30 27 0 0 0 0 3 0 13B 1/5/2003 R 283 21 21 0 0 0 0 0 0 13B 1/9/2003 R 292 20 20 0 0 0 0 0 0 13B 1/9/2003 A 294 0 0 0 0 0 0 0 0 13C 12/30/2002 A 261 7 7 0 0 0 0 0 0 13C 1/2/2003 A 264 8 7 0 0 0 0 1 0 13C 1/3/2003 R 270 27 27 0 0 0 0 0 0 13C 1/4/2003 R 274 21 21 0 0 0 0 0 0 13C 1/5/2003 A 281 1 1 0 0 0 0 0 0 13C 1/5/2003 R 282 34 34 0 0 0 0 0 0 13C 1/8/2003 R 290 9 9 0 0 0 0 0 0 13C 1/9/2003 R 298 27 27 0 0 0 0 0 0 13C 1/11/2003 R 300 28 28 0 0 0 0 0 0 13E 12/30/2002 A 260 13 13 0 0 0 0 0 0 13E 1/2/2003 A 265 4 4 0 0 0 0 0 0 13E 1/3/2003 R 268 17 17 0 0 0 0 0 0 13E 1/4/2003 R 272 25 25 0 0 0 0 0 0 13E 1/5/2003 A 278 1 1 0 0 0 0 0 0 13E 1/5/2003 R 279 28 28 0 0 0 0 0 0 13E 1/6/2003 A 285 3 3 0 0 0 0 0 0 13E 1/6/2003 R 286 39 39 0 0 0 0 0 0 13E 1/8/2003 R 288 9 9 0 0 0 0 0 0 13E 1/9/2003 R 295 9 9 0 0 0 0 0 0 13E 1/9/2003 A 297 0 0 0 0 0 0 0 0 14A 3/2/2003 A 318 16 16 0 0 0 0 0 0 14A 3/6/2003 R 304 39 38 0 0 1 0 0 0 14A 3/6/2003 R 306 42 40 0 0 2 0 0 0 14A 3/6/2003 A 308 16 11 0 0 5 0 0 0 14A 3/8/2003 R 314 15 15 0 0 0 0 0 0 14A 3/8/2003 R 316 26 26 0 0 0 0 0 0 14B 3/5/2003 A 301 20 19 0 0 0 0 1 0 14B 3/5/2003 R 302 21 21 0 0 0 0 0 0 14E 3/1/2003 A 300 11 10 0 0 0 0 1 0 14E 3/7/2003 R 309 23 23 0 0 0 0 0 0 14E 3/7/2003 A 311 3 2 0 0 1 0 0 0

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79 PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 14E 3/7/2003 R 312 0 0 0 0 0 0 0 0 15A 4/1/2003 A 320 13 13 0 0 0 0 0 0 15A 4/2/2003 A 324 21 20 0 0 1 0 0 0 15A 4/4/2003 A 330 18 6 12 0 0 0 0 0 15A 4/5/2003 R 340 23 21 0 0 2 0 0 0 15A 4/5/2003 A 342 12 4 8 0 0 0 0 0 15A 4/6/2003 A 352 16 7 6 0 3 0 0 0 15A 4/6/2003 R 353 2 0 0 0 2 0 0 0 15B 4/1/2003 A 321 9 9 0 0 0 0 0 0 15B 4/2/2003 A 325 26 25 0 0 1 0 0 0 15B 4/4/2003 R 331 28 20 1 0 7 0 0 0 15B 4/4/2003 A 333 13 11 2 0 0 0 0 0 15B 4/5/2003 R 343 24 19 2 0 3 0 0 0 15B 4/5/2003 A 345 6 3 1 0 2 0 0 0 15C 4/1/2003 A 319 10 10 0 0 0 0 0 0 15C 4/2/2003 A 323 21 21 0 0 0 0 0 0 15C 4/4/2003 A 327 20 15 3 0 2 0 0 0 15C 4/4/2003 R 329 52 50 0 1 1 0 0 0 15C 4/5/2003 R 337 0 0 0 0 0 0 0 0 15C 4/5/2003 A 339 21 7 10 0 4 0 0 0 15C 4/6/2003 A 349 0 0 0 0 0 0 0 0 15C 4/6/2003 R 350 13 3 4 0 6 0 0 0 15E 4/1/2003 A 322 11 9 2 0 0 0 0 0 15E 4/2/2003 A 326 10 10 0 0 0 0 0 0 15E 4/4/2003 R 334 23 23 0 0 0 0 0 0 15E 4/4/2003 A 336 12 10 1 0 1 0 0 0 15E 4/5/2003 R 346 28 21 5 0 2 0 0 0 15E 4/5/2003 A 348 6 5 1 0 0 0 0 0 15E 4/6/2003 R 355 36 1 17 0 18 0 0 0 15E 4/6/2003 A 357 2 1 1 0 0 0 0 0 16B 4/26/2003 A 358 31 31 0 0 0 0 0 0 16B 4/27/2003 A 365 11 11 0 0 0 0 0 0 16B 4/28/2003 A 372 13 9 3 0 1 0 0 0 16B 4/28/2003 R 373 33 28 5 0 0 0 0 0 16B 4/29/2003 R 381 34 34 0 0 0 0 0 0 16B 4/29/2003 A 389 6 5 1 0 0 0 0 0 16B 4/30/2003 A 388 19 19 0 0 0 0 0 0 16C 4/26/2003 A 360 14 13 1 0 0 0 0 0 16C 4/27/2003 A 361 21 9 12 0 0 0 0 0 16C 4/28/2003 A 366 14 1 11 1 1 0 0 0 16C 4/28/2003 R 367 34 32 2 0 0 0 0 0 16C 4/29/2003 A 375 11 2 8 0 1 0 0 0 16C 4/29/2003 R 376 63 0 59 0 4 0 0 0 16C 4/30/2003 A 383 5 0 5 0 0 0 0 0 16C 4/30/2003 R 384 85 0 84 0 1 0 0 0 16C 5/1/2003 A 393 0 0 0 0 0 0 0 0 16E 4/26/2003 A 359 16 16 0 0 0 0 0 0 16E 4/27/2003 A 362 13 6 7 0 0 0 0 0 16E 4/27/2003 R 363 19 19 0 0 0 0 0 0

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80 PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 16E 4/28/2003 A 369 11 4 6 0 1 0 0 0 16E 4/28/2003 R 370 33 33 0 0 0 0 0 0 16E 4/29/2003 A 378 0 0 0 0 0 0 0 0 16E 4/29/2003 R 379 32 32 0 0 0 0 0 0 16E 4/30/2003 R 386 26 4 18 0 4 0 0 0 17B 6/12/2003 A 396 16 8 8 0 0 0 0 0 17B 6/13/2003 A 399 23 22 1 0 0 0 0 0 17B 6/14/2003 A 400 30 0 20 0 10 0 0 0 17B 6/14/2003 R 401 10 1 9 0 0 0 0 0 17B 6/15/2003 R 419 20 0 14 6 0 0 0 0 17B 6/15/2003 A 421 13 0 13 0 0 0 0 0 17C 6/12/2003 A 394 46 27 19 0 0 0 0 0 17C 6/13/2003 A 398 0 0 0 0 0 0 0 0 17C 6/14/2003 R 403 0 0 0 0 0 0 0 0 17C 6/14/2003 A 405 18 6 10 2 0 0 0 0 17C 6/15/2003 R 414 5 0 1 4 0 0 0 0 17C 6/15/2003 A 416 12 0 10 2 0 0 0 0 17C 6/15/2003 R 417 32 0 0 32 0 0 0 0 17E 6/12/2003 A 395 31 19 11 1 0 0 0 0 17E 6/13/2003 A 397 0 0 0 0 0 0 0 0 17E 6/13/2003 R 401 21 7 14 0 0 0 0 0 17E 6/14/2003 R 406 28 0 28 0 0 0 0 0 17E 6/14/2003 A 408 13 0 13 0 0 0 0 0 17E 6/15/2003 R 409 24 0 24 0 0 0 0 0 17E 6/15/2003 A 411 19 0 19 0 0 0 0 0 17E 6/15/2003 R 412 10 1 7 2 0 0 0 0 18A 12/8/2003 A 424 23 23 0 0 0 0 0 0 18A 12/10/2003 A 428 10 10 0 0 0 0 0 0 18A 12/13/2003 R 439 9 9 0 0 0 0 0 0 18A 12/13/2003 A 441 14 14 0 0 0 0 0 0 18A 12/14/2003 R 448 22 22 0 0 0 0 0 0 18A 12/14/2003 A 450 4 4 0 0 0 0 0 0 18A 12/16/2003 R 457 24 24 0 0 0 0 0 0 18A 12/16/2003 A 459 9 7 0 0 2 0 0 0 18A 12/20/2003 R 464 23 21 0 0 2 0 0 0 18B 12/8/2003 A 422 10 10 0 0 0 0 0 0 18B 12/10/2003 A 427 13 13 0 0 0 0 0 0 18B 12/11/2003 A 429 2 0 0 0 0 0 2 0 18B 12/12/2003 A 430 6 5 0 0 1 0 0 0 18B 12/12/2003 R 432 23 23 0 0 0 0 0 0 18B 12/13/2003 A 436 20 17 0 0 3 0 0 0 18B 12/13/2003 R 437 16 16 0 0 0 0 0 0 18B 12/14/2003 R 445 25 25 0 0 0 0 0 0 18B 12/14/2003 A 447 4 2 0 1 1 0 0 0 18B 12/16/2003 A 454 8 5 0 1 2 0 0 0 18B 12/16/2003 R 455 25 25 0 0 0 0 0 0 18C 12/8/2003 A 423 9 9 0 0 0 0 0 0 18C 12/10/2003 A 426 9 9 0 0 0 0 0 0 18C 12/12/2003 A 425 0 0 0 0 0 0 0 0

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81 PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 18C 12/13/2003 A 433 14 13 0 1 0 0 0 0 18C 12/13/2003 R 434 20 20 0 0 0 0 0 0 18C 12/14/2003 R 442 20 20 0 0 0 0 0 0 18C 12/14/2003 A 444 16 13 1 1 1 0 0 0 18C 12/16/2003 A 451 15 13 0 0 2 0 0 0 18C 12/16/2003 R 452 20 11 0 9 0 0 0 0 18C 12/17/2003 R 460 19 19 0 0 0 0 0 0 18C 12/17/2003 R 462 20 20 0 0 0 0 0 0 19A 1/23/2004 A 468 14 14 0 0 0 0 0 0 19A 1/25/2004 A 469 19 19 0 0 0 0 0 0 19A 1/26/2004 A 474 16 16 0 0 0 0 0 0 19A 1/27/2004 R 476 0 0 0 0 0 0 0 0 19A 1/27/2004 A 477 21 19 0 0 2 0 0 0 19A 1/28/2004 R 490 31 31 0 0 0 0 0 0 19A 1/29/2004 R 492 27 27 0 0 0 0 0 0 19A 1/29/2004 A 494 2 1 0 0 1 0 0 0 19B 1/24/2004 A 467 7 6 0 0 0 0 1 0 19B 1/25/2004 A 470 22 22 0 0 0 0 0 0 19B 1/26/2004 A 473 27 25 0 0 1 0 1 0 19B 1/27/2004 R 478 45 45 0 0 0 0 0 0 19B 1/27/2004 A 480 15 15 0 0 0 0 0 0 19B 1/28/2004 R 487 32 30 0 0 2 0 0 0 19B 1/28/2004 A 489 1 0 0 0 1 0 0 0 19B 1/29/2004 R 495 37 36 0 0 1 0 0 0 19B 1/30/2004 A 502 3 2 0 0 1 0 0 0 19B 1/31/2004 A 507 1 1 0 0 0 0 0 0 19C 1/24/2004 A 466 11 11 0 0 0 0 0 0 19C 1/25/2004 A 471 0 0 0 0 0 0 0 0 19C 1/26/2004 A 472 42 39 0 0 2 0 1 0 19C 1/27/2004 R 482 41 41 0 0 0 0 0 0 19C 1/27/2004 A 483 8 4 0 0 4 0 0 0 19C 1/28/2004 R 484 55 42 0 0 13 0 0 0 19C 1/28/2004 A 486 2 1 0 0 0 1 0 0 19C 1/29/2004 R 497 26 25 0 0 1 0 0 0 19C 1/30/2004 A 500 8 7 0 0 1 0 0 0 19C 1/31/2004 A 505 1 1 0 0 0 0 0 0 20A 3/6/2004 A 512 31 29 0 0 2 0 0 0 20A 3/7/2004 A 516 21 18 0 0 3 0 0 0 20A 3/8/2004 A 518 17 4 0 0 13 0 0 0 20A 3/8/2004 R 519 47 47 0 0 0 0 0 0 20A 3/9/2004 A 530 4 1 0 0 3 0 0 0 20A 3/9/2004 R 531 14 2 0 0 12 0 0 0 20A 3/14/2004 A 548 0 0 0 0 0 0 0 0 20B 3/5/2004 A 536 1 1 0 0 0 0 0 0 20B 3/6/2004 A 511 17 12 0 0 5 0 0 0 20B 3/7/2004 A 515 22 6 2 0 14 0 0 0 20B 3/8/2004 A 521 14 0 1 0 13 0 0 0 20B 3/8/2004 R 523 0 0 0 0 0 0 0 0 20B 3/9/2004 A 527 7 0 0 0 7 0 0 0

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82 PIG: DATE: A, R sample # # adults P. coerul. C. macellaria C. ruffifaces P. regina C. megaceph. C. livida C. vicina 20B 3/9/2004 R 528 47 1 0 0 46 0 0 0 20B 3/10/2004 R 537 35 0 0 0 35 0 0 0 20B 3/13/2004 A 543 8 3 3 0 2 0 0 0 20B 3/14/2004 A 545 23 1 5 0 17 0 0 0 20B 3/14/2004 R 547 6 0 0 1 5 0 0 0

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83 APPENDIX B RAW DATA: PRESERVED SPECIMENS PIG: DATE: sample # 1st instar 1-2 trans. 2nd instar 2-3 trans. 3rd instar total P. coerul. C. mac. C. ruf. P. regina ? 3rd 2B 11/21/2001 10B 0 0 0 0 0 0 0 0 0 0 0 3A 12/29/2001 11A 0 0 0 0 0 0 0 0 0 0 0 3A 1/11/2002 14 0 0 42 11 219 272 108 0 0 98 13 3B 12/29/2001 12A 0 0 0 0 90 90 90 0 0 0 0 3B 1/11/2002 18 0 0 0 1 152 153 148 0 0 4 0 3C 1/11/2002 16 0 0 8 2 107 117 107 0 0 0 0 4E 2/9/2002 29 19 4 64 16 151 254 107 0 0 44 0 5A 3/17/2002 34 66 17 56 14 0 153 0 0 0 0 0 5A 3/18/2002 46 0 0 17 19 79 115 76 0 0 0 3 5B 3/17/2002 39 82 35 80 2 0 199 0 0 0 0 0 5C 3/17/2002 37 34 3 129 5 14 185 14 0 0 0 0 5C 3/18/2002 43 254 52 16 0 0 322 0 0 0 0 0 5E 3/17/2002 41 82 10 24 0 0 116 0 0 0 0 0 5E 3/18/2002 45 0 0 88 6 7 101 7 0 0 0 0 5E 3/19/2002 48 0 2 61 12 34 109 34 0 0 0 0 6A 5/1/2002 54 17 8 73 10 13 121 13 0 0 0 0 6A 5/1/2002 58 1 2 75 8 59 145 59 0 0 0 0 6B 5/1/2002 58 0 0 0 0 0 0 0 0 0 0 0 6B 5/1/2002 60 174 1 0 0 0 175 0 0 0 0 0 6C 5/1/2002 62 5 2 59 48 10 124 10 0 0 0 0 6C 5/1/2002 64 1 0 14 7 63 85 63 0 0 0 0 6E 5/1/2002 67 12 5 73 0 10 100 10 0 0 0 0 6E 5/1/2002 69 78 5 39 10 8 140 8 0 0 0 0 6E 5/1/2002 71 0 0 17 4 33 54 33 0 0 0 0 7A 5/23/2002 79 1 0 19 1 175 196 175 0 0 0 0 8A 7/23/2002 88 138 7 0 0 0 145 0 0 0 0 0 8A 7/24/2002 93 2 0 31 6 65 104 65 0 0 0 0 8A 7/24/2002 95 51 20 50 0 1 122 1 0 0 0 0 8A 7/24/2002 96 8 1 89 8 2 108 1 0 1 0 0 8A 7/25/2002 106 0 0 14 12 164 190 0 0 164 0 0 8A 7/25/2002 108 0 0 47 8 241 296 0 0 241 0 0 8C 7/23/3003 90 0 0 0 0 0 0 0 0 0 0 0 8C 7/24/2002 104 9 1 76 40 77 203 77 0 0 0 0 8C 7/24/2002 105 15 1 41 28 54 139 54 0 0 0 0 8E 7/23/2002 89 10 4 40 0 0 54 0 0 0 0 0 8E 7/24/2002 99 14 1 111 6 16 148 15 0 1 0 0 8E 7/24/2002 100 400 53 49 0 0 502 0 0 0 0 0 8E 7/24/2002 101 43 14 28 1 10 96 8 0 2 0 0 8E 7/25/2002 110 0 0 3 2 70 75 0 0 70 0 0

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84 PIG: DATE: sample # 1st instar 1-2 trans. 2nd instar 2-3 trans. 3rd instar total P. coerul. C. mac. C. ruf. P. regina ? 3rd 9A 8/21/2002 120 2 1 126 7 17 153 17 0 0 0 0 9A 8/21/2002 120B 6 2 115 15 11 149 11 0 0 0 0 9A 8/21/2002 121 102 23 55 0 0 180 0 0 0 0 0 9A 8/21/2002 122 108 3 45 0 2 158 2 0 0 0 0 9A 8/22/2002 155 4 6 52 5 34 101 34 0 0 0 0 9A 8/22/2002 156 8 0 1 3 14 26 1 0 13 0 0 9A 8/22/2002 158 0 0 7 1 49 57 48 0 0 1 0 9B 8/21/2002 126 29 2 39 24 17 111 17 0 0 0 0 9B 8/21/2002 126B 16 2 42 18 10 88 9 0 0 1 0 9B 8/21/2002 127 183 0 0 0 0 183 0 0 0 0 0 9B 8/21/2002 128 12 0 49 12 2 75 2 0 0 0 0 9B 8/22/2002 150 0 0 6 6 54 66 53 0 0 1 0 9B 8/22/2003 152 13 2 36 22 1 74 0 0 1 0 0 9C 8/22/2002 136 3 0 8 10 32 53 5 0 27 0 0 9C 8/22/2002 138 0 0 2 1 40 43 33 0 0 7 0 9C 8/22/2002 139 9 2 3 3 9 26 4 0 5 0 0 9C 8/22/2002 140 116 17 23 9 18 183 18 0 0 0 0 9C 8/22/2002 141 0 0 11 4 40 55 38 0 1 1 0 9E 8/21/2002 131 0 0 8 3 31 42 31 0 0 0 0 9E 8/21/2002 131B 0 0 15 1 29 45 24 0 3 2 0 9E 8/21/2002 132 0 0 0 0 0 0 0 0 0 0 0 9E 8/21/2002 133 34 4 111 21 11 181 11 0 0 0 0 9E 8/22/2002 144 3 3 42 7 17 72 13 2 0 1 1 9E 8/22/2002 146 1 0 61 9 20 91 2 0 18 0 0 9E 8/22/2002 147 0 0 19 8 41 68 41 0 0 0 0 10A 9/25/2002 164 0 0 20 49 23 92 23 0 0 0 0 10A 9/26/2002 179 0 0 0 0 137 137 135 2 0 0 0 10A 9/26/2002 181 33 8 186 30 32 289 32 0 0 0 0 10B 9/25/2002 168 1 3 50 74 25 153 25 0 0 0 0 10B 9/25/2002 169 3 11 189 17 0 220 0 0 0 0 0 10B 9/26/2002 182 15 1 8 0 73 97 71 0 2 0 0 10B 9/26/2002 184 0 0 0 0 8 8 0 3 5 0 0 10B 9/26/2002 186 89 20 87 0 1 197 1 0 0 0 0 10C 9/25/2002 176 0 0 45 20 9 74 9 0 0 0 0 10C 9/27/2002 189 0 0 29 24 88 141 43 43 0 2 0 10E 9/25/2002 172 0 0 35 17 35 87 35 0 0 0 0 10E 9/25/2002 173 3 2 32 7 86 130 86 0 0 0 0 10E 9/27/2002 190 0 0 0 0 72 72 0 0 72 0 0 10E 9/27/2002 192 0 0 30 7 108 145 1 102 0 5 0 11A 10/26/2002 194 0 0 0 0 0 0 0 0 0 0 0 11A 10/27/2002 202 0 1 7 24 82 114 54 4 24 0 0 11A 10/27/2002 204 94 28 31 30 14 197 13 0 1 0 0 11A 10/28/2002 211 0 0 14 14 98 126 98 0 0 0 0 11A 10/28/2002 213 0 0 28 48 22 98 0 22 0 0 0 11B 10/26/2002 197 170 13 56 2 9 250 9 0 0 0 0 11C 10/28/2002 207 0 0 1 0 69 70 0 0 69 0 0 12A 12/7/2002 220 1 3 111 45 62 222 62 0 0 0 0 12A 12/8/2002 229 0 0 86 5 33 124 33 0 0 0 0 12A 12/12/2002 254 0 4 52 48 52 156 52 0 0 0 0

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85 PIG: DATE: sample # 1st instar 1-2 trans. 2nd instar 2-3 trans. 3rd instar total P. coerul. C. mac. C. ruf. P. regina ? 3rd 12A 12/13/2002 256 0 0 4 0 85 89 85 0 0 0 0 12A 12/13/2002 257 0 0 10 6 61 77 61 0 0 0 0 12A 12/14/2002 259 0 0 10 5 99 114 99 0 0 0 0 12B 11/30/2002 219 0 0 0 0 0 0 0 0 0 0 0 12B 12/7/2002 223 3 2 126 23 6 160 6 0 0 0 0 12B 12/8/2002 232 0 0 59 2 25 86 25 0 0 0 0 12B 12/10/2002 242 7 4 46 14 149 220 146 3 0 0 0 12B 12/10/2002 243 0 0 1 1 111 113 111 0 0 0 0 12B 12/10/2002 244 1 0 3 4 135 143 135 0 0 0 0 12B 12/12/2002 250 3 3 47 12 22 87 22 0 0 0 0 12B 12/12/2002 252 0 6 109 50 24 189 24 0 0 0 0 12C 12/7/2002 227 0 0 55 17 33 105 33 0 0 0 0 12C 12/10/2002 236 22 5 53 16 27 123 27 0 0 0 0 12C 12/10/2002 237 0 0 9 1 56 66 56 0 0 0 0 12C 12/10/2002 238 0 0 6 2 103 111 103 0 0 0 0 12C 12/12/2002 246 2 1 10 0 24 37 24 0 0 0 0 12E 12/7/2002 225 5 8 161 11 5 190 5 0 0 0 0 12E 12/10/2002 241 89 5 24 17 34 169 34 0 0 0 0 12E 12/12/2002 248 0 0 13 1 102 116 102 0 0 0 0 13B 1/3/2003 267 0 0 0 0 0 0 0 0 0 0 0 13B 1/4/2003 277 0 0 0 0 0 0 0 0 0 0 0 13B 1/5/2003 284 2 1 109 2 0 114 0 0 0 0 0 13B 1/9/2003 293 2 1 19 6 69 97 69 0 0 0 0 13C 1/3/2003 271 12 5 76 6 1 100 0 1 0 0 0 13C 1/4/2003 275 0 1 29 25 51 106 51 0 0 0 0 13C 1/5/2003 283 0 0 72 16 33 121 33 0 0 0 0 13C 1/8/2003 291 0 0 6 1 101 108 101 0 0 0 0 13C 1/9/2003 299 0 0 38 5 42 85 42 0 0 0 0 13C 1/11/2003 301 3 0 84 7 3 97 2 0 0 0 1 13E 1/3/2003 269 2 0 69 15 8 94 8 0 0 0 0 13E 1/4/2003 273 21 3 75 46 28 173 28 0 0 0 0 13E 1/5/2003 280 0 0 12 3 61 76 61 0 0 0 0 13E 1/6/2003 287 0 0 0 1 83 84 83 0 0 0 0 13E 1/8/2003 289 0 0 23 37 47 107 42 0 0 0 5 13E 1/9/2003 296 0 0 3 0 54 57 54 0 0 0 0 14A 3/6/2003 305 0 0 2 1 71 74 0 0 0 71 0 14A 3/6/2003 307 44 0 263 4 7 318 6 0 0 0 1 14A 3/8/2003 315 0 0 8 0 97 105 97 0 0 0 0 14A 3/8/2003 317 0 0 2 0 47 49 47 0 0 0 0 14B 3/5/2003 303 0 0 10 2 52 64 27 0 0 25 0 14E 3/7/2003 310 2 0 73 4 37 116 37 0 0 0 0 14E 3/7/2003 313 5 0 60 6 29 100 29 0 0 0 0 15A 4/5/2003 341 1 0 0 11 27 39 27 0 0 0 0 15A 4/6/2003 354 0 0 1 1 66 68 49 0 0 17 0 15B 4/4/2003 332 32 6 130 13 0 181 0 0 0 0 0 15B 4/5/2003 344 105 0 89 8 4 206 4 0 0 0 0 15C 4/4/2003 328 60 5 92 0 0 157 0 0 0 0 0 15C 4/4/2003 329 0 0 0 0 0 0 0 0 0 0 0 15C 4/5/2003 338 15 1 102 1 7 126 7 0 0 0 0

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86 PIG: DATE: sample # 1st instar 1-2 trans. 2nd instar 2-3 trans. 3rd instar total P. coerul. C. mac. C. ruf. P. regina ? 3rd 15C 4/6/2003 351 1 0 20 2 70 93 70 0 0 0 0 15E 4/4/2003 335 44 10 122 8 2 186 0 0 0 2 0 15E 4/5/2003 347 5 0 30 4 67 106 67 0 0 0 0 15E 4/6/2003 356 0 0 0 0 58 58 10 25 0 23 0 16B 4/28/2003 374 54 10 15 0 0 79 0 0 0 0 0 16B 4/29/2003 382 0 0 106 0 104 210 104 0 0 0 0 16C 4/28/2003 368 1 0 59 6 10 76 6 3 0 1 0 16C 4/29/2003 377 0 0 63 0 32 95 23 9 0 0 0 16C 4/30/2003 385 0 0 4 0 105 109 0 105 0 0 0 16C 5/1/2003 390 0 0 0 0 67 67 0 67 0 0 0 16C 5/1/2003 391 0 0 0 1 61 62 2 58 0 1 0 16C 5/1/2003 392 0 0 0 0 41 41 0 41 0 0 0 16E 4/27/2003 364 28 9 13 0 0 50 0 0 0 0 0 16E 4/28/2003 371 0 0 34 0 43 77 43 0 0 0 0 16E 4/29/2003 380 0 2 2 1 134 139 134 0 0 0 0 16E 4/30/2003 387 0 0 1 0 54 55 17 29 0 7 1 17B 6/14/2003 402 99 17 113 13 22 264 22 0 0 0 0 17B 6/15/2003 420 0 1 12 3 44 60 44 0 0 0 0 17C 6/14/2003 404 192 0 9 0 40 241 40 0 0 0 0 17C 6/15/2003 415 0 0 1 1 29 31 7 4 18 0 0 17C 6/15/2003 418 0 0 6 0 75 81 11 64 0 0 0 17E 6/13/2003 400A 153 6 3 0 0 162 0 0 0 0 0 17E 6/14/2003 407 8 3 73 3 73 160 73 0 0 0 0 17E 6/15/2003 410 0 0 77 1 45 123 35 2 3 5 0 17E 6/15/2003 413 0 0 14 4 60 78 54 5 0 0 1 18A 12/13/2003 440 50 10 88 2 0 150 0 0 0 0 0 18A 12/14/2003 449 28 0 270 2 4 304 2 0 0 2 0 18A 12/16/2003 458 4 0 121 6 14 145 10 0 0 4 0 18A 12/20/2003 465 1 8 31 0 52 92 50 0 2 0 0 18B 12/12/2003 431 129 21 9 3 0 162 0 0 0 0 0 18B 12/13/2003 438 0 0 68 0 2 70 2 0 0 0 0 18B 12/14/2003 446 4 0 84 20 36 144 12 0 0 24 0 18B 12/16/2003 456 0 0 92 8 85 185 77 0 0 8 0 18C 12/13/2003 435 10 2 178 4 0 194 0 0 0 0 0 18C 12/14/2003 443 2 0 0 8 192 202 176 0 0 16 0 18C 12/16/2003 453 0 0 19 2 94 115 88 0 6 0 0 18C 12/17/2003 461 0 0 0 0 70 70 70 0 0 0 0 18C 12/17/2004 463 0 0 15 2 99 116 99 0 0 0 0 19A 1/27/2004 475 122 12 110 0 0 244 0 0 0 0 0 19A 1/28/2004 491 12 0 138 6 18 174 4 0 0 14 0 19A 1/29/2004 493 4 0 68 0 38 110 24 0 0 14 0 19A 1/30/2004 503 0 0 11 0 57 68 57 0 0 0 0 19A 1/31/2004 508 0 0 2 1 58 61 58 0 0 0 0 19B 1/27/2004 479 224 26 44 0 0 294 0 0 0 0 0 19B 1/28/2004 488 107 0 3 0 0 110 0 0 0 0 0 19B 1/29/2004 496 46 2 128 6 12 194 0 0 0 12 0 19B 1/30/2004 501 10 0 70 12 38 130 20 0 0 18 0 19B 1/31/2004 506 0 0 23 1 61 85 61 0 0 0 0 19C 1/27/2004 481 88 0 0 0 0 88 0 0 0 0 0

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87 PIG: DATE: sample # 1st instar 1-2 trans. 2nd instar 2-3 trans. 3rd instar total P. coerul. C. mac. C. ruf. P. regina ? 3rd 19C 1/28/2004 485 76 0 74 0 0 150 0 0 0 0 0 19C 1/29/2004 498 48 0 150 10 2 210 0 0 0 2 0 19C 1/30/2004 499 0 0 56 1 17 74 17 0 0 0 0 19C 1/31/2004 504 0 0 70 4 46 120 46 0 0 0 0 20A 3/8/2004 520 0 0 138 4 16 158 14 0 0 2 0 20A 3/9/2004 532 13 0 67 2 45 127 38 0 0 7 0 20B 3/8/2004 522 0 0 280 20 28 328 28 0 0 0 0 20B 3/9/2004 529 24 0 70 8 22 124 22 0 0 0 0 20B 3/10/2004 538 0 0 20 0 68 90 64 2 0 2 0 20B 3/13/2004 544 0 0 10 0 61 71 0 4 0 57 0 20B 3/14/2004 546 0 0 0 1 65 66 1 3 0 61 0

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88 LIST OF REFERENCES Abell, D. H., S. S. Wasti, and G. C. Hartmann. 1982. Saprophagous arthropod fauna associated with turtle carrion. Appl. Entomol. Zool. 17: 301-307. Adair, T. W. 1999. Three species of blowfly (Diptera: Calliphoridae) collected from a human stillborn infant in the Rocky Mount ains of Colorado. J. Med. Entomol. 36: 236-237. Aldrich, J. M. 1916. Sarcophaga and Allies. Thomas Say Foundation, Lafayette, IN. 301 pp. Anderson, G. S. 1998. Wildlife forensic ento mology: determining time of death in two illegally killed black bear cubs J. Forensic Sci. 44: 856-859. Anderson, G. S. 2001. Insect succession on carri on and its relationship to determining time of death. In J. H. Byrd and J. L. Castner [eds.], Forensic Entomology: The Utility of Arthropods in Legal Invest igations. CRC Press, Boca Raton. pp. 143-175. Anderson, G. S., and S. L. VanLaerhoven. 1996. Initial studies on insect succession on carrion in southwestern British Columbia. J. Forensic Sci. 41: 617-625. Ashworth, J. R., and R. Wall. 1994. Responses of the sheep blowflies Lucilia sericata and L. cuprina to odour and the development of semiochemical baits. Med. Vet. Entomol. 8: 303-309. Bass, W. M. 2001. Preface. In J. H. Byrd and J. L. Castner [eds.], Forensic Entomology: The Utility of Arthropods in Legal Investigations. CRC Press, Boca Raton. pp. ixx. Baumgartner, D. L. 1986. The hairy maggot blow fly Chrysomya Rufifaces (Maquart) confirmed in Arizona. J. Entomol. Sci. 21: 130-132. Baumgartner, D. L. 1988. Spring season surv ey of the urban bl owflies (Diptera: Calliphoridae) of Chicago, Illinois. Great Lakes Entomol. 21: 119-121. Baumgartner, D. L., and B. Greenberg. 1984. The Genus Chrysomya (Diptera: Calliphoridae) in the new worl d. J. Med. Entomol. 21: 105-113. Benecke, M. 1998. Six forensic entomology cases: description and commentary. J. Forensic Sci. 43: 797-805.

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89 Benecke, M. 2001. A brief history of forensic entomology. Forensic Sci. Int. 120: 2-14. Blackith, R. E., and R. M. Blackith. 1989. Insect infestations of small corpses. J. Nat. Hist. 24: 699-709. Bornemissza, G. F. 1957. An analysis of arth ropod succession in carrion and the effect of its decomposition on the soil fauna. Aust. J. Zool. 5: 1-12. Bourel, B., L. Martin-Bouyer, V. Hedouin, J. -C. Cailliez, D. Derout, and D. Gosset. 1999. Necrophilous insect succession on ra bbit carrion in sand dune habitats in northern France. J. Med. Entomol. 36: 420-425. Braack, L. E. O. 1981. Visitation patterns of pr incipal species of the insect-complex at carcasses in the Kruger Nationa l Park. Koedoe 24: 33-49. Byrd, J. H. 1998. Temperature dependent deve lopment and computer modeling of insect growth: its application to forensic entomology. Unpublished Dissertation, Department of Entomology and Nematology, University of Florida, Gainesville, FL. 196 pp. Byrd, J. H., and J. F. Butler. 1996. Effects of temperature on Cochliomyia macellaria (Diptera: Calliphoridae) developmen t. J. Med. Entomol. 33: 901-905. Byrd, J. H., and J. F. Butler. 1997. Effects of temperature on Chrysomya rufifacies (Diptera: Calliphoridae) developmen t. J. Med. Entomol. 34: 353-357. Byrd, J. H., and J. F. Butler. 1998. Effects of temperature on Sarcophaga haemorrhoidalis (Diptera: Sarcophagidae) development. J. Med. Entomol. 35: 694698. Byrd, J. H., and J. C. Allen. 2001. The development of the black blow fly, Phormia regina. Forensic Sci. Int. 120: 79-88. Byrd, J. H., and J. L. Castner [eds.]. 2001a. Forensic Entomology: The Utility of Arthropods in Legal Investigations. CRC Press, Boca Raton. 418 pp. Byrd, J. H., and J. L. Castner. 2001b Insects of forensic importance. In J. H. Byrd and J. L. Castner [eds.], Forensic Entomology: The Utility of Arthropods in Legal Investigations. CRC Press, Boca Raton. pp. 43-79. Campobasso, C. P., and F. Introna. 2001. The fore nsic entomologist in the context of the forensic pathologist's role. Fo rensic Sci. Int. 120: 132-139. Campobasso, C. P., G. Di Vella, and F. In trona. 2001. Factors affecting decomposition and Diptera colonization. Fore nsic Sci. Int. 120: 18-27.

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90 Castner, J. L. 2001. General Entomology and Arthropod Biology. In J. H. Byrd and J. L. Castner [eds.], Forensic Entomology: The Utility of Arthropods in Legal Investigations. CRC Press, Boca Raton. pp. 17-41. Catts, E. P., and N. H. Haskell. 1990. Entomo logy & Death-A Procedural Guide. Joyce's Print Shop, Inc., Clemson, SC. 182 pp. Catts, E. P., and M. L. Goff. 1992. Forensic entomology in criminal investigations. Annu. Rev. Entomol. 37: 253-272. Coe, M. 1978. The decomposition of elephant car cases in the Tsavo (East) National Park, Kenya. J. Arid. Environ. 1: 71-86. Cornaby, B. W. 1974. Carrion reduction by animal s in contrasting tropical habitats. Biotropica 6: 51-63. Cragg, J. B. 1956. The olfactory behavior of Lucilia species (Diptera) under natural conditions. Ann. Appl. Biol. 44: 467-477. Cragg, J. B., and B. A. Thurston. 1949. The r eactions of blowflies to organic sulphur compounds and other materials used in traps. Parasitology 40: 187-194. Cragg, J. B., and P. Cole. 1956. Laboratory st udies on the chemosensory reactions of blowflies. Ann. Appl. Biol. 44: 478-491. Cushing, E. C., and H. E. Parish. 1938. Seasonal variations in the abundance of Cochliomyia spp., Phormia spp., and other flies in Menard County, Tex. J. Med. Entomol. 31: 764-769. Davidson, J. 1944. On the relationship between temperature and rate of development of insects at constant temperatur es. J. Anim. Ecol. 13: 26-38. Davis, J. B., and M. L. Goff. 2000. Decompos ition patterns in terres trial and intertidal habitats on Oahu Island and Coconut Island, Hawaii. J. Forensic Sci. 45: 836-842. deCarvalho, L. M. L., and A. X. Linhares. 2001. Seasonality of insect succession and pig carcass decomposition in a natural forest ar ea in southeastern Brazil. J. Forensic Sci. 46: 604-608. deCarvalho, L. M. L., P. J. Thyssen, A. X. Linhares, and F. A. B. Palhares. 1999. A checklist of arthropods associated wi th pig carrion and human corpses in southeastern Brazil. Mem. In st. Oswaldo Cruz. 95: 135-138. Denno, R. F., and W. R. Cothran. 1975. Co mpetitive interactions and ecological strategies of sarcophagid and callipho rid flies inhabiting rabbit carrion. Ann. Entomol. Soc. Amer. 69: 109-113.

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92 Goodbrod, J. R., and M. L. Goff. 1990. Effect s of larval population density on rates of development and interactions between two species of Chrysomya (Diptera: Calliphoridae) in laboratory cultur e. J. Med. Entomol. 27: 338-343. Gordh, G., and D. Headrick. 2001. A Dictiona ry of Entomology. CA BI Publishing, New York. Green, A. A. 1951. The control of blowflies infesting slaughter-houses. Ann. Appl. Biol. 38: Greenberg, B. 1985. Forensic entomology: case studies. Bull. Entomol. Soc. Amer. 2528. Greenberg, B. 1988. Chrysomya megacephala (F.) (Diptera:Callipho ridae) collected in North America and notes on Chrysomya species present in the New World. J. Med. Entomol. 25: 199-200. Greenberg, B. 1990. Nocturnal oviposition behavior of blow flies (Diptera: Calliphoridae). J. Med. Entomol. 27: 807-810. Greenberg, B. 1991. Flies as forensic i ndicators. J. Med. Entomol. 28: 565-577. Greenberg, B., and J. C. Kunich. 2002. Ento mology and the Law: Flies as Forensic Indicators. Cambridge University Press, Cambridge. 306 pp. Hall, D. G. 1948. The Blowflies of North America. Thomas Say Foundation, Lafayette, IN. 477 pp. Hall, M. 2003. President's letter, Europ ean Association of Forensic Entomology. www.eafe.org/index.php?section=main&p age=Letter. September 17, 2003. Hall, M. J., R. Farkas, F. Kelemen, M. Ho sier, and J. M. El-Khoga. 1995. Orientation of agents of wound myiasis to hosts and ar tificial stimuli in Hungary. Med. Vet. Entomol. 9: 77-84. Hall, R. D. 2001. Introduction: perceptions and status of forensic entomology. In J. H. Byrd and J. L. Castner [eds.], Forensic Entomology: The Utility of Arthropods in Legal Investigations. CRC Press, Boca Raton. pp. 1-15. Hall, R. D., and L. H. Townsend. 1977. The Insects of Virginia: No. 11. In The Blow Flies of Virginia (D iptera: Calliphoridae). Virginia Polytechnic Institute and State University, Blacksburg, VA. pp. 48 Hall, R. D., and K. E. Doisy. 1993. Length of time after death: effect on attraction and oviposition or larviposition of midsummer blow flies (D iptera: Calliphoridae) and flesh flies (Diptera: Sarcophagidae) of medicolegal importance in Missouri. Ann. Entomol. Soc. Amer. 86: 589-593.

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93 Hammack, L., and G. G. Holt. 1983. Responses of gravid screwworm flies, Cochliomyia hominivorax to whole wounds, wound fluid, and a standard blood attractant in olfactometer tests. J. Chem. Ecol. 9: 913-922. Haskell, N. H. 1989. Calliphoridae of pig car rion in northwest Indiana: a seasonal comparative study. Unpublished Thesis, Colleg e of Agriculture, Purdue University, Lafayette. 57 pp. Haskell, N. H., R. D. Hall, V. J. Cerve nka, and M. A. Clark. 1997. On the body: insect's life stage presence and their postmortem artifacts. In W. D. Haglund and M. H. Sorg [eds.], Forensic Taphonomy. CRC Press, Boca Raton. pp. 415-448. Introna, F., Jr., T. W. Suman, and J. E. Smialek. 1991. Sarcosaprophagous fly activity in Maryland. J. Forensic Sci. 36: 238-243. James, M. T. 1947. The Flies That Cause Myia sis in Man. United States Department of Agriculture, Washi ngton, D.C. 175 pp. James, M. T. 1955. The blowflies of California (Diptera: Calliphoridae ). Bull. Calif. Ins. Surv. 4: 1-34. Jiron, L., and V. Cartin. 1981. Insect succession in the decomposition of a mammal in Costa Rica. N. Y. Ento mol. Soc. 89: 158-165. Johnson, M. D. 1975. Seasonal and microseral va riations in the inse ct populations on carrion. Am. Midl. Nat. 93: 79-90. Joy, J. E., M. L. Herrell, and P. C. Roge rs. 2002. Larval fly activity on sunlit versus shaded raccoon carrion in southwestern West Virginia with special reference to the black blowfly (Diptera: Calliphorida e). J. Med. Entomol. 39: 392-397. Kamal, A. S. 1958. Comparative study of thirteen species of sarcosaphrophagous Calliphoridae and Sarcophagi dae (Diptera) I. Bionomics Ann. Entomol. Soc. Am. 51: Kashyap, V. K., and V. V. Pillay. 1989. Effi cacy of entomological method in estimation of postmortem interval: A comparative anal ysis. Forensic Sci. Int. 40: 245-250. Kocarek, P. 2001. Diurnal patterns of postfeedi ng larval dispersal in carrion blowflies (Diptera: Calliphoridae). Eur. J. Entomol. 98: 117-119. Knipling, E. F. 1936. Some specifi c taxonomic characters of common Lucilia larvae-Calliphorniae--Diptera. Iowa St ate Coll. J. Sci. 10: 275-293. Lane, R. P. 1975. An investigation into bl owfly (Diptera: Calliphoridae) succession on corpses. J. Nat. Hist. 9: 581-598.

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94 Lewerenz, D. 2003. 'CSI' boosts interest in forensic science, Associated Press. http://www.sacticket.com/24hour/ent ertainment/tv/news/story/972073p. September 16, 2003. 2003. Lord, W. D. 1990. Case histories of th e use of insects in investigations. In N. H. Haskell and E. P. Catts [eds.], In Entomology and Death: A procedur al guide. Forensic entomology specialties, Clemson, SC. pp. 9-37. Lord, W. D., and J. F. Burger. 1984a. Ar thropods asscoiated with herring gull ( Larus argentatus ) and great black-backed gull ( Larus marinus ) carrion on islands in the gulf of Maine. Environ. Entomol. 13: 1261-1268. Lord, W. D., and J. F. Burger. 1984b. Arth ropods associated with harbor seal ( Phoca vitulina ) carcasses stranded on islands along the New England coast. Int. J. Entomol. 26: 282-285. Mann, R. W., W. M. Bass, and L. Meadow s. 1990. Time since death and decomposition of the human body: variables and observat ions in case and experimental field studies. J. Forensic Sci. 35: 103-111. Martin, C. S., C. E. Carlton, and C. L. M eek. 1996. New distribution record for the hairy maggot blow fly Chrysomya rufifacies Southw. Entomol. 21: 4. McKnight, B. E. 1981. The Washing Away of Wr ongs: Forensic Medicine in ThirteenthCentury China. The University of Mich igan Center for Chinese Studies, Ann Arbor. 181 pp. Mgnin, P. 1894. La faune de cadavres. Applic ation de l'entomologie la mdicine lgale, Encyclopedie scientifique de s Aides-Mmoire, Masson, Paris. Morris, M. C., L. Morrison, M. A. Joyce, and B. Rabel. 1998. Trapping sheep blowflies with lures based on bacterial cultures. Aust. J. Exp. Agri. 38: 125-130. Nelson, E. L. 1999. Estimation of short-term postmortem interval utilizing core body temperature: a new algorithm. Forensic Sci. Int. 109: 31-38. Nilssen, A. C., B. . Tmmers, R. Sc hmid, and S. B. Evensen. 1996. Dimethyl trisulphide is a strong attractant for some calliphorids and a muscid but not for the reindeer oestrids Hypoderma tarandi and Cephenemyia trompe Entomol. Exp. Appl. 79: 211-218. Nuorteva, P. 1977. Sarcosaphrophagous insects as forensic indicators. In C. G. Tedeschi, W. G. Eckert and L.G. Tedeshi [eds.], Forensic Medicine: A Study in Trauma and Environmental Hazards. W. B. Sa unders and Company, Toronto. pp. 1072-1095. Parish, H. E., and E. C. Cushing. 1938. Locations for blowfly traps: abundance and activity of blowflies and ot her flies in Menard County, Tex. J. Econ. Entomol. 31: 750-763.

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95 Payne, J. A. 1965. A summer carrion study of the baby pig Sus scrofa Linnaeus. Ecology 46: 592-602. Payne, J. A., and E. W. King. 1970. Coleoptera associated with pig carrion. Entomol. Monthly Mag. 105: 224-232. Peters, S. L. 2003. Temperature variations of dipteran larval masses analyzed on Florida black bear carcasses. Unpublished Th esis, Department of Entomology and Nematology, University of Florida, Gainesville. 93 pp. Portrait of a Prince. 2004. Asso ciated Press. Gainesville S un. Gainesville, Florida. April 29, 2004. Page 2. Putnam, R. J. 1978. The role of carrion-frequenting arthropods in the decay process. Ecol. Entomol. 3: 133-139. Reed, J., H. B. 1958. A study of dog carcass communities in Tennessee, with special reference to the insects. Am. Midl. Nat. 59: 213-245. Richards, E. N., and M. L. Goff. 1997. Arth ropod succession on exposed carrion in three contrasting tropical habitats on Hawaii Is land, Hawaii. J. Med. Entomol. 34: 328338. Rodriguez, W. C., and W. M. Bass. 1983. Inse ct activity and its relationship to decay rates of human cadavers in east Tennessee. J. Forensic Sci. 28: 423-432. Seago, J. M. 1953. Fly larvae: pictor ial key to some common species. In U.S. Department of Health, Education, and Welfare. Shahid, S. A., R. D. Hall, N. H. Haskell, and R. W. Merritt. 1999. Chrysomya rufifacies (Macquart)(Diptera: Calli phoridae) established in th e vicinity of Knoxville, Tennessee, USA. J. Forensic Sci. 45: 896-897. Shean, B. S., L. Messinger, and M. Papw orth. 1993. Observations of differential decomposition on sun exposed v. shaded pig ca rrion in coastal Washington state. J. Forensic Sci. 38: 938-949. Sherman, R.A., and Edward A. Pechter. 1988. Maggot therapy: A review of the therapeutic applications of fly larvae in human medicine, especially for treating osteomyelitis. Med. Vet. Entom. 2: 225-230. Singh, D. 2001. Further observations on the nocturn al oviposition behavior of blow flies (Diptera: Calliphoridae). Forens ic Sci. Int. 120: 124-126. Smith, K. G. V. 1975. The faunal succession of insects and other inve rtebrates on a dead fox. Entomol. Gaz. 26: 277-287.

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96 Smith, K. G. V. 1986. A Manual of Forensic Entomology. British Museum of Natural History, London. 207 pp. Stensmyr, M., I. Urru, I. Collu, M. Cela nder, B. S. Hansson, and A.-M. Angioy. 2002. Rotting smell of dead-horse ar um florets. Nature 420: 625. Tantawi, T. I., and B. Greenberg. 1993. Chrysomya albiceps and C. rufifacies (Diptera: Calliphoridae): contribution to an ongoi ng taxonomic problem. J. Med. Entomol. 30: 646-648. Tantawi, T. I., E. M. El-Kady, B. Gree nberg, and H. A. El-Ghaffar. 1996. Arthropod succession on exposed rabbit carrion in Al exandria, Egypt. J. Med. Entomol. 33: 566-580. Tenorio, F. M., J. K. Olson, and C. J. Coates. 2003. Decomposition studies, with a catalog and description of forensically impor tant blow flies (Dip tera: Calliphoridae) in central Texas. Southw. Entomol. 28: 37-45. Tessmer, J. W., and C. L. Meek. 1996. Disp ersal and distribution of Calliphoridae (Diptera) immatures from animal carcasse s in southern Louisiana. J. Med. Entomol. 33: 665-669. Tessmer, J. W., C. L. Meek, and V. L. Wr ight. 1995. Circadian patterns of oviposition by necrophilous flies (Diptera: Calliphorida e) in southern Louisiana. Southw. Entomol. 20: 439-445. Tomberlin, J. K., and P. H. Adler. 1998. S easonal colonization and decomposition of rat carrion in water and on land in an open fi eld in South Carolina. J. Med. Entomol. 35: 704-709. Tullis, K., and M. L. Goff. 1987. Arthropod succ ession in exposed carrion in a tropical rainforest on O'ahu Island, Hawai'i. J. Med. Entomol. 24: 332-339. Wall, R., and M. L. Warnes. 1994. Responses of the sheep blowfly Lucilia sericata to carrion odour and carbon dioxide. Entomol. Exp. Appl. 73: 239-246. Wall, R., and P. Fisher. 2001. Visual and olf actory cue interaction in resource-location by the blowfly, Lucilia sericata Physiol. Entomol. 26: 212-218. Wall, R., C. H. Green, N. French, and K. L. Morgan. 1992. Development of an attractive target for the sheep blowfly Lucilia sericata Med. Vet. Entomol. 6: 67-74. Wardle, R. A. 1921. The protection of meat commodities against blowflies. Ann. Appl. Biol. 9: 1-9. Watson, E. J., and C. E. Carlton. 2003. Spring succession of necrophilous insects on wildlife carcasses in Louisiana. J. Med. Entomol. 4: 338-347.

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97 Wells, J. D., and J. King. 2001. Incidence of precocious egg development in flies of forensic importance (Calliphoridae). Pan Pac. Entomol. 77: 235-239. Wells, J. D., J. H. Byrd, and T. I. Tant awi. 1999. Key to third-instar Chrysomyinae (Diptera: Calliphoridae) from carrion in the continental United States. J. Med. Entomol. 36: 638-641. White, S. R., D. Aubertin, and J. Smart. 1940. The Fauna of British India including the remainder of the Oriental region. Ta ylor and Francis, Ltd., London. 288 pp. Wolff, M., A. Uribe, A. Ortiz, and P. Duque. 2001. A preliminary of study forensic entomology in Medellin, Columbia. Forensic Sci. Int. 120: 53-59.

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98 BIOGRAPHICAL SKETCH Susan V. Gruner was born July 5, 1956, in Wiesbaden, Germany. After her father-an O.S.I. agent--retired from the Air For ce, her family moved to the small town of Mattapoisett, MA, a small coastal town not far from Cape Cod. Sue graduated from Old Rochester Regional High School in 1974. Her firs t attempt at a college career was a brief but unsuccessful stint at EmbryRiddle Aeronautical University, Daytona Beach, Florida. After the academic attempt in Daytona B each crashed and burned, Susan got a job in the audio industry in which she worked until 1993. In 1994, she decided to go back to college and enrolled at Santa Fe Community College in Gainesville, FL. Before classes began, Susan received an official letter i ndicating that she was on academic probation due to the unfortunate academic history that came back to haunt her after 20+ years when she flunked out of Embry-Riddle. Determined to succeed in the face of adversity, Sue forged ahead. She hit the books hard, managed to learn the metric system, successfully plodded through 3 semesters of chemistry involvi ng hair loss and sleepless nights, and was eventually inducted into the Phi Theta Kappa Honor Society in 1995. Both the academic probation letter and Phi Th eta Kappa certificate have been saved for posterity. Sue graduated from S.F.C.C. in 1996 and was accepted at UF immediately, where she began working on her B.S. in the College of Agriculture with a major in entomology. During one of her first entomology classes with Dr. John Strayer, she received a hand-out about human decomposition studies at the Body Fa rm in TN. At that point, Susan knew what she wanted to do: forensic entomology. But, how?

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99 Sue graduated with her B.S. in the College of Agricultural and Life Sciences in the last semester of the millennium, December, 1999. Dr. Jon Allen signed off on her graduate school entrance form, Dan Slone gave her part of a National Institute of Justice grant and introduced her to Neal Haskell in February of 2001, and the rest has yet to be written.


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Title: The Forensically Important Calliphoridae (Insecta: Diptera) of Pig Carrion in Rural North-Central Florida
Physical Description: Mixed Material
Copyright Date: 2008

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THE FORENSICALLY IMPORTANT CALLIPHORIDAE (INSECTA: DIPTERA) OF
PIG CARRION IN RURAL NORTH-CENTRAL FLORIDA
















By

SUSAN V. GRUNER


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA


2004






























Copyright 2004

by

Susan V. Gruner

































For Jack, Rosamond, and Michael















ACKNOWLEDGMENTS

The successful completion of this thesis would not have been possible without the

support, encouragement, understanding, guidance, and physical help of many colleagues,

friends, and family. My mother, Rosamond, edited my thesis despite her obvious distaste

for anything related to maggots. My husband, Michael, cheerfully allowed more than

most spouses could bear. And when it got really cold outside, he only complained two or

three times when I kept multiple containers of stinking liver and writhing maggots on the

kitchen counter. He also took almost all of the fantastic photos presented in this thesis

and for my presentation. Finally, Michael had the horrible job of inserting an arrow with

twelve temperature probes into pig "E."

I am also indebted to Dan and Jenny Slone, Jon Allen, Debbie Hall, Jane and

Buthene Haskell, Dan and Zane Greathouse, owners of Greathouse Butterfly Farm,

Aubrey Bailey, and the National Institute of Justice.

My professional colleagues, John Capinera, Marjorie Hoy, and Neal Haskell,

guided and encouraged me through my research and writing every step of the way.

Although I doubt that John Capinera was greatly interested in maggots, his support and

enthusiasm in response to my enthusiasm are very much appreciated. Marjorie Hoy's

beneficial advice was always appreciated and on occasion, she offered a shoulder on

which to cry. Neal Haskell, as usual, went above and beyond the call of duty in every

aspect of my time spent as an M.S. graduate student. I thank them all for their support.

Finally, I thank the pigs which were sacrificed in the name of forensic science.

















TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ................................................................................................. iv

L IST O F TA B L E S ......... ............ .......... .......................... ...... .. .............. .. vii

LIST OF FIGURES .................. ....................... ................. viii

ABSTRACT ........ .............. ............. ...... ...................... xi

CHAPTER

1 INTRODUCTION AND LITERATURE REVIEW ....................................................1

Definition of and Scope of Forensic Entomology .................................................. 1
H history of Forensic Entom ology ........................................................ ...............2
Current Status of Forensic Entom ology ............................................ .....................6
B biology of C alliphoridae .............................................................. ....................... 8
Insect Succession on Carrion.................. .............................. ............................. 10
Factors that Affect Blow Fly Succession on Carrion ...........................................12
Biology of Hum an D ecom position................................... ............................. ....... 14
F further A areas for Stu dy ...................................................................... ..................15

2 M ATERIALS AND M ETHOD S ........................................ ......................... 17

S tu d y S ite ........................................................................................1 7
D ata Collection ................................................................ .. ... ............ 18
P ro to c o l fo r D ay 1 ......................................................................................... 18
Protocol for D ay 2 Onw ard.......................................... ......... 19
M eteorological M easurements ................................ ............... 19
R hearing Procedures.......... ........................................................ .............. ... 20
Pupation Substrate .......................................................................... ............... 21

3 R E SU LT S ................................................... ............. ........29

Species of Calliphoridae Collected on Pig Carrion............................... ..................29
Seasonal Distribution and Succession of Calliphoridae Species.......................30
Collection 1, November 16-21, 2001 ................................... ............... 31
Collection 2, December 29, 2001-January 11, 2002 ............... ............ .....31









Collection 3, February 5-9, 2002.................................... ........ ............... 31
Collection 4, M arch 15-19, 2002.................................... ......... ............... 32
Collection 5, A pril 29-M ay 1, 2002 ....................................... ............... 32
Collection 6, M ay 20-23, 2002......... ......... ........... ... .................... 32
Collection 7, July 22-25, 2002 ........................................ ........................ 32
Collection 8, A ugust 19-23, 2002.................................... ........................ 33
Collection 9, September 23-27, 2002 ..... ................................33
Collection 10, October 26-28, 2002 ........................................ ............... 34
Collection 11, November 30-December 14, 2002 ....................................34
Collection 12, December 30, 2002-January 11, 2003 ...................................... 35
Collection 13, M arch 2-8, 2003 ............ ......... .. ......... ............. ............... 35
Collection 14, April 1-6, 2003 ............ ........ .. ......... ............... ............... 35
Collection 15, April 26-M ay 1, 2003 ...................................... ............... 36
Collection 16, June 12-15, 2003 .......... ......... ..................... ............ ... 36
Collection 17, D ecem ber 8-20, 2003........... .............................. ...... ............. 36
Collection 18, January 23-31, 2004.................................... ...... ............... 37
Collection 19, M arch 5-14, 2004.................................... ......... ............... 37

4 D ISCU SSION ......... .. ....... ............................................................ ........ 68

APPENDIX

A RAW DATA: FLIES COLLECTED AS ADULTS OR REARED FROM
LARVAE .................. ........ ................................ ........ ... ...... .. 75

B RAW DATA: PRESERVED SPECIMENS................... .. .................... 83

L IST O F R E F E R E N C E S ......... ................. ...................................................................88

BIO GRAPH ICAL SK ETCH .................................................. ............................... 98
















LIST OF TABLES

Table p

2-1 Dates of pig deposition, approximate pig weight, and time each pig was placed
on site for sampling dates from November, 2001 to October, 2002......................27

2-2 Mean temperatures ( SD) during the study at the Earleton, Florida site................28
















LIST OF FIGURES


Figure page

2-1 A caged pig carcass is shown at Greathouse Butterfly Farm property in Earleton,
F lo rid a ........................................................... ................ 2 3

2-2 Wire cage with bungee cords and tent stakes hammered into the ground ...............24

2-3 A HOBO temperature data logger is sealed inside a Gladware container ..........25

2-4 The data logger was hung from bush at left and temperature probe was placed
on the ground (circled) to record ground temperature near the pig carcass in
E arleton, F lorida............ .......... ................... ... .. ........ ..........................26

3-1 Relative abundance of calliphorid adults aerially collected.................................38

3-2 A calliphorid first-instar larva hatching from its egg .............................................39

3-3 The two sets of spiracles of a first-second transitional larva .................................40

3-4 The two inner slits in the spiracles of a second-instar calliphorid larva. .................41

3-5 The two sets of spiracles of a second-third transitional calliphorid larva ..............42

3-6 The three inner splits in the spiracles of a third-instar calliphorid larva .................43

3-7 Chrysomya rufifaces, third-instar larvae ...................................... ............... 44

3-8 Relative abundance of third-instar calliphorid larvae preserved from pig carrion
during the study (N=8253) in Earleton, Florida....................................... .......... 45

3-9 Calliphorid activity for year 1 of the study. .................................. .................46

3-10 Calliphorid activity for year 2 of the study. .................................. .................47

3-11 Mean daily and mean low temperatures (SD) for year 1, November 16, 2001 to
October 26, 2002 (Part A) and year 2, November 30, 2002 to March 14, 2004
(Part B) of study. ........................................... .....................48

3-12 Reared adults, N=232, (Part A), and preserved larvae, N=30, (Part B), from
collection 1, November 16 to November 21, 2001. ........................................ ......49









3-13 Reared adults, N=362, (Part A), and preserved larvae, N=568, (Part B), from
collection 2, December 29, 2001 to January 11, 2002. ..........................................50

3-14 Adults aerially collected, N=39, (Part A), reared adults, N=305, (Part B), and
preserved larvae, N=151, (Part C), from collection 3, February 1-9, 2002. ............51

3-15 Adults collected aerially, N=71, (Part A), reared adults, N=326, (Par B), and
preserved larvae, N=134, (Part C), from collection 4, March 15-19, 2002. ............52

3-16 Adults aerially collected, N=107, (Part A), reared adults, N=332, (Part B), and
preserved larvae, N=196, (Part C), from collection 5, April 29 to May 1, 2002.....53

3-17 Adults aerially collected, N=51, (Part A), reared adults, N=67, (Part B), and
preserved larvae, N=175 (Part C), from collection 6, May 20 to May 23, 2002.....54

3-18 Adults aerially collected, N=99, (Part A), reared adults, N=223, (Part B), and
preserved larvae, N=700, (Part C), from collection 7, July 22 to July 24, 2002......55

3-19 Adults aerially collected, N=76, (Part A), reared adults, N=248, (Part B), and
preserved larvae, N=499, (Part C), from collection 8, August 19-23, 2002............56

3-20 Adults aerially collected, N=76, (Part A), reared adults, N=323, (Part B), and
preserved larvae, N=499, (Part C), from collection 9, September 23-27, 2002.......57

3-21 Adults aerially collected, N=87, (Part A), reared adults, N=131, (Part B), and
preserved larvae, N=294, (Part C), from collection 10, October 26-28. 2002.........58

3-22 Adults aerially collected, N=60, (Part A), reared adults, N= 384, (Part B), and
preserved larvae, N=1248, (Part C), from collection 11, November 30-
D december 8, 2002. .................................... .. ... .... ... ........ .... 59

3-23 Adults aerially collected, N=56, (Part A), reared adults, N= 371, (Part B), and
preserved larvae, N=581, (Part C), from collection 12, December 30, 2002 to
January 11, 2003 .............................................................................. .... 60

3-24 Adults aerially collected, N=66, (Part A), reared adults, N=166, (Part B), and
preserved larvae, N=340, (Part C), from collection 13, March 1-8, 2003. ..............61

3-25 Adults aerially collected, N=247, (Part A), reared adults, N=229, (Part B), and
preserved larvae, N=301, (Part C), from collection 14, April 1-6, 2003 ................62

3-26 Adults aerially collected, N=185, (Part A), reared adults, N=359, (Part B), and
preserved larvae, N=651, (Part C), from collection 15, April 26-May 1, 2003......63

3-27 Adults aerially collected, N=221, (Part A), reared adults, N=150, (Part B), and
preserved larvae, N=388, (Part C), specimens from collection 16, June 12-15,
2003 .......................................................................... .................. ............64









3-28 Adults aerially collected, N=186, (Part A), reared adults, N=266, (Part B), and
preserved larvae, N=648, (Part C), from collection 17, December 8-20, 2003......65

3-29 Adults aerially collected, N=220, (Part A), reared adults, N=294, (Part B), and
preserved larvae, N=347, (Part C), from collection 18, January 23-31, 2004. .......66

3-30 Adults aerially collected, N=316, (Part A), reared adults, N=639, (Part B), and
preserved larvae, N=675, (Part C), from collection 19, March 5-14, 2004. ............67















Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

THE FORENSICALLY IMPORTANT CALLIPHORIDAE (INSECTA: DIPTERA) OF
PIG CARRION IN RURAL NORTH-CENTRAL FLORIDA


By

Susan V. Gruner

August 2004

Chair: John Capinera
Major Department: Entomology and Nematology

The use of insect life stages in the determination of postmortem intervals in crime

scene investigations is an important forensic science tool used by coroners, medical

examiners, and police investigators. For estimation of postmortem interval, basic

distribution data for the major indicator species of insects are required. It is apparent that

the seasonality and species assemblage vary in different geographical areas.

A study to determine possible indicator species of Calliphoridae present in rural

north-central Florida was conducted using pig carrion as models representing human

bodies. A wooded habitat was used as the site for placement of the pigs. The study

involved 19 batches of pigs placed in a wooded site over a period of time including spring,

summer, fall, and winter collections from November 16, 2001, to March 2004

(approximately monthly). Larval and adult calliphorid flies were collected, as were

meteorological data relating to the study site.









Seven species of Calliphoridae were collected from the pig carrion. Relative

abundance of each species as a percentage of the total adult Calliphoridae assemblage (%

aerially collected/% reared) for the study was Phaenicia coeruleiviridis, 68.1 vs. 77.9%;

Cochliomyia macellaria, 16.0 vs. 8.5%; Chrysomya rufifaces, 7.0 vs. 8.0 %; Phormia

regina, 8.2 vs. 3.9%; Chrysomya megacephala, 0.3 vs. 1.6 %; Calliphora livida, 0.4 vs.

0.1%; and Calliphora vicina, 0.0 vs. 0.02%. There were obvious seasonal and successional

variations of the species assemblage. Phaenicia coeruleiviridis (Macquart) was the

predominant species year-round but was lower in abundance during the summer, mid-June

to mid September. Only a few specimens of C. vicina Robineau -Desvoidy (= C.

erythrocephala Meigen) and C. livida Hall were found during the coldest months,

November to February, while C. megacephala (Fabricus) was collected during the hottest

months, June to September. Cochliomyia macellaria (Fabricus) was found during the

warm months, April to June, when the temperature did not rise above 30.00 C. Chrysomya

rufifaces (Macquart) was found in all but the coldest months of the year, mid December to

mid March. Phormia regina (Meigen) was not found during the winter.

Different species of calliphorid flies arrived at the pig carrion at different stages

during the decomposition process. Within minutes of placing the pig carcass on the ground,

P. coeruleiviridis usually began to arrive. Cochliomyia macellaria, C. rufifaces, P. regina,

C. vicina and C. livida arrived at the carcass after a delay of about 24 hours.














CHAPTER 1
INTRODUCTION AND LITERATURE REVIEW

Definition of and Scope of Forensic Entomology

Forensic entomology is an extensive discipline where arthropod science and the

judicial system interact (Hall 2001). The field of forensic entomology has been divided

into three areas: medicocriminal entomology (also referred to as medicolegal

entomology), urban entomology and stored product entomology. Information gained

from medicolegal entomology typically is used to determine time of death, place of death

and other issues of medical or legal importance (Gordh and Headrick 2001). Urban

entomology concentrates mainly on controversies involving termites, cockroaches, and

other insect problems accruing to the human environment, whereas stored product

entomology involves disputes over arthropods and arthropod parts in food and other

products (Hall 2001).

When human remains are found, the most important questions usually are how,

when, where and why the person died. Historically, determination of the postmortem

interval (PMI) has been estimated through observation and measurement of body

conditions such as core body temperature (Nelson 1999), muscular flaccidity, rigor

mortis, lividity, pallor of the skin and others (Smith 1986, Bass 2001, Byrd and Castner

2001 a). Entomological specimens in medicolegal death investigations can be reliable

indicators for estimating the PMI in both early and advanced stages of cadaver

decomposition (Nuorteva 1977, Smith 1986, Goff et al. 1988, Kashyap and Pillay 1989,

Greenberg 1991, Byrd 1998).









Insects and other invertebrates feeding on carrion form a distinct faunal succession

associated with the various stages of decay (Smith 1986). Recognition of the different

immature stages of each species involved, together with the knowledge of their rates of

development, can give an indication of the PMI (Smith 1986). A forensic (= medicolegal)

entomologist can also determine the age of immature insects, based upon knowledge of

the variables regarding insect invasion of human remains. Evaluation and interpretation

of entomological evidence at a crime scene can address other complicated issues

including season of death, geographic location of death, movement or storage of the

remains following death, location of specific sites of trauma on the body, sexual

molestation and use of drugs (Haskell et al. 1997).

In case studies conducted in varying temperate and tropical climates where human

remains were exposed to the environment for 2.5 months or less, entomology-based PMI

estimates differed by + 48 hours when compared with the intervals determined by

independent corroboration such as confessions and eyewitness testimony (Greenberg

1985, Goff et al. 1988, Lord 1990, Byrd 1998). Entomological evidence is statistically the

most reliable scientific means of estimating PMI when compared to other methods such

as police reports and autopsy results (Kashyap and Pillay 1989, Catts and Haskell 1990,

Anderson 2001).

History of Forensic Entomology

The first documented forensic entomology case is from thirteenth century China in

a book entitled "Hsi yuan chi lu" which can be translated as "The Washing Away of

Wrongs." The author, Sung Tz'u, was an educated man. He was a doctor, a sheriff and

eventually a Judicial Intendant. The book describes applications of forensic entomology

used in criminal cases during that period. A man was murdered by the roadside









apparently by an assailant with a sickle. Sung Tz'u made a proclamation that the nearest

neighbors were to bring all their sickles to him for examination (McKnight 1981). At

inquest time, the weather was hot and blow flies were attracted to one sickle only, even

though it had no discernable traces of blood. The owner of the sickle confessed to the

murder.

In addition to medical and legal experts, sculptors, painters and poets have closely

observed the decomposition of human bodies, noting, in particular, the effects of feeding

maggots. Artwork from the Middle Ages accurately depicts the insect-mediated pattern of

body mass reduction, particularly the early skeletonization of the skull and the reduction

of internal organs, with large parts of the skin left intact (Benecke 2001). In May 2004, a

new painting of Prince Philip entitled "Portrait of a Prince" was released by artist Stuart

Pearson Wright. The painting shows Prince Philip with a bluebottle fly sitting on his left

shoulder, which represents a memento mori; the prince's mortality (The Associated Press

2004).

In 1855, Dr. Bergeret, a French physician, used insect succession as a tool

(incorrectly) to solve a case (Benecke 2001). In the mid-1880s, J.P. Megnin, also in

France, published La Faune des Cadavres: Application de Entomologie a la Medicin

Legale. The recognition by Megnin of a sequence and progression of decomposition of a

corpse was recorded in this work and in association with this decomposition progression,

he observed changes in the insect assemblages as the corpse aged (Haskell et al. 1997,

Benecke 1998).

This early interest in insects and decomposition led to a study on insect succession

on human corpses in Quebec, Canada, in 1897 by Wyatt Johnston and Geoffrey









Villeneuve (Anderson 2001, Benecke 2001). At the same time in the United States,

Murray Motter systematically tabulated the insect fauna from 150 exhumed corpses from

the Washington, D.C. area (Haskell et al. 1997, Benecke 2001).

Species identification of the most important fly groups, Calliphoridae (blow flies)

and Sarcophagidae (flesh flies), used in forensic cases could not have been accomplished

had it not been for Aldrich's (1916) monograph on the Sarcophagidae which illustrated

the distinctive male genitalia of adult flies. Knipling (1936) initiated taxonomic work on

the larvae of sarcophagids and calliphorids. Hall's 1948 book, The Blowflies of North

America, made it possible to identify the mature larvae of most species of calliphorids.

In northern Europe, the blow fly Phaenicia sericata (Meigen) is the most

economically important ectoparasite of domesticated sheep. Sheep myiasis is a

widespread disease and can cause high levels of mortality. The desire to develop control

methods against sheep myiasis led to studies of calliphorid attractants (Wardle 1921,

Cragg and Thurston 1949, Hammack and Holt 1983, Ashworth and Wall 1994, Wall and

Warnes 1994, Morris et al. 1998). The attractant studies prompted additional studies on

blow fly distribution and ecology (Parish and Cushing 1938, James 1947, Green 1951,

Wolff et al. 2001) and were followed by studies that addressed effects of temperature on

developmental time of blow fly life cycles (Davidson 1944, Kamal 1958, Nuorteva 1977,

Greenberg 1991, Byrd and Butler 1996, 1997, 1998).

Regional successional studies of Calliphoridae in the United States have been

conducted in California (James 1955), Hawaii (Goff et al. 1986, Goff et al. 1988, Goff

1991), Mississippi (Goddard and Lago 1985), Missouri (Hall and Doisy 1993), Virginia

(Hall and Townsend 1977), Indiana (Haskell 1989), Illinois (Baumgartner 1988), Arizona









(Deonier 1942, Baumgartner 1986, Galloway et al. 1989), Colorado (Adair 1999),

Maryland (Introna et al. 1991), West Virginia (Joy et al. 2002), Louisiana (Tessmer et al.

1995, Watson and Carlton 2003), and South Carolina (Tomberlin and Adler 1998). Four

species--two of which are now found in Florida--of Old World blow flies have been

confirmed from South or North America (Baumgartner and Greenberg 1984,

Baumgartner 1986, Greenberg 1988, Tantawi and Greenberg 1993, Martin et al. 1996):

Chrysomya rufifaces (Maquart), C. albiceps (Wiedemann), C. megacephala (F.) and C.

putoria (Wiedmann). Studies by Byrd (1998) and Peters (2003) were conducted in the

Gainesville, FL (= north-central Florida) area. This study will add rural north-central

Florida to the list.

The study of insects important to forensic entomology has been conducted mainly

through the use of non-human animal models. Decomposition studies worldwide have

used a variety of different carcass types and sizes, including dogs (Jiron and Cartin 1981,

Early and Goff 1986, Richards and Goff 1997), cats (Early and Goff 1986), alligators

(Watson and Carlton 2003), voles (Lane 1975), rats (Greenberg 1990, Tomberlin and

Adler 1998, Faucherre et al. 1999, Kocarek 2001), squirrels (Johnson 1975), deer

(Watson and Carlton 2003), foxes (Easton and Smith 1970, Smith 1975), harbor seals

(Lord and Burger 1984b), herring gulls (Lord and Burger 1984a), guinea pigs

(Bornemissza 1957), mice (Putnam 1978, Blackith and Blackith 1989), lizards and toads

(Cornaby 1974), raccoons (Joy et al. 2002), turtles (Abell et al. 1982), poultry (Hall and

Doisy 1993, Tessmer et al. 1995), sheep (Deonier 1940), rabbits (Denno and Cothran

1975, Tantawi et al. 1996, Bourel et al. 1999), elephants (Coe 1978), opossums (Goddard

and Lago 1985), black bears (Anderson 1998, Peters 2003, Watson and Carlton 2003),









impala (Braack 1981), and pigs (Payne 1965, Tullis and Goff 1987, Haskell 1989,

Anderson and VanLaerhoven 1996, Tessmer and Meek 1996, Richards and Goff 1997,

Byrd 1998, deCarvalho et al. 1999, Shahid et al. 1999, Davis and Goff 2000, deCarvalho

and Linhares 2001, Wolff et al. 2001, Tenorio et al. 2003, Watson and Carlton 2003).

The only faunal succession research on human remains was conducted in Tennessee

(Rodriguez and Bass 1983, Catts and Haskell 1990).

Human cadavers are not easily obtainable for detailed decomposition studies. Pigs,

Sus scrofa, are omnivorous, have similar gut fauna, are relatively hairless and have skin

that is very similar to that of humans (Anderson and VanLaerhoven 1996). The

putrefaction of pigs proceeds approximately at the same rate as for human bodies that are

of the same weight (Campobasso et al. 2001). Haskell's 1989 study in Tennessee

(Schoenly and Haskell 2000) compared the insect community structure and

decomposition rates between adult and infant human remains to a pig model and found

no significant difference in the composition of the insect communities in human and pig

carcasses (Campobasso et al. 2001). Therefore, twenty-two kg pigs have been

recommended as suitable human models for adult decomposition (Catts and Goff 1992).

Current Status of Forensic Entomology

The popularity of television shows such as C.S.I. (Crime Scene Investigation),

Forensic Files, and Court TV have created a recent surge of interest in the forensic

sciences. Several colleges report long waiting lists for forensic science courses, and

dozens of others are developing courses or entire programs in the science of crime

fighting (Lewerenz 2003). Purdue University instituted its first forensic science course in

the fall of 2003, formatted by Neal Haskell and Ralph Williams. Assuming that the

course would not generate much enthusiasm, a 25-student capacity room was assigned









for the class. Once fall registration was completed, the room had to be changed to a

lecture hall to accommodate the 425 students who registered for the class (Haskell 2003,

personal communication).

Lord and Stevenson's 1986 directory (the only one ever published) of forensic

entomologists listed only 62 scientists involved in this field of study; of the 62, only

about a third were linked solely with the "medicolegal" subdiscipline (Catts and Haskell

1990).

In 1996, the American Board of Forensic Entomologists was created. Currently,

there are only 8 members. However, forensic entomologists have no special group within

the American Academy of Forensic Science or the Entomological Society of America.

The European Association for Forensic Entomology was created in May, 2002, at

the First European Forensic Entomology Seminar, held at the headquarters of the

National Gendarmerie in Rosny sous Bois, France. This association was created to

promote forensic entomology in Europe (Hall 2003). In August, 2003, the first annual

meeting of the American Association of Forensic Entomologists was held in Las Vegas,

NV; approximately 45 people attended the meeting. The second annual meeting is

scheduled for July 24-27, 2004 and will be conducted at the University of California-

Davis.

The highly specialized field of forensic entomology has never had a large

following. Several reasons may explain this lack of interest, including:

* It involves having a close relationship with the larval stages of flies, commonly
known as maggots. Most people think that these creatures, along with insects in
general, are disgusting.

* Only a small number of colleges or universities offer a course in the specific field
of forensic entomology, and none offer majors or minors.









* Historically, there has been little opportunity for full-time employment in this field.

Biology of Calliphoridae

Two major groups of insects are predictably attracted to cadavers and provide the

majority of information in forensic investigation; the flies and the beetles (Castner 2001).

This research focused on the Family Calliphoridae, commonly called the blow flies

(blowflies if you live outside the USA), which are the first insects to find and colonize

human corpses. Experimental studies indicate that these flies arrive at carcasses within

minutes of their exposure (Byrd and Castner 200 b, Watson and Carlton 2003).

There are more than 1000 species of blow flies throughout the world; about 90

species exist in North America (Haskell 2003, personal communication). This family

includes the green bottle flies (genus Phaenicia), blue bottle flies (genus Calliphora), the

screwworm flies (genus Cochliomyia) and the black blow flies (tribe Phormiini).

According to (Hall 1948), "to blow" is an ancient term that refers to depositing of eggs.

The family name means 'beauty bearer' in Greek (Greenberg and Kunich 2002).

Common blow flies carry at least 2,000,000 bacteria per specimen externally.

Internally, each individual fly can carry from eight to ten times as many (Hall 1948).

They can carry typhoid, cholera, the plague, anthrax, tuberculosis, tularemia,

trypanosomes, leishmanias and can cause primary, secondary and tertiary myiasis

(infestations of human skin).

Some species of blow flies, such as Phaenicia sericata, are used for cleaning of

non-healing wounds. This healing method is called "maggot therapy" and has been in use

for hundreds of years (Sherman and Pechter 1988). Sterile maggots debride the wounds

by eating necrotic tissue. Urea, ammonium carbonate and allantoin secreted by the larvae

disinfect deep tissue wounds and healing is stimulated. Maggot therapy is appropriate for









cases where antibiotics are ineffective and surgery is impracticable (Sherman and Pechter

1988).

Calliphorid flies have highly specialized sense organs on their antennae that are

stimulated by putrefaction odors and gases which are released during post-mortem

decomposition of organic matter. Some species of Phaenicia are attracted to various

organic sulphur compounds, either alone or in combination with hydrogen sulphide, and

also by ammonia (Cragg 1956, Cragg and Cole 1956, Ashworth and Wall 1994, Wall and

Warnes 1994). Flight traps baited with dimethyl trisulphide are strong attractants for

some calliphorids (Nilssen et al. 1996). Odors from Proteus mirabilis Hauser, a

bacterium that causes infections in the fleece of sheep, are attractants to some calliphorid

flies (Morris et al. 1998).

Some plants have developed a pollination strategy that targets calliphorid flies. The

dead-horse arum (Helicodiceros muscivorus L. fil.) floret emits an odor that smells like a

dead animal. Blow flies are deceived into pollinating the plant. The volatile compounds

of the dead-horse arum and of a carcass were identified by gas chromatography as three

structurally similar oligosulphides: dimethyl mono-, di- and trisulphide. When calliphorid

flies were exposed to each of the odors, identical antennal response patterns were elicited

(Stensmyr et al. 2002).

Landing behavior of calliphorids may be affected by visual cues such as white and

yellow colors (Wall et al. 1992, Hall et al. 1995). Oviposition is elicited primarily by the

presence of ammonia-rich compounds, moisture, pheromones, and tactile stimuli

(Ashworth and Wall 1994) yet is rarely stimulated by chemicals alone (Cragg 1956).









Unfortunately, the complex interaction of semiochemical and visual cues used for

resource location remains little studied in calliphorids (Wall and Fisher 2001).

Blow flies are heliotropic and usually rest at night. Eggs are not usually laid at

night although clearly there are exceptions. Green (1951, page 484) observed that

Calliphora deposited eggs at night under artificial light in slaughter houses. He wrote that

"under laboratory conditions it has been found that Calliphora erythrocephala (now

called C. vicina), Lucilia sericata and Phormia terrae-novae will all oviposit in total

darkness, although Wardle (1921) asserts that blowflies do not oviposit in the complete

absence of light. Greenberg (1990) observed that Phaenicia sericata, Phormia regina

(Meigen) and Calliphora vicina (Robineau-Desvoidy) oviposited a very small number of

eggs on rat carrion at night. Singh (2001) pointed out that the flies in Greenberg's

experiment probably were resting on a nearby bush and literally crawled over to oviposit

on the rat carrion, thus indicating that blow flies were not actively searching for an

oviposition site. Nocturnal oviposition has not been observed in large-scale studies in

other areas (Greenberg 1990, Byrd and Butler 1997, Haskell et al. 1997).

Other factors that affect blow fly activity are temperature, size of the carcass,

geographical location, humidity, light and shade, seasonal and daily periodicity,

availability of food and competition, maggot mass temperature and manner of death

(Rodriguez and Bass 1983).

Insect Succession on Carrion

The first organisms to arrive on a body after death are usually the insects. They

arrive at predictable times during the decomposition process. Each decomposition stage is

attractive to a different group of sarcosaprophagous arthropods. Smith (1986, page 13), in









A Manual ofForensic Entomology, defines four ecological categories in the carrion

community:

1. Necrophagous species: Feed on the carrion itself and constitute the most important
category in establishing time of death, e.g., Diptera:
Calliphoridae (blow flies); Coleoptera (beetles): Silphidae
(in part), Dermestidae.

2. Predators and parasites: Second most important forensic category, e.g., Coleoptera:
Silphidae (in part), Staphylinidae; Diptera: some carrion
feeders become predaceous in later instars, e.g., Chrysomya
(Calliphoridae), Ophyra and Hydrotaea (Muscidae) on the
necrophagous species.

3. Omnivorous species: Wasps, ants and some beetles feed both on the corpse
and its inhabitants.

4. Adventive species Use the corpse as an extension of their environment, e.g.
Collembola springtailss), spiders (which may become
incidental predators).

Blow flies are attracted by the odors and gases which are released during the onset

of autolysis and putrefaction, depending on time of year and situation of the corpse

(Smith 1986). As decomposition proceeds, the odors emanating from the corpse change,

making the cadaver more attractive to some species of blow flies and less attractive to

others. Once the dry decay stage has been reached, blow flies are no longer attracted to

the corpse (Nuorteva 1977, Anderson 2001). After the invasion of North America by the

Chrysomyinae in the 1980's, the blow fly sequence in North America may be Phaenicia,

Cynomyopsis, Chrysomyinae, Calliphora, and Cochliomyia (Campobasso et al. 2001).

Obviously, species lists will differ by region.

Beginning with Megnin's (1894) work, eight waves of arthropod invasion on

human bodies have been described. Other forensic entomologists reduced the number of

stages in attempts to define biological communities, but ultimately this reduction

complicated and lessened the forensic applicability. Payne (1965) defined the associated









insect community and analyzed the percentage abundance of species attracted to the

various stages of decay. He condensed eight to six stages of decay: fresh, bloated, active,

advanced, dry and remains. Lord and Burger (1984a) and Bornemissza (1957) recognized

five stages of carcass decomposition. According to Smith (1986, page 17), there exists a

broad general agreement in the observations of Megnin, Bornemissza (1957), Reed

(1958) and the series of publications by Payne (1965) and (Payne 1965, Payne and King

1970) as follows:

1. Initial decay stage (0-2 days). Carcass appears fresh externally but is decomposing
internally due to the activities of bacteria, protozoa and nematodes present in the
animal before death.

2. Black putrefaction stage (12-20 days). Flesh of creamy consistency with exposed
parts black. Body collapses as gases escape. Odor of decay very strong.

3. Butyric fermentation stage (20-40 days). Carcass drying out. Some flesh remains at
first and cheesy odor develops. Ventral surface of body moldy from fermentation.

4. Dry decay stage (40-50 days). Carcass almost dry; slow rate of decay.

The aforementioned stages of decay are not easy to delineate and there is

controversy regarding these definitions, but they are useful in describing the sequence of

decomposition. For example, the number of days varies considerably with temperature.

Factors that Affect Blow Fly Succession on Carrion

Temperature and access to a body are the two most important factors affecting

insect succession. Temperature is the most important variable influencing the rate of

maggot development. High temperatures generally reduce development time of Diptera.

Large aggregations of dipteran larvae (maggot masses) develop heat due to their frenetic

activity and fast metabolism, thus raising the microenvironmental temperature

(Campobasso et al. 2001). The heat of the maggot mass is related to the density of the

mass and the size of the carcass (weight and mass). The size of maggot masses and the









degree to which the corpse is either exposed to, or insulated from, the environment

affects the amount of heat absorbed or dissipated, which in turn has a significant effect

upon the rate of larval development and the decomposition of a corpse. Goodbrod and

Goff (1990) studied the effects of maggot-generated heat during the development cycle in

experimental cultures of C. megacephala and C. rufifaces and found an inverse

relationship between density and the duration of larval stage.

The insects that colonize corpses vary in species depending on the biogeoclimatic

zone in which the remains are found. Each zone has different habitat types, vegetation,

soil pH, soil type, flora and fauna, altitude and climatic conditions that affect the species

of insects present. Decomposition also is affected by the time of year, and the location in

which remains are found (Anderson 2001).

Many blow fly species vary in abundance depending on season and even time of

day. Presence or absence of sunlight or shade can have an effect on which blow fly

species will colonize a corpse. Cragg (1956) demonstrated that P. sericata prefer heated

surfaces and will not oviposit on carcasses that have surface temperatures below 300 C.

Results of a sun-exposed versus shaded pig carrion study indicated that more Lucilia

illustris (Meigen) and Phormia regina (Meigen) were observed at the sun-exposed pig

whereas Calliphora vomitoria (Linnaeus) were observed in greater numbers at the shaded

pig (Shean et al. 1993).

Blow flies can be found in both urban and rural areas but some species may be

found only in wooded areas. Flies primarily associated with human refuse (synanthropy)

are usually found in urban areas. Presence of certain species of blow flies found on a

body may indicate that the body was moved from an urban to a rural environment or vice









versa (Erzinclioglu 1985, Catts and Haskell 1990). Blow flies are capable of colonizing

corpses inside dwellings and cars, depending upon how well they are sealed.

Competition for the food source is the most important factor affecting size and

completion of the life history of carrion. Smith (1986, page 34) listed ways in which this

occurs:

1. Intraspecific competition may reduce the size of the larvae and can reduce the
number and fecundity of individual adults

2. Interspecific competition has similar results as to intraspecific competition, plus a
possibility of total elimination

3. Predators and parasites-selective predation or parasitization of one species can be
advantageous to a competing species.

The early arrivers (such as Calliphora and Phaenicia) at a corpse may have an

advantage. Some female blow flies (Tribe Calliphorini) are capable of moving a single

egg from one of the ovaries into the vagina (termed "precocious egg development")

where it is fertilized before an oviposition site has been found (Wells and King 2001).

Delayed arrivers such as Chrysomya may compensate for this by being viviparous and

deposit larvae on the carcass. Some dipteran larvae begin life as carrion feeders and

become predators after the second molt.

Biology of Human Decomposition

Campobasso et al. (2001, page 18) describe the decomposition of corpses as

a mixed process ranging from autolysis of individual cells by internal chemical
breakdown, to tissue autolysis from liberated enzymes and from external processes
introduced by bacteria and fungi from both the intestine and outer environment.
The bacterial enzymatic structures cause putrid liquefaction of tissues by the
breakdown of proteins, carbohydrates and lipids into their basic components
(amino acids, water and carbon dioxide, fat acids and volatile substances) with gas
formation (nitrogen, methane, hydrogen sulphide, ammonia, etc.). As the tissues
are digested to a fluid consistency with the production of large amounts of foul-
smelling gas; they become moist and gas-ridden, and eventually liquefy down to
the skeleton.










Calliphorid larvae hasten tissue decay by dissemination of bacteria. The larvae also

have digestive enzymes in their saliva that liquefy carcass tissues. As the larvae burrow

into and consume the carcass, tissues further disintegrate (Easton and Smith 1970).

Usually, tissues that conduct the highest rates of ATP synthesis, biosynthesis, and

membrane transport decompose first. The intestines begin to decompose first, followed

by the stomach, accessory organs of digestion, heart, blood and circulation, heart muscle,

air passages and lungs, kidneys and bladder, brain and nervous tissues, skeletal muscles

and finally, connective tissues and integument (Gill-King 1997).

Over a seven-year period, researchers at the Anthropology Research Facility

(ARF-also known as The Body Farm) in east Tennessee studied 150 bodies in various

stages of decay, including homicide victims, bodies donated to science, and unidentified

persons. They found that variables affecting decay rates of human bodies are (in order of

importance): temperature, access to the body by insects, burial and depth, carnivore and

rodent activity, trauma (penetrating/crushing), humidity, rainfall, size and weight of the

body, embalming, clothing, type of surface body was placed on and the soil pH (Mann et

al. 1990).

Further Areas for Study

Entomological studies of Calliphoridae associated with carrion should include

investigations of seasonal changes in the fauna and successional events which are

influenced by weather factors. Data are needed from different geographic regions

including species composition of sarcosaprophagous arthropods and the weather

parameters associated with those fauna (including their maximum/minimum threshold









temperature data). Reliable taxonomic keys are needed for identification of early larval

instars and of puparia.

The assessment of species composition within rural north-central Florida is

addressed in the present study by determining the calliphorid species associated with

carrion in north-central Florida and the influence of seasons on the calliphorid species

assemblage. The objectives of the study were:

1. To identify the species of Calliphoridae associated with pig carrion in rural north-
central Florida.

2. To determine how season (spring, summer, fall and winter) affects the assemblage
of calliphorid species associated with pig carrion in rural north-central Florida.

3. To determine the daily succession of calliphorid species associated with pig
carrion during each collection period in rural north-central Florida.














CHAPTER 2
MATERIALS AND METHODS

The use of human corpses for field studies is illegal in Florida. As determined in

other decomposition studies (Haskell 1989, Anderson and VanLaerhoven 1996,

Campobasso et al. 2001) the rate of pig decomposition is very similar to that of humans.

Dead pigs, Sus scrofa L., were used as animal models for this study.

All pigs were purchased from North Florida Livestock Market, Lake City, Florida.

Prior to purchase, the pigs were killed by a lateral, transverse shot into the top of the head

with a .22 caliber rifle. This method resulted in instant death of the animal. Each dead pig

was immediately double-bagged in a heavy-duty plastic trash bag and was transported

from Lake City to the Greathouse Butterfly Farm in Earleton, Florida.

Study Site

Greathouse Butterfly Farm is located 19.3 km east of Gainesville, near Earleton, on

the southeast corner of N.E. SR 26 and SR1469. The property consists of 48.6 hectares of

north Florida flatwoods (Figure 2-1). Pig carcasses were placed in a wooded habitat

where sunlight was somewhat restricted. In some cases, the pigs received direct sunlight

during certain parts of the day, while at other times they were shaded. The study site

mainly consisted of a moderate stand of live oak, Quercus virginiana Mill., and slash

pine, Pinus elliottii Engelman, with an understory of various saw palmettos and grasses

(Florida Chapter 1989).

Pigs were placed at least 18.3 meters apart. Wire cages (86.5 cm long x 50.8 cm

wide x 61 cm high) were placed over the pigs to protect them from large vertebrate









scavengers. The cages were constructed of wire mesh (5 cm x 5 cm). At least four metal

or plastic tent stakes were driven into the ground around the cage, and bungee cords were

attached tightly to the cages and then to tent stakes to minimize disturbance (Figure 2-2).

The cages were lifted off and set aside during sampling times.

Data Collection

Trials were conducted from October 14, 2001 to March 5, 2004; however, data

from October 2001 were incomplete and therefore not included. The raw data, starting

with the second collection on November, 2001, are in appendices A and B. Each trial

consisted of 3-4 pigs, with trials conducted throughout the year (Table 2-1). Observations

and collections were made daily during the afternoon, if possible, when flies were most

active. The duration of each trial lasted until the first wave of maggot dispersal occurred

(termed "maggot migration" by forensic entomologists). After the third-instar larvae have

reached a certain size, they leave the carcass en mass to pupariate in the soil. The pigs

were not moved or disturbed in any way during the study. Digital pictures of the pigs and

of the expanding maggot masses were taken at each collection time. At the beginning of

each collection time, a laminated identification sheet was placed in front of each pig,

indicating the date, time and identification number. A meter-stick was placed below the

identification sheet to give document size. Sampling usually occurred in the middle of the

afternoon when the adult flies were most active.

Protocol for Day 1

On day 1, the pigs were removed from bags and placed in a location on the site, and

the GPS (global positioning system) was noted. After all the pigs were in position, data

were collected including pig number, time of death (TOD), date, time, sample number

(SA#), ambient temperature (AT0), ground/pig interface temperature (GPI), ground









temperature at 5 cm depth about 3 m from the pig, a brief description of the weather

(sunny, cloudy, rain, etc.) and wind velocity in meters per second (M/S).

An aerial collection of adult flies was made over each pig with an insect net, with a

target of at least 10 adult calliphorid flies (although this was not always possible to

achieve). Flies were placed in vials with 70% isopropyl alcohol and later pinned and

identified. A data logger (Figure 2-3) was affixed to a nearby tree or bush and the

temperature probe was placed on the ground 1-2 m from each pig. Therefore, the ground

temperature being recorded was similar to that which the pig was exposed.

Protocol for Day 2 Onward

Adult flies were sampled with a net (if possible) and paired samples of

approximately 50-500 larvae (L = live, P = preserved) were collected from the growing

maggot mass, usually located on the head. The location where the sample was taken on

the body area (or ground area) was noted on the collection sheet. About half of the

sample specimens were boiled in water for about 2 minutes using a camp stove, and then

placed in vials with 70% isopropyl alcohol for preservation. The remainder of larvae

were placed in containers and reared to the adult stage in order to compare the identity

and relative abundance of adult and larval flies.

Meteorological Measurements

Temperatures determined with a Taylor 9841 digital thermometer (Forestry

Suppliers, Jackson, MS) shielded from direct rays of the sun (when necessary) include

ambient air, ground/pig interface, external tissue, oral cavity of the pig, soil at 5 cm depth

3 m from the pig, and soil at 5 cm depth under the maggot mass. Maggot mass

temperatures were taken (MM T) but will not be addressed in this thesis. Ground

temperatures were taken with HOBO (Onset Computer Corp., Bourne, MA) data









loggers. These units were not 100% waterproof, so a plastic housing was made for each

of the loggers (Figure 2-3). Three hangars were cut and cemented with marine glue into

one end of each container. A hole was drilled at the bottom of the container (for the

temperature probe cable) and foam was packed around each logger. The HOBO unit

was set to record temperature every 30 minutes for 3 weeks. A temperature probe

connected to each logger was set on the ground within 1-2 m of each pig (Figure 2-4).

Table 2-4 lists the mean temperatures that were taken during the collection intervals

during the months of the project. The mean temperatures were taken with the Taylor

thermometer during the collection times. The temperature at 3:56 AM, taken from the

data logger's data, was used as the mean low temperature (Figures 3-5 A and B). There

was no solar interference at this time. For collections 2 and 3, the data logger was not

used and the mean low temperatures were taken from the NOAA (National Oceanic and

Atmospheric Administration) daily data available for Gainesville Regional Airport.

Rearing Procedures

Maggots were removed from the carcass with a plastic spoon and placed

immediately into rearing pouches constructed from aluminum foil (Catts and Haskell

1990) and placed into Rubbermaid 28 L or plastic Gladware containers half-filled with

substrate for pupation. The container lids each had approximately ten 2-mm holes

randomly drilled for air circulation. A paper towel was placed under each lid to prevent

larvae/adult flies from escaping through lids and also to prevent other flies from getting

into the containers.

Each foil pouch was filed with 100-200 g of calves' liver, which served as food.

The containers of live maggots were kept outside-except when temperatures were

below 4.5C-on shelving placed in a screened enclosure. The maggots were checked









daily. Once the maggots migrated off the liver, the pouch was removed from the

container. On occasion, the maggots would not move out of the foil pouch. When this

occurred, a small amount of fresh liver was placed directly onto the substrate, and the

maggots were gently moved from the foil pouch onto the liver. The liver would be

removed a day or two later after the maggots migrated into the substrate to pupate.

At adult emergence, the containers were placed into the freezer for 15 minutes to

kill the flies. Adults were put in labeled vials of 70% isopropyl alcohol, then later pinned

and identified (White et al. 1940, Dodge 1953, Seago 1953, Furman and Catts 1982,

Wells et al. 1999). The preserved maggots were separated into first, first-second

transitional, second, second-third transitional and third instars. Only third-instar larvae

were identified to species because there are no reliable taxonomic keys for first- and

second-instar calliphorid larval identification. Gary Steck, dipterist at Florida Division of

Plant Industry (DPI, Department of Agriculture and Consumer Services, Gainesville, FL),

verified the identifications of representative samples of the flies. All data were entered

into a Microsoft Office XP Excel version 2002 spreadsheet (Microsoft Corp.). The

percentages of adult and larval specimens were transformed to their square roots to run

the Pearson's correlation analysis. The raw data are presented in Appendices 1 and 2.

Voucher specimens are being kept in lab 2209 at the University of Florida Department of

Entomology and Nematology and will be deposited with the museum at DPI.

Pupation Substrate

Pupation substrate consisted of about 90% dried laurel oak leaves (Quercus

laurifolia Michx.) and branches, 2 % rye grass (Lolium perenne Lam.) cuttings, 8 %

magnolia leaves (Magnolia grandiflora L.), and a minute amount of sandy soil. The plant

mixture was ground up with a Troy-Bilt Model 4731-10 HP chipper/shredder with a






22


mulching blade. The substrate was stored in a 200 L (liter) Rubbermaid trash container

with a tight fitting top. On most occasions, larvae that migrated into the substrate

successfully produced adults in containers half-filled with substrate. On a few occasions

(less than 3), no adults were obtained due to mortality from fungi. No information is

available; however, on the proportion of pupae that successfully reached adulthood.


















































Figure 2-1. A caged pig carcass is shown at Greathouse Butterfly Farm property in
Earleton, Florida.






























Figure 2-2. Wire cage with bungee cords and tent stakes hammered into the ground. The
cage protected the carcass from scavengers.







































Figure 2-3. A HOBO temperature data logger is sealed inside a Gladware container.
Data loggers (one for each pig) were used to obtain ground temperatures near
pig carcasses at Greathouse Butterfly Farm, Earleton, Florida.
































S*WFWiff a&qC7 !I ME.F il'EM12iMillP=C lll GOza 7 ='IM&
Figure 2-4. The data logger was hung from bush at left and temperature probe was placed
on the ground (circled) to record ground temperature near the pig carcass in
Earleton, Florida.











Table 2-1. Dates of pig deposition, approximate pig weight, and time each pig was
placed on site for sampling dates from November, 2001 to October, 2002.

Date of Pig Deposition Approx. pig weight (kg) Number of pigs per date Approx. deposition time (h)
15-Nov-2001 20 3 1730
23-Dec 20 3 1500
1-Feb-2002 25 4 1530
15-Mar 24 4 1545
29-Apr 30 4 1445
20-May 32 4 1630
22-Jul 20 4 1550


19-Aug
23-Sep
24-Oct
27-Nov
30-Dec
28-Feb-2003
31-Mar
25-Apr
12-Jun
8-Dec
23-Jan-2004
5-Mar


1445
1450
1730
1500
1530
1545
1445
1630
1550
1440
1545
1530











Table 2-2. Mean temperatures ( SD) during the study at the Earleton, Florida site. Mean
temperatures were taken during collection times with the Taylor digital
thermometer while facing away from sun. Mean low temperatures were taken
from the HOBO 9 data at the 3:56 AM mark. November 2001 through
January 2002 temperature data were obtained from National Oceanic and
Atmospheric Administration.


Nov. 16-21, 2001
Dec. 29, 2001-Jan.11, 2002
February 5-9, 2002
March 15-19, 2002
April 29 -May 1, 2002
May 20-23, 2002
July 22-25, 2002
Aug. 19-23, 2002
September 23-27, 2002
October 26-28, 2002


Mean Temp. 0 C
23.6
20.2
19.9
30.4
32.8
25.9
33.2
30.5
29.3
28.4


SD
1.1
3.2
4.1
1.2
1.1
0.9
1.3
2.9
3.2
2.2


Mean Low Temp.0 C
5.2
0.9
8.1
16.5
19.4
17.4
22.4
23.9
23.7
19.1


Nov. 30-December 14, 2002
December 30, 2002-January 11, 2003
March 2-8, 2003
April 1-6, 2003
April 26-May 1, 2003
June 12-15, 2003
Dec. 8-20, 2003
January 23-31, 2004
March 5-14, 2004


SD
2.8
5.8
4.5
3.0
1.5
3.2
0.7
1.3
1.7
0.5


16.8
17.0
22.1
25.0
27.4
30.3
17.4
18.9
24.5


9.7
5.7
16.9
10.8
18.4
23.0
9.4
8.9
13.3














CHAPTER 3
RESULTS

Species of Calliphoridae Collected on Pig Carrion

The seven species of Calliphoridae collected on pig carrion between November 16,

2001 and March 14, 2004 in rural north-central Florida were Phaenicia coeruleiviridis

(Macquart), Cochliomyia macellaria (Fabricus), Chrysomya rufifaces (Macquart),

Phormia regina (Meigen), Chrysomya megacephala (Fabricus), Calliphora livida Hall,

and Calliphora vicina Robineau-Desvoidy (= C. erythrocephala Meigen) (Figure 3-1).

Assessment of the fly assemblage by three collection methods (aerial collection of

adults, adults reared from larvae, and preserved larvae) produced similar results (Figure

3-1). Pearson's correlation analysis of aerial collections made when adult flies were most

active (usually during the early afternoon) indicated a high degree of correlation with

maggot abundance for the same time period. The analysis of aerial and reared specimens

indicated a high degree of correlation (r = 0.9512, 95% confidence interval = 0.6981 to

0.9930, two-tailed P= 0.0010). The analysis of aerial and preserved specimens (r =

0.9744, 95 % confidence interval 0.8315 to 0.9964, two-tailed P = 0.0002) also indicated

a high degree of correlation. Finally, the analysis of reared and preserved specimens (r =

0.9783, 95% confidence interval 0.8557 to 0.9969, two-tailed P = 0.0001) indicated a

high degree of correlation. It is much easier to catch adult flies than it is to rear larvae, so

an aerial sample taken during the middle of the day represents a good estimation of

species present, although, in practice, both adults and larvae should continue to be

collected for legal reasons.









Preserved larvae collected during the study were identified and counted by instar.

Examined through a microscope, each of the different instars is easily recognizable. First-

instar larvae have Y-V shaped spiracular slits (Figures 3-2 A and B). First-second

transitional instars have a second set of spiracles behind the Y-V slits (Figure 3-3). The

second-instar larvae have two inner spiracular slits (Figure 3-4). The second-third

transitional instars have two sets of spiracular slits (Figure 3-5), and the third instars have

three inner spiracular slits (Figure 3-6). Chrysomya rufifaces larvae are recognizable

without the aid of a microscope; they have rows of conspicuous tubercles (Figure 3-7).

Chrysomya megacephala larvae have a distinguishing accessory oral sclerite, but none

were collected in this study. A total of 23,960 larvae were collected. Of these, 8253 were

third-instar larvae and their relative abundance was: P. coeruleiviridis, 76.5%; C.

macellaria, 7.3%; C. rufifaces, 9.1%; P. regina, 6.8%; and 0.3% were unknown (Figures

3-1 and 3-8).

Seasonal Distribution and Succession of Calliphoridae Species

Adult calliphorids collected and reared in year 1 (N=3197) and year 2 (N= 3992)

are shown in Figures 3.9 and 3.10, respectively. Differences in seasonal phenology are

evident (spring is March 20 to June 20; summer is June 21 to September 21; fall is

September 22 to December 20; winter is December 21 to March 19). The mean

temperatures taken during afternoon samples and the mean low temperatures for year 1

and year 2 of the study are in Figures 3-11 A and B. Two species, Calliphora livida and

Calliphora vicina, were found only during the winter, from mid-December to mid-March

(Figs. 3-9 and 3-10). One species, Chrysomya megacephala, was found only in the

summer, from mid-June to late-September (Figs. 3-9 and 3-10). Cochliomyia macellaria,

C. rufifaces and P. regina and others were not found during the winter (Figs. 3-9 and 3-









10). Phaenicia coeruleiviridis generally were found in great abundance year round, but

were lower in abundance during the summer (Figures 3-9 and 3-10).

Different species of calliphorid flies arrived at the pig carrion at different stages

during the decomposition process. Within minutes of removing the pig from the plastic

bag and placing it on the ground P. coeruleiviridis usually began to arrive. Cochliomyia

macellaria, C. rufifaces, P. regina, C. livida and C. vicina arrived at the carcass after a

delay of about 24 hours.

The data below are discussed in terms of total insects collected, reared, or

preserved per collection interval. The figures are shown graphically in an effort to

evaluate succession.

Collection 1, November 16-21, 2001

No adult calliphorids were aerially collected. Of the total reared adults (N=232),

94% were P. coeruleiviridis, 5.6% C. rufifaces and 0.4% were C. macellaria (Figure 3-12

A). One hundred percent of the preserved third-instar larvae (N=30) were P.

coeruleiviridis (Figure 3-12 B).

Collection 2, December 29, 2001-January 11, 2002

No adult calliphorids were aerially collected. Of the total reared adults (N=362),

99.4 % were P. coeruleiviridis, while C. livida and C. vicina each comprised of 0.3%

(Figure 3-13 A). Preserved larvae (N=568) consisted of 79.8% P. coeruleiviridis, 18.0%

P. regina, and 2.3% were unknown (Figure 3-13 B).

Collection 3, February 5-9, 2002

One hundred percent of the adults aerially collected (N=39) and adults reared

(N=305) were P. coeruleiviridis (Figures 3-14 A and B). Preserved larvae (N=151)

consisted of 70.9% P. coeruleiviridis and 29.1% P. regina (Figure 3-14 C).









Collection 4, March 15-19, 2002

Adults aerially collected (N=71) consisted of 97.2% P. coeruleiviridis and 2.8% P.

regina (Figure 3-15 A). Of the reared adults (N=326), 100% were P. coeruleiviridis

(Figure 3-15 B). Preserved larvae (N=134) consisted of 97.8% P. coeruleiviridis and

2.2% were unknown (Figures 3-15 C).

Collection 5, April 29-May 1, 2002

Adults aerially collected (N=107) consisted of 86.0% P. coeruleiviridis, 0.9% C.

macellaria and 13.1% P. regina (Figure 3-16 A). Of the reared adults (N=332), 90.1%

were P. coeruleiviridis, 2.7% were C. macellaria and 7.2% were P. regina (Figure 3-16

B). Preserved larvae (N=196) consisted of 100% P. coeruleiviridis (Figure 3-16 C).

Collection 6, May 20-23, 2002

Figure 3-17 A illustrates successional data. Four species of calliphorids were

captured on the first day. Cochliomyia macellaria was the most abundant species, but

apparently they were not depositing eggs on the first day because the larvae collected on

day 3 were 100 % P. coeruleiviridis. The abundance of P. coeruleiviridis declined (adults

and larvae) as decomposition progressed. The abundance of C. rufifaces remained about

equal, but P. regina increased in abundance as decomposition progressed. Aerially

collected specimens (N=51) consisted of 17.6% P. coeruleiviridis, 39.2% C. macellaria,

21.6% C. rufifaces and 21.6% P. regina (Figure 3-17 A). Of the reared adults (N=67)

and preserved larvae (N=175), 100% were P. coeruleiviridis (Figures 3-17 B and C).

Collection 7, July 22-25, 2002

Figures 3-18 A-C illustrate successional data. P. coeruleiviridis was the most

abundant calliphorid present on day 1 and declined in abundance as decomposition

progressed while C. macellaria, C. rufifaces, and P. regina arrived in fewer numbers on









day 1 but increased in abundance as decomposition progressed. Chrysomya megacephala

arrived when the carcass was in the active decay state on day 3. Adults aerially collected

(N=99) consisted of 58.6% P. coeruleiviridis, 5.1% C. macellaria, 31.3% C. rufifaces,

2.0% P. regina, and 3.0% C. megacephala (Figure 3-18 A). Of the reared adults

(N=223), 47.1% were P. coeruleiviridis and 52.9% were C. rufifaces (Figure 3-18 B).

Preserved larvae (N=700) consisted of 31.6% P. coeruleiviridis and 68.4% C. rufifaces

(Figure 3-18 C).

Collection 8, August 19-23, 2002

Figures 3-19 A-C illustrate successional data. Phaenicia coeruleiviridis was the

most abundant species for the first 2 days but decreased in abundance as decomposition

progressed. Chrysomya rufifaces and C. macellaria did not arrive at the carcass until day

2 (Figure 3-19 A), but increased in abundance as decomposition progressed. Chrysomya

megacephala arrived when the carcass was in the active decay state, after 3 days. Adults

aerially collected (N= 18) consisted of 67.8% P. coeruleiviridis, 6.8% C. macellaria,

23.7% C. rufifaces, and 1.7% C. megacephala (Figure 3-19 A). Of the reared adults

(N=248), 43.1% were P. coeruleiviridis, 1.2% C. macellaria, 42.3% C. rufifaces, and

13.3% C. megacephala (Figure 3-19 B). Preserved larvae (N=499) consisted of 83.0% P.

coeruleiviridis, 0.4% C. macellaria, 13.6% C. rufifaces, 2.8% P. regina, and 0.2% were

unknown (Figure 3-19 C).

Collection 9, September 23-27, 2002

Figures 3-20 A-C illustrate the successional data. Phaenicia coeruleiviridis was the

only calliphorids species collected on day 1 and abundance declined as decomposition

progressed. Cochliomyia macellaria and C. rufifaces arrived at the carcass on day 2, and

increased in abundance as decomposition progressed. Chrysomya megacephala adults









were reared from the September 26 larval sample, but no adults were aerially collected

(Figure 3-20 B). Adults aerially collected (N=76) consisted of 44.7% P. coeruleiviridis,

34.2% C. macellaria, and 21.1% C. rufifaces (Figure 3-20 A). Of the reared adults

(N=323), 54.2% were Phaenicia coeruleiviridis, 30.7% C. macellaria, 1.2% C. rufifaces,

and 13.0% C. megacephala (Figure 3-20 B). Preserved larvae (N=697) consisted of

66.1% P. coeruleiviridis, 21.5% C. macellaria, 11.3% C. rufifaces, and 1.0% P. regina

(Figure 3-20 C). No P. regina adults were aerially collected or reared (Figure 3-20 B).

Collection 10, October 26-28, 2002

Figures 3-21 A-C illustrate successional data. Phaenicia coeruleiviridis, C.

macellaria, and C. rufifaces were collected on day 1. Phaenicia coeruleiviridis was not

the most abundant species of adults on day 1, but was the most dominant species of

larvae. Their abundance declined as decomposition progressed. Adults aerially collected

(N=87) consisted of 12.6% P. coeruleiviridis, 35.6% C. macellaria and 51.7% C.

rufifaces (Figure 3-21 A) Of the reared adults (N=131), 52.7% were P. coeruleiviridis,

7.6% C. macellaria and 39.7% C. rufifaces (Figure 3-15 B). Preserved larvae (N=294)

consisted of 59.2% P. coeruleiviridis, 8.8% C. macellaria and 32.0% C. rufifaces (Figure

3-21 C).

Collection 11, November 30-December 14, 2002

Phaenicia coeruleiviridis was the most dominant calliphorid species during this

sampling period. Adults aerially collected specimens (N=60) consisted of 91.7% P.

coeruleiviridis, 1.7% C. macellaria, 5.0% C. rufifaces, and 1.7% P. regina (Figure 3-22

A). Reared adults (N=351) consisted of 88.0% P. coeruleiviridis and 12.0% P.

coeruleiviridis, 19.0% C. macellaria (Figure 3-22 B). Preserved specimens (N=1248)

consisted of 99.8% P. coeruleiviridis and 0.2% C. macellaria (Figure 3-22 C).









Collection 12, December 30, 2002-January 11, 2003

Phaenicia coeruleiviridis was the most dominant calliphorid species during this

sampling period. Adults aerially collected (N=56) consisted of 98.2% P. coeruleiviridis

and 1.8 % C. livida (Figure 3-23 A). Reared adults (N=371) consisted of 99.2% P.

coeruleiviridis and 0.8% C. livida (Figure 3-23 B). Preserved larvae (N=1248) were

comprised of 99.8% P. coeruleiviridis, 0.2% C. macellaria and 1.0% were unknown

(Figure 3-23 C).

Collection 13, March 2-8, 2003

Phaenicia coeruleiviridis was the most dominant calliphorid species during this

sampling period. Adults aerially collected (N=66) consisted of 87.9% P. coeruleiviridis,

9.1% P. regina and 3.0% C. livida (Figure 3-24 A). Reared adults (N=166) consisted of

98.2% P. coeruleiviridis and 1.8% P. regina (Figure 3-24 B). Preserved larvae (N=340)

consisted of 71.5% P. coeruleiviridis, 28.2% P. regina and 0.3% were unknown (Figure

3-24 C).

Collection 14, April 1-6, 2003

Figures 3-25 A-C illustrate successional data. Phaenicia coeruleiviridis was the

most dominant calliphorid species for the first 4 days of the sample period but decreased

in abundance as decomposition progressed. Cochliomyia macellaria, C. rufifaces and P.

regina arrived after a delay of about 24 hours and increased in abundance as

decomposition progressed. Adults aerially collected (N=247) consisted of 75.3% P.

coeruleiviridis, 19.0% C. macellaria and 5.7% P. regina (Figure 3-25 A). Reared adults

(N=229) consisted of 69.0% P. coeruleiviridis, 12.7% C. macellaria, 0.4% C. rufifaces,

and 17.9% P. regina (Figure 3-25 B). Preserved larvae (N=301) consisted of 77.7% P.

coeruleiviridis, 8.3% C. macellaria and 14.0% P. regina (Figure 3-25 C).









Collection 15, April 26-May 1, 2003

Figures 3-26 A-C illustrate successional data. Phaenicia coeruleiviridis was the

most dominant calliphorid species for the first 4 days of the sample period but decreased

in abundance as decomposition progressed. Cochliomyia macellaria, C. rufifaces and P.

regina arrived after a delay of about 24 hours and increased in abundance as

decomposition progressed. Adults aerially collected (N=185) consisted of 68.1% P.

coeruleiviridis, 29.2% C. macellaria, 0.5% C. rufifaces, and 2.2% P. regina (Figure 3-26

A). Reared adults (N=359) consisted of 50.7% P. coeruleiviridis, 46.8% C. macellaria,

2.5% P. regina (Figure 3-26 B). Preserved larvae (N=651) consisted of 50.5% P.

coeruleiviridis, 47.9% C. macellaria, 1.4% P. regina and 0.2% were unknown (Figure 3-

26 C).

Collection 16, June 12-15, 2003

Figures 3-27 A-C illustrate successional data. Phaenicia coeruleiviridis was the

most abundant species of adults aerially collected on the first 2 days of the sampling

period, but C. macellaria was the most abundant species of calliphorid overall during the

sampling period (Figure 3-27 A). Adults aerially collected (N=221) consisted of 37.1%

P. coeruleiviridis, 56.1% C. macellaria, 2.3% C. rufifaces, and 4.5% P. regina (Figure 3-

27 A). Reared adults (N=150) consisted of 6.0% P. coeruleiviridis, 64.7% C. macellaria

and 29.3% C. rufifaces (Figure 3-27 B). Preserved larvae (N=388) consisted of 73.7% P.

coeruleiviridis, 19.3% C. macellaria, 5.4% C. rufifaces, 1.3% P. regina and 0.3% were

unknown (Figure 3-27 C).

Collection 17, December 8-20, 2003

Phaenicia coeruleiviridis was the most dominant calliphorid species during the

sampling period (Figures 3-28 A-C). Adults aerially collected (N=186) consisted of









89.8% P. coeruleiviridis, 0.5% C. macellaria, 2.2% C. rufifaces, 6.5% P. regina, and

1.1% C. livida (Figure 3-28 A). Reared adults (N=266) consisted of 95.9% P.

coeruleiviridis, 3.4% C. rufifaces and 0.8% P. regina (Figure 3-28 B). Preserved larvae

(N=648) consisted of 90.4% P. coeruleiviridis, 1.2% C. rufifaces and 8.3% P. regina

(Figure 3-28 C).

Collection 18, January 23-31, 2004

Figures 3-29 A-C illustrate the successional data. Phaenicia coeruleiviridis was the

most abundant dominant species during the sampling period. Phormia regina, C.

megacephala and C. livida arrived after a delay of about 24 hours. Adults aerially

collected (N=220) consisted of 92.3% P. coeruleiviridis, 5.9% P. regina, 0.5% C.

megacephala, 1.4% C. livida (Figure 3-29 A). Reared adults (N=294) consisted of 94.2%

P. coeruleiviridis and 5.8% P. regina (Figure 3-29 B). Preserved larvae (N=347)

consisted of 82.7% P. coeruleiviridis and 17.3% P. regina (Figure 3-29 C).

Collection 19, March 5-14, 2004

Figures 3-30 A-C illustrate the successional data. Phaenicia coeruleiviridis was the

most abundant dominant species during the first few days of the sampling period, but

decreased in abundance as decomposition progressed. Cochliomyia macellaria and P.

regina arrived after a 24 hour delay and increased in abundance as decomposition

progressed. Adults aerially collected (N=165) consisted of 45.5% P. coeruleiviridis, 6.7%

C. macellaria, and 47.9% P. regina. (Figure 3-30 A). Reared adults (N=149) consisted of

33.6% P. coeruleiviridis, 0.7% C. macellaria, 65.8% P. regina (Figure 3-30 B).

Preserved specimens (N=305) were comprised of 54.8% P. coeruleiviridis, 3.0% C.

macellaria and 42.3% P. regina (Figure 3-30 C).













E Percentage aerial

O Percentage reared

* Percentage preserved
3rd-instar larvae


80.0


70.0


60.0


50.0


< 40.0


30.0


20.0


10.0


0.0


Phaenicia Cochliomyia Chrysomya
coeruleiviridis macellaria rufifaces


16
03 o00


04 01 00 0000200 0000003


Phormia regina Chrysomya Calliphora livida
megacephala


Figure 3-1. Relative abundance of calliphorid adults aerially collected, adults reared from
larvae and preserved third-instar larvae collected from pig carrion during the
entire study (N=15,396) between November 16, 2001 and March 14, 2004 in
Earleton, Florida.


779
-765


16 0


70 80 82 6


kAi.


Calliphora
vicina


unknown









A






















(B)











t1 i :**** ,

Figure 3-2. A calliphorid first-instar larva hatching from its egg, (A), and the tiny Y-V
shaped spiracles of a first-instar larva (B).




































Figure 3-3. The two sets of spiracles of a first-second transitional larva. The first-instar
Y-V slits (small, dark) can be seen below the two slit spiracles (larger, light-
colored) in this molting larva.


























Figure 3-4. The two inner slits in the spiracles of a second-instar calliphorid larva.






























Figure 3-5. The two sets of spiracles of a second-third transitional calliphorid larva. The
second-instar spiracles (small, dark) have two slits, while the third-instar
spiracles have three (larger, light-colored) in this molting larva.





































Figure 3-6. The three inner splits in the spiracles of a third-instar calliphorid larva.






































Figure 3-7. Chrysomya rufifaces, third-instar larvae. The obvious fleshy protuberances
found on the second- and third-instar larvae of this species are easy to see
without a microscope.

















80 0
76 5






60 0



C,


S 40 0
.o






20 0


91
73 68


03

Phaenicia Cochliomyia Chrysomya rufifaces Phormia regina unknown
coeruleiviridis macellaria



Figure 3-8. Relative abundance of third-instar calliphorid larvae preserved from pig

carrion during the study (N=8253) in Earleton, Florida.
















100.0

II
z






-0
S80.0





60.0
ro









S60.0






0.0


Nov. 16-21, Dec. 29, February 5- March 15- April29- May 20-23, July22-25, Aug. 19-23, September October26-
2001 2001- 9, 2002 19, 2002 May 1,2002 2002 2002 2002 23-27, 2002 28, 2002
Jan.11, 2002


O PFaericia coeruleiviridis U Cochliomyia macellaria
o Chrysomya megacephala U Calliphora livida


Figure 3-9. Calliphorid activity for year 1 of the study.


[ Chrysomya rufifaces
* Calliphora vicina


M Phormia regina








47





100.0



z

80.0






. 60.0

0

o


S40.0

0



S20.0




0 .0 ,


November 30- December 30, March 2-8, April 1-6, April 26-May June 12-15, December8- January23- March 5-14,
December 14, 2002-January 2003 2003 1, 2003 2003 20, 2003 31, 2004 2004
2002 11,2003

0 Phaenicia coeruleiviridis U Cochliomyia macellaria E Chrysomya rufifaces
I Phormia regina 0 Chrysomya megacephala U Calliphora livida


Figure 3-10. Calliphorid activity for year 2 of the study.









48




(A)
40

Mean Temperature


Mean Low Temperature 328 332

30 304 305 93
SI 293 28
S28


236 23 9 237
I i ~224
20 202 199 194 19

< 174








5 52 18
1 15-










0 09

Nov 16-21, Dec 29, February 5- March 15-19, April29-May May20-23, July22-25, Aug 19-23, September October2E
2001 2001-Jan 11, 9, 2002 2002 1, 2002 2002 2002 2002 23-27, 2002 28, 2002
2002



(B)

350 -

Mean Temperature

300 Mean Low Temperature 4 30 3

II 27 4

250 I I 250 24
SI 230
I I 22 1

200 1
184 17 1189
11168 1170 16974
S150
S133
E 108
I-I

100 97 9489



50



00
Nov 30- December 30, March 2-8, April 1-6, April 26-May June 12-15, Dec 8-20, January 23- March 5-14
December 14, 2002-January 2003 2003 1, 2003 2003 2003 31, 2004 2004
2002 11,2003



Figure 3-11. Mean daily and mean low temperatures (SD) for year 1, November 16,

2001 to October 26, 2002 (Part A) and year 2, November 30, 2002 to March

14, 2004 (Part B) of study.


6-


,














(A) Reared Adults


100.0





80.0


November 16, 2001


November 21, 2001


(B) Preserved Larvae
100.0


U.U
November 21, 2001


Figure 3-12. Reared adults, N=232, (Part A), and preserved larvae, N=30, (Part B), from

collection 1, November 16 to November 21, 2001.


B Phaenicia coeruleiviridis
* Cochliomyia maceflaria
E Chrysomya rufifaces


E Phaenicia coeruleiviridis


e:
W
a
E

a,


e:
W
a
E

a,














(A) Reared Adults
100.0 i


80.0


e:
0-
QJ
CJ

.3

,0


u.u
January 11, 2002


(B) Preserved Larvae
100.0 -


80.0


v?
0^-
1)




p:


December 29, 2001 January 11, 2002


Figure 3-13. Reared adults, N=362, (Part A), and preserved larvae, N=568, (Part B), from

collection 2, December 29, 2001 to January 11, 2002.


O Pinaerca coe, uleiviridis















Collected Aerially


February 1, 2002


(B) Reared Adults
100.0 1


February 5, 2002


February 9, 2002


(C) Preserved Larvae

80.0 .


February 9, 2002

Figure 3-14. Adults aerially collected, N=39, (Part A), reared adults, N=305, (Part B),

and preserved larvae, N=151, (Part C), from collection 3, February 1-9, 2002.


(A) Adults
100.0-


a

o
W
a
a

a,


p:


0.0


a


8




.i

p:


e:
B





4















(A) Adults Aerially Collected
100.0 -


March 15, 2002


March 17, 2002


(B) Reared Adults
100.0 1


v?
0^-
1)



.

I:


0.0 !rsre Ira


(C) Preserved Larvae
100.0o


e:




rt
T3


March 17, 2002


March 17, 2003


March 18, 2002


March 18, 2003


March 19, 2003


Figure 3-15. Adults collected aerially, N=71, (Part A), reared adults, N=326, (Par B), and

preserved larvae, N=134, (Part C), from collection 4, March 15-19, 2002.


e:




rt
T3


C -J


I I O unknown


0.0


0.0















(A) Adults Aerially Collected
100.0 .


3 Phaenicia coeruleiviridis
I Cochliomyia macellaria
12 Phormia regina
















April 29, 2002 May 1, 2002


(B) Reared Adults
100.0 I


April 29, 2002 May 1, 2002


(C) Preserved Larvae
100.0 -


May 1, 2002

Figure 3-16. Adults aerially collected, N=107, (Part A), reared adults, N=332, (Part B),

and preserved larvae, N=196, (Part C), from collection 5, April 29 to May 1,

2002.


e:
0-
QJ
CJ



T3


v?
0^
1)
CJ
aT



I:


e:




rt
T3


0.0














(A) Adults Aerially Collected
60.0 .


May 20, 2002 May 22, 2002 May 23, 2002


(B) Reared Adults
100.0 1


May 23, 2002


(C) Preserved larvae
100.0 .


May 23, 2002

Figure 3-17. Adults aerially collected, N=51, (Part A), reared adults, N=67, (Part B), and

preserved larvae, N=175 (Part C), from collection 6, May 20 to May 23, 2002.


e:
0-
QJ
CJ



T3


v?
0^-
1)



.

I:


e:




rt
T3


0.0














(A) Adults Aerially Collected


July 22, 2002


July 23, 2002


July 24, 2002


(B) Reared Adults
100.0 1


July 24, 2002


(C) Preserved Larvae
100.0


July 25, 2002


July 24, 2002


July 25, 2002


Figure 3-18. Adults aerially collected, N=99, (Part A), reared adults, N=223, (Part B),

and preserved larvae, N=700, (Part C), from collection 7, July 22 to July 24,

2002.


f Phaenicia coeruleiviridis
Hi Cochliomyia macellaria
B Chrysomya rufifaces
E Phormia regina
B Chrysomya megacephala







IinJI


I _


h

e:
a
a
E


.

a


0.0


3
a,
8




.

p:


e:
a





t~
a


0.0


00.0













(A) Adults Aerially Collected
100.0 .--


August 19, 2002


(B)




o^
1)
CJ
aT

I,

p)


August 20, 2002


O Phaenicia coeruleiviridis
2 Cochliomyia macellaria
m Chrysomya rufifaces
O Chrysomya megacephala










JKb


August 21, 2002


August 21, 2002


* unknown


August 21, 2002


August 22, 2002


Figure 3-19. Adults aerially collected, N=76, (Part A), reared adults, N=248, (Part B),
and preserved larvae, N=499, (Part C), from collection 8, August 19-23, 2002.


August 22, 2002


(C) Preserved Larvae
100.0 .


August 22, 2002 August 23, 2002


e:
W
a
E




a


I I


0.0


h

e:
a
a
E


.

a


80.0
















(A) Adults Aerially Collected
100.0 .


September 25, 2002


e:
W
a
E






a


OJ ,chaenicia coerdileiviridis
SCochionmyia ,naceh'aria


O Ghryso,,ya ,,egaoephaIa


September 26, 2002 September 27, 2002


O Phaenicia coeruleiviridis
O3 Coch/iomyia macellaria
m Chrysomya rufifaces
El Phormia regina


September 25, 2002


September 26, 2002


ISept
September 27, 2002


Figure 3-20. Adults aerially collected, N=76, (Part A), reared adults, N=323, (Part B),

and preserved larvae, N=499, (Part C), from collection 9, September 23-27,

2002.


September 23, 2002


(B) Reared Adults
100.0 I


(C) Preserved Larvae
100.0


September 25, 2002 September 26, 2002 September 27, 2002


0.0


3

a,
o
W
a
a


a,


p:


0.0


e:
a
a
E



.

a


80.0 -














(A) Adults Aerially Collected
70.0 .


October 26, 2002


[ Phaenicia coeruleiviridis
I Coch/iomyia macellaria
B Chrysomya rufifaces


October 27, 2002 October 28, 2002


(B) Reared Adults
70.0 1


00ctober 27, 2002
October 27, 2002


October 28, 2002


(C) Preserved Larvae
100.0 --


October 26, 2002 October 27, 2002 October 28, 2002


Figure 3-21. Adults aerially collected, N

and preserved larvae, N=294,

2002.


=87, (Part A), reared adults, N=131, (Part B),

(Part C), from collection 10, October 26-28.


e:
a




t~
a


0.0


3
a,
8




.

p:


e:
a




t~
a


80.0
















(A) Adults Aerially Collected
100.0 .


0.0 I I




(B) Reared Adults
100.0


November 30, 2002


December 8, 2002


December 7, 2002 December 8, 2002 December 10, December 12, December 14,
2002 2002 2002


(C) Preserved Larvae
100.0



80.0 -







400-



U 20.0 -




December 7, December 8, December 10, December 12, December 13, December 14,
2002 2002 2002 2002 2002 2002


Figure 3-22. Adults aerially collected, N=60, (Part A), reared adults, N= 384, (Part B),

and preserved larvae, N=1248, (Part C), from collection 11, November 30-

December 8, 2002.


Ml Phaenicia coeruleiviridis

H Cochliomyia macellaria


D; ~1


I _ _ _ I I___


e:
a





t~
a


80.0-



60.0-



40.0-



20.0


3

a,
8







p:















(A) Adults Aerially Collected
100.0 [I I I I I


December 30, 2002

(B) Reared Adults
100.0 --


January 3, January 4, January 5, January 6, January 8,
2003 2003 2003 2003 2003

(C) Preserved Larvae
100.0


January 3, January 4,
2003 2003


January 9, January 11,
2003 2003


Figure 3-23. Adults aerially collected, N=56, (Part A), reared adults, N= 371, (Part B),

and preserved larvae, N=581, (Part C), from collection 12, December 30, 2002

to January 11, 2003.


E Phaenicia coeruleiviridis

M Calliphora livida


E Phaenicia coeruleiviridis
a Coch/iomyia macellaria
L unknown














* H


January 5, January 6, January 8, January 9, January 11,
2003 2003 2003 2003 2003


' ' '


e:
a





t~
a


0.0


January 2, 2003


January 5, 2003


January 6, 2003


3

a,
8




.

p:


e:
a





t~
a


0.0
















(A) Adults Aerially Collected
100.0 .


March 1, 2002 March 2, 2003 March 5, 2003 March 6, 2003 March 7, 2003


(B) Reared Adults
100.0 .


March 5, 2003


(C) Preserved Larvae
100.o .-


March 6, 2003


March 7, 2003


March 8, 2003


March 5, 2003


March 6, 2003


- unknown


March 7, 2003


March 8, 2003


Figure 3-24. Adults aerially collected, N=66, (Part A), reared adults, N=166, (Part B),

and preserved larvae, N=340, (Part C), from collection 13, March 1-8, 2003.


0.0 I


e:
W
a
E






a


0.0


3

a,
8





.

p:


e:
a
a
E



.

a


80.0-



60.0-



40.0-



20.0-














(A) Adults Aerially Collected
100.0 .


April 1, 2003 April 2, 2003 April 4, 2003 April 5, 2003 April 6, 2003


(B) Reared Adults
100.0 I


April 4, 2003

(C) Preserved Larvae


April 4, 2003


fl Phaenicia coeruleiviridis
I Cochliomyia macellaria
B Chrysomya rufifaces
II Phormia regina


April 5, 2003


April 6, 2003


_1 I


April 5, 2003


April 6, 2003


Figure 3-25. Adults aerially collected, N=247, (Part A), reared adults, N=229, (Part B),

and preserved larvae, N=301, (Part C), from collection 14, April 1-6, 2003.


I I 1 IIIII ~NNNNNN~ I


e:
W
a
E





a


0.0


3
a,
8




.

p:


0.0


100.0 -


e:
a




t~
a












(A) Adults Aerially Collected
100.0 -


April 26, 2003


El Phaenicia coeruleiviridis
I Cochliomyia macellaria
I Chrysomya rufifaces
1 Phormia regina









irL/rL


April 27, 2003


April 28, 2003


April 29, 2003


April 30, 2003


(B) Reared Adults
100.0 i


April 27, 2003 April 28, 2003


April 29, 2003
April 29, 2003


April 3U, 2UU3


(C) Preserved Larvae
100.0


O 80.0


60.0


- 40.0-
A 40.0
p:


April 28, 2003 April 29, 2003 April 30, 2003 May 1, 2003


Figure 3-26. Adults aerially collected, N=185, (Part A), reared adults, N=359, (Part B),
and preserved larvae, N=651, (Part C), from collection 15, April 26-May 1,
2003.


e:
0-
QJ
CJ

.3
,0


-3

I,
I)
_^
rt
1)
a,


0.0


0.0
















(A) Adults Aerially Collected
100.0 .


e:
0-
QJ
CJ



T3


June 12, 2003 June 13, 2003 June 14, 2003 June 15, 2003


(B) Reared Adults
100.0 1


-3

I,
I)
_^
rt
1)

a,


June 13,2003


June 14, 2003


June 15, 2003


(C) Preserved Larvae
100.0 -


June 14,2003


June 15, 2003


Figure 3-27. Adults aerially collected, N=221, (Part A), reared adults, N=150, (Part B),

and preserved larvae, N=388, (Part C), specimens from collection 16, June 12-

15, 2003.


0.0 ---


Y?
0--
QJ
CJ
T3


.

<;


I unknown


0.0


80.0
















(A) Adults Aerially Collected
100.0 .O


December 8, December 10, December 11, December 12,
2003 2003 2003 2003


(B) Reared Adults
100.0 I I


December 13, December 14, December 16,
2003 2003 2003


December 12, December 13, December 14, December 16, December 17, December 20,
2003 2003 2003 2003 2003 2003


(C) Preserved Larvae
100.0 -


December 13,
2003


December 14,
2003


December 16,
2003


December 17,
2003


December 20,
2003


Figure 3-28. Adults aerially collected, N=186, (Part A), reared adults, N=266, (Part B),

and preserved larvae, N=648, (Part C), from collection 17, December 8-20,

2003.


U I L-


~ Phaenicia coerulervlridls
I Cochllomyla macellaria
B Chrysomya ruflfaces
ad Phorrma regina
I Calllphora Irvida


e:
W
a
E






a


0.0


3

a,
o
W
a
a


a,


p:


0.0


h


e:
a
a
E



.

a














(A) Adults Aerially Collected
100.0 -----


January January January January January January January January January
23, 2004 24, 2004 25, 2004 26, 2004 27, 2004 28, 2004 29, 2004 30, 2004 31, 2004


(B) Reared Adults
100.0 r


January 27, 2004

(C) Preserved Larvae
100.0


January 28, 2004


January 29, 2004


January 28, 2004 January 29, 2004


January 30, 2004


January 31, 2004


Figure 3-29. Adults aerially collected, N=220, (Part A), reared adults, N=294, (Part B),

and preserved larvae, N=347, (Part C), from collection 18, January 23-31,

2004.


e:
0-
QJ
CJ


T3


-3

I,
I)
_^
rt
1)
a,


I I I _I I _


v?
0^-
1)



.

I:


0.0


0.0









67




(A) Adults Aerially Collected
100.o0
i Phaenicia coeruleiviridis

80.0 m Coch/iomyia macellaria

o 1 Phormia regina
-60.0


40.0

20.0 -





March 5, March 6, March 7, March 8, March 9, March 13, March 14,
2004 2004 2004 2004 2004 2004 2004


(B) Reared Adults
100.o



80.0 -



-oo.o



S 40.0



20.0 -



0.0
March 8, 2004 March 9, 2004 March 10, 2004 March 14, 2004


(C) Preserved Larvae






S 80.0 -



-00.0o







20.0 -




March 8, 2004 March 9, 2004 March 10, 2004 March 13, 2004 March 14, 2004


Figure 3-30. Adults aerially collected, N=316, (Part A), reared adults, N=639, (Part B),

and preserved larvae, N=675, (Part C), from collection 19, March 5-14, 2004.














CHAPTER 4
DISCUSSION

The most common calliphorid species collected on pig carrion during the study

aerially or by rearing was Phaenicia coeruleiviridis. Of the total adult specimens

(N=6971) identified during the study, 75.0% were P. coeruleiviridis. Similarly, of the

identifiable third-instar larvae (N=8253), 76.5% were P. coeruleiviridis. This species was

most abundant from late October to the end of May (Figures 3-9 and 3-10). Chrysomya

rufifaces and C. macellaria were also collected during those months, but in much smaller

numbers. Phaenicia coeruleiviridis was less abundant during the summer months-when

the temperatures were over 25.00 C. Cochliomyia macellaria was not present during the

winter and was the most abundant species collected in June 2003 (Figures 3-9 and 3-10).

Phormia regina was present from mid-November 2001 until the end of July 2002

(Figures 3-9 and 3-10). No specimens were collected in August or September in 2002. It

was the most abundant species collected in March 2004, but that was unusual compared

to the rest of the study because P. regina was quite low in abundance compared to the

other calliphorids. Hall (1948) found that P. regina is abundant in the spring months in

the southern states (location not defined by Hall) but apparently this was not the case

during my study in rural north-central Florida.

Of the seven species of calliphorids collected during the study, all were consistent

with published distribution records, seasonality, and successional patterns (Campobasso

et al. 2001). However, Byrd (1998) collected Phaenica cuprina (Wiedemann) (=

Phaeniciapallescens Shannon) from pig carcasses in June and September 1996, and also









in September 1997, but this species was not found during this study. Byrd's study was

conducted at an undesignated location in the Gainesville area, a "woodland habitat" (page

28 of dissertation), which consisted of a Live Oak hammock with no understory or

roosting sites (for female blow flies) within close proximity to the pigs. Phaenicia

cuprina is an urban fly that prefers excrement to carrion (Byrd 2001), which could

explain why it was not found on pig carrion in rural Earleton.

During this study, the species and relative abundances of calliphorids found during

adult aerial, reared and larval collections made during December 2002 and 2003 (Figures

3-23 A and B and 3-28 A-C) were similar, as were the collections made during

November 2001 and November 2002 (Figures 3-12 A and B, 3-22 A-C). Finally, data

obtained for three years (2002, 2003, and 2004) in March (Figures 3-15 A-C, 3-24 A-C,

and 3-30 A-C) also yielded similar species and relative abundances of species. The

congruence between these samples suggests that there is a consistent pattern in species

composition and relative abundance through time in this sample site.

Common calliphorid species such as Phaenicia sericata and Phaenicia eximia

apparently are not normally found in rural north-central Florida, but were found in the

urban Gainesville area by Byrd (1998) and Peters (2003). Different calliphorids are

associated with different habitats. An urban area, with odors from human refuse, cooking,

garbage dumps, and improper sanitation will have different species assemblages than

rural, wooded areas or arid regions. Some calliphorid species were not expected to be

found because they are not associated with carrion. For example, Hall (1948) stated that

the calliphorid Pollenia rudis (Fabricus) has been collected in northern Florida, but this

species is exclusively a parasite of earthworms.









Calliphorid species (both adults and larvae) found on human remains (in cases from

1991) in Palm Beach and Lake Counties, Florida were identified as C. rufifaces and C.

macellaria (Haskell 2003, personal communication). The evidentiary specimens were

identified by Neal Haskell during the court cases. These two species were collected on

pig carrion in rural north-central Florida during my study, on pig carrion in Gainesville

by Byrd (1998) and on bear carrion in Gainesville by Peters (2003).

In northwest Indiana, Haskell (1989) found P. regina to be the most abundant

species of Calliphoridae from late spring to early autumn. Phaenicia coeruleiviridis was

dominant during the spring and autumn. Haskell also found a few C. vicina and C. livida

specimens during the spring and autumn.

In West Virginia, Joy et al. (2002) found P. regina to be the dominant calliphorid

in May 2002 on raccoon carrion. They found only a few Phaenicia species on these

raccoons, which were killed by cars so that the time of death was unknown. The dead

raccoons were frozen and transported to the site in garbage bags.

In Menard County Texas, Cushing and Parish (1938) captured Cochliomyia and

Phaenicia species from April to November 1933, but P. regina was far more abundant.

They caught hundreds of thousands of flies with pit-fall traps and traps baited with beef.

No P. coeruleiviridis were found.

Calliphora vicina, C. livida, P. coeruleiviridis, C. macellaria, C. rufifaces, P.

regina and C. megacephala were collected at both rural and urban sites during a recent

pig carrion study in central Texas (Tenorio et al. 2003). Phaenicia coeruleiviridis was

found from March until May while C. livida and C. vicina were found from December to

May. Cochliomyia macellaria were found all year and C. rufifaces was found from









March until November. Chrysomya megacephala was found from September to

November, and P. regina was not found from June through August.

In northern Mississippi, P. coeruleiviridis, P. regina and C. macellaria were the

dominant species from April to September (Goddard and Lago 1985). Phormia regina

was dominant from October to March on carcasses of rabbits, opossum, and fish

carcasses.

Deonier (1937, 1938) collected hundreds of thousands of calliphorids with beef-

baited traps in Arizona. Phormia regina, C. macellaria and Phaenicia species were

found. Cochliomyia macellaria were captured in enormous numbers all year. A single P.

regina specimen was collected in August and again in September of 1937, while almost

293,000 C. macellaria and 1000 Phaenicia species were collected at the same time

(Deonier 1942).

Denno and Cothran (1975) found P. sericata and P. regina to be the dominant

calliphorids on rabbit carrion in Davis, CA during June to September. Hall and Doisy

(1993) found C. macellaria, P. regina, and P coeruleiviridis during their field studies in

Missouri from mid-June to late August 1992.

Reed (1958) found C. livida, C. vicina, C. macellaria, P. coeruleiviridis, and P.

regina during his dog carcass study in Tennessee. Phormia regina was the most abundant

calliphorid species on the dog carcasses (Reed 1958).

Watson and Carlton (2003) conducted research in Louisiana from April 1 to July

1999 using bear, deer, alligator, and pig carcasses and found similar species to those of

my study. The most common species on all four carcasses (in order of occurrence) were

P. regina, P. coeruleiviridis, and P. sericata. Of note was that P. coeruleiviridis landed









on carcasses within minutes of deposition. Cochliomyia macellaria and C. rufifaces were

also found on the pig carcasses, but in smaller numbers.

Peters (2003, page 78) conducted a carrion study using three dead bears (Ursus

americanusfloridanus) in the Gainesville area and concluded that "the most abundant

blow fly found in north central Florida is the Hairy Maggot Blow Fly, Chrysomya

/ IIJif e'\". Only 8 P. coeruleiviridis adults were among 94 calliphorid specimens

collected by Peters (2003). This is contrary to my results, which indicate that P.

coeruleiviridis is most abundant on pig carrion in this area while C. rufifaces consisted of

less than 10 % of the total specimens collected during the entire study. The differences in

our data are even greater when comparing abundances of C. megacephala, which

consisted of less than 2.0 % of the total flies during my study while Peters found this

species to comprise 44.7 % of adult calliphorids collected in her study.

The differences in our findings may be attributed to the following:

* The actual time of death of the bears was approximated; time "0" was not the
actual time of death. Once retrieved by the wildlife officer, the dead bears were not
protected in plastic or placed in containers while being transferred to the study site.
At least 48 hours passed before any sample collections were taken on Bear 1.
Therefore, each bear had a collection delay of at least 48 hours. These differences
in the method of procuring and placement of carcasses may have resulted in
completely missing the first wave of carrion insects attracted to fresh carcasses,
specifically P. coeruleiviridis. Apparently, P. coeruleiviridis larvae were collected
on all three bears, which clearly indicate that the adults had been present.

* Bears may decompose differently than do pigs or humans; bears have thick fur,
thick skin, and layers of adipose tissue that are unique to bears (Watson and Carlton
2003). These differences may make bear carcasses more attractive to C. rufifaces
and C. megacephala or less attractive to P. coeruleiviridis. Watson and Carlton
(2003) noticed that P. coeruleiviridis arrived earlier on the carcasses of deer,
alligator and pigs than on the carcass of the bear.

* The number of larval specimens collected or reared (if any) is not disclosed by
Peters (2003). Apparently, only 94 adult calliphorid specimens were aerially
collected, of which almost half were C. megacephala and 19 were C. rufifaces.









Additionally, aerial collections were made late in the day (between 5-6 PM) when
some calliphorid flies are less active.

* The three bears were each placed at the site by Peters (2003) during different
months of the year. As a result, there could be variability due to a lack of
replications or different seasons.

In my study, P. coeruleiviridis was the most abundant species of calliphorid found

on pig carrion in rural north-central Florida between 2001and 2004 during most of the

year. This species was always the first to arrive at fresh pig carrion, the first to deposit

eggs, the first to complete development and the first to migrate off the carcass to

pupariate in the soil. In the spring and summer, these events took 5 days, or fewer, to

complete. Depending on the season, and almost always after an approximate 24-h delay,

C. macellaria, C. rufifaces, P. regina and C. megacephala arrived at the pig carcasses.

Calliphora livida and C. vicina arrived at the carcasses also after a delay, but only a few

specimens of each were collected (Figures 3-3 and 3-4).

Despite the fact that P. coeruleiviridis is commonly found in many other states,

there are no developmental rate data for this species because, until now, no one has been

able to successfully rear the larvae to adulthood (Haskell 1989, Hall and Doisy 1993).

During this study, I discovered an organic pupation substrate that results in successful

rearing of P. coeruleiviridis larvae to adulthood. Now that we can rear P. coeruleivirdis,

we need detailed developmental rate data for this species so a forensic entomologist can

determine accumulated degree hours (ADH) or days (ADD) to determine the postmortem

interval.

The fact that we do not have developmental data for P. coeruleiviridis is very

unfortunate because it is an important indicator species for determining PMI, especially

in rural north-central Florida. Standardized rearing data are needed for P. coeruleiviridis,






74


as are rearing data for other calliphorids that are attracted to human remains including

Calliphora vicina, C. vomitoria, Eucalliphora sp., Cynomyopsis cadaverina, Phaenicia

sericata, Lucilia Illustris, Phormia regina, Cochliomyia macellaria and Paralucilia

wheeleri.




















APPENDIX A
RAW DATA: FLIES COLLECTED AS ADULTS OR REARED FROM LARVAE


PIG: DATE:
2A 11/16/2001
2B 11/16/2001
2B 11/21/2001
2C 11/16/2001
3A 1/11/2002
3B 1/11/2002
3C 1/11/2002
4A 2/1/2002
4A 2/5/2002
4B 2/1/2002
4B 2/5/2002
4C 2/1/2002
4C 2/5/2002
4E 2/1/2002
4E 2/5/2002
4E 2/9/2002
5A 3/15/2002
5A 3/17/2002
5A 3/17/2002
5A 3/18/2002
5B 3/15/2002
5B 3/17/2002
5C 3/15/2002
5C 3/17/2002
5E 3/15/2002
5E 3/17/2002
5E 3/18/2002
6A 4/29/2002
6A 5/1/2002
6A 5/1/2002
6B 4/29/2002
6B 5/1/2002
6B 5/1/2002
6B 5/1/2002
6C 4/29/2002
6C 5/1/2002
6C 5/1/2002
6C 5/1/2002
6E 4/29/2002
6E 4/29/2002


sample #
A, R # adults
R 7A 21
R 8A 29
R 10A 157
R 9A 25
R 13A 145
R 17A 115
R 15A 102
A 23 4
R 24 54
A 22 12
R 25 71
A 21 12
R 26 38
A 20 11
R 27 55
R 28 87
A 30 17
R 35 59
A 36 22
R 47 65
A 31 15
R 40 56
A 32 7
R 38 42
A 33 10
R 42 47
R 44A 57
A 49 11
R 53 58
A 55 13
A 50 17
R 57 33
R 59 42
A 61 6
A 51 44
R 63 56
R 65 38
A 66 0
A 52 16
R 70 69


P.
coerul.
8
28
157
25
145
115
100
4
54
12
71
12
38
11
55
87
17
59
22
65
15
56
7
42
8
47
57
11
58
0
17
33
42
4
44
56
38
0
16
36


C.
macellaria
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
9


C.
ruffifaces
13
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0


P.
regina
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
13
0
0
0
1
0
0
0
0
0
24


C.
megaceph. C. livida C. vicina













PIG: DATE:
6E 5/1/2002
7A 5/20/2002
7A 5/22/2002
7A 5/23/2002
7A 5/23/2002
7B 5/20/2002
7C 5/20/2002
7E 5/20/2002
8A 7/22/2002
8A 7/23/2002
8A 7/24/2002
8A 7/24/2002
8A 7/24/2002
8A 7/25/2002
8B 7/22/2002
8B 7/23/2002
8C 7/22/2002
8C 7/23/2002
8C 7/24/2002
8C 7/24/2002
8E 7/22/2002
8E 7/24/2002
8E 7/24/2002
8E 7/24/2002
8E 7/25/2002
9A 8/19/2002
9A 8/20/2002
9A 8/21/2002
9A 8/21/2002
9A 8/22/2002
9A 8/22/2002
9A 8/23/2002
9B 8/19/2002
9B 8/20/2002
9B 8/21/2002
9B 8/21/2002
9B 8/22/2002
9B 8/22/2002
9B 8/23/2002
9C 8/19/2002
9C 8/20/2002
9C 8/20/2002
9C 8/22/2002
9C 8/22/2002
9E 8/20/2002
9E 8/21/2002
9E 8/21/2002
9E 8/22/2002
9E 8/22/2002


sample #
A, R # adults
R 68 36
A 72 11
A 76 18
A 77 8
R 78 67
A 73 12
A 74 2
A 75 0
A 80 0
A 84 11
A 91 11
R 92 21
R 94 25
R 107 57
A 81 9
A 87 13
A 82 11
A 86 7
A 102 5
R 103 42
A 85 8
A 83 17
A 97 7
R 98 24
R 109 54
A 111 14
A 115 18
R 119 5
A 123 10
A 153 2
R 157 11
R 154 27
A 112 2
A 116 10
R 124 43
A 125 10
A 148 4
R 149 14
R 151 38
A 113 1
A 117 12
A 134 0
R 135 13
R 137 16
A 118 11
A 129 8
R 130 27
A 114 11
A 142 5


P.
coerul.
36
0
3
1
67
4
1
0
0
4
0
21
24
0
8
10
11
5
0
42
3
17
0
18
0
14
18
2
3
0
8
4
2
10
16
3
1
12
0
1
11
0
4
14
10
0
27
7
0


C.
macellaria
0
11
6
1
0
2
0
0
0
3
0
0
0
0
1
0
0
0
0
0
0
0
1
0
0
0
0
0
1
2
0
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
2
2


C.
ruffifaces
0
0
3
2
0
6
0
0
0
3
9
0
1
57
0
2
0
2
5
0
5
0
5
6
54
0
0
1
6
0
1
20
0
0
7
6
2
1
38
0
1
0
7
0
1
7
0
2
3


P.
regina
0
0
6
4
0
0
1
0
0
1
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0


C.
megaceph. C. livida


C. vicina
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0








77



sample # P. C. C. P. C.
PIG: DATE: A, R # adults coerul. macellaria ruffifaces regina megaceph. C. livida C. vicina
9E 8/22/2002 R 145 13 10 0 2 0 1 0 0
9E 8/23/2002 R 143 41 10 2 28 0 1 0 0
10A 9/23/2002 A 159 0 0 0 0 0 0 0 0
10A 9/25/2002 A 163 10 3 4 3 0 0 0 0
10A 9/25/2002 R 165 32 32 0 0 0 0 0 0
10A 9/26/2002 A 177 13 0 8 5 0 0 0 0
10A 9/26/2002 R 178 22 22 0 0 0 0 0 0
10A 9/26/2002 R 180 22 15 0 0 0 7 0 0
10B 9/23/2002 A 160 6 6 0 0 0 0 0 0
10B 9/25/2002 A 166 10 3 5 2 0 0 0 0
10B 9/25/2002 R 167 32 27 5 0 0 0 0 0
10B 9/26/2002 R 183 16 16 0 0 0 0 0 0
10B 9/26/2002 R 185 55 0 28 0 0 27 0 0
10C 9/23/2002 A 162 10 10 0 0 0 0 0 0
10C 9/25/2002 A 174 9 5 1 3 0 0 0 0
10C 9/25/2002 R 175 26 26 0 0 0 0 0 0
10C 9/27/2002 A 187 6 0 4 2 0 0 0 0
10C 9/27/2002 R 188 42 12 17 4 0 9 0 0
10E 9/23/2002 A 161 7 7 0 0 0 0 0 0
10E 9/25/2002 A 170 5 0 4 1 0 0 0 0
10E 9/25/2002 R 171 25 25 0 0 0 0 0 0
10E 9/27/2002 R 191 51 0 49 0 0 2 0 0
11A 10/26/2002 A 193 8 2 1 5 0 0 0 0
11A 10/27/2002 A 200 3 0 2 1 0 0 0 0
11A 10/27/2002 R 201 43 20 9 14 0 0 0 0
11A 10/27/2002 R 203 31 15 0 16 0 0 0 0
11A 10/28/2002 A 209 8 0 1 7 0 0 0 0
11A 10/28/2002 R 210 14 14 0 0 0 0 0 0
11A 10/28/2002 R 212 28 20 1 7 0 0 0 0
11B 10/26/2002 A 195 17 3 5 9 0 0 0 0
11B 10/26/2002 R 196 0 0 0 0 0 0 0 0
11C 10/26/2002 A 199 19 2 8 9 0 0 0 0
11C 10/28/2002 A 205 6 1 1 4 0 0 0 0
11C 10/28/2002 R 206 15 0 0 15 0 0 0 0
11E 10/26/2002 A 198 17 3 7 7 0 0 0 0
11E 10/28/2002 A 208 9 0 6 3 0 0 0 0
12A 11/30/2002 A 214 10 10 0 0 0 0 0 0
12A 12/7/2002 R 221 33 33 0 0 0 0 0 0
12A 12/8/2002 R 228 34 34 0 0 0 0 0 0
12A 12/8/2002 A 230 19 14 1 3 1 0 0 0
12A 12/12/2002 R 253 29 29 0 0 0 0 0 0
12A 12/13/2002 A 255 0 0 0 0 0 0 0 0
12A 12/14/2002 R 258 21 21 0 0 0 0 0 0
12B 11/30/2002 A 215 6 6 0 0 0 0 0 0
12B 11/30/2002 R 218 0 0 0 0 0 0 0 0
12B 12/7/2002 R 222 33 33 0 0 0 0 0 0
12B 12/8/2002 R 231 25 25 0 0 0 0 0 0
12B 12/8/2002 A 233 0 0 0 0 0 0 0 0
12B 12/12/2002 R 249 9 9 0 0 0 0 0 0













PIG: DATE:
12B 12/12/2002
12C 11/30/2002
12C 12/7/2002
12C 12/10/2002
12C 12/10/2002
12C 12/12/2002
12E 11/30/2002
12E 12/7/2002
12E 12/8/2002
12E 12/10/2002
12E 12/12/2002
13B 12/30/2002
13B 1/2/2003
13B 1/3/2003
13B 1/4/2003
13B 1/5/2003
13B 1/9/2003
13B 1/9/2003
13C 12/30/2002
13C 1/2/2003
13C 1/3/2003
13C 1/4/2003
13C 1/5/2003
13C 1/5/2003
13C 1/8/2003
13C 1/9/2003
13C 1/11/2003
13E 12/30/2002
13E 1/2/2003
13E 1/3/2003
13E 1/4/2003
13E 1/5/2003
13E 1/5/2003
13E 1/6/2003
13E 1/6/2003
13E 1/8/2003
13E 1/9/2003
13E 1/9/2003
14A 3/2/2003
14A 3/6/2003
14A 3/6/2003
14A 3/6/2003
14A 3/8/2003
14A 3/8/2003
14B 3/5/2003
14B 3/5/2003
14E 3/1/2003
14E 3/7/2003
14E 3/7/2003


sample #
A, R # adults
R 251 2
A 216 10
R 226 24
R 235 27
R 239 46
R 245 9
A 217 7
R 224 38
A 234 8
R 240 36
R 247 18
A 262 2
A 263 17
R 266 27
R 276 30
R 283 21
R 292 20
A 294 0
A 261 7
A 264 8
R 270 27
R 274 21
A 281 1
R 282 34
R 290 9
R 298 27
R 300 28
A 260 13
A 265 4
R 268 17
R 272 25
A 278 1
R 279 28
A 285 3
R 286 39
R 288 9
R 295 9
A 297 0
A 318 16
R 304 39
R 306 42
A 308 16
R 314 15
R 316 26
A 301 20
R 302 21
A 300 11
R 309 23
A 311 3


P.
coerul.
2
10
24
27
0
9
7
38
8
36
18
2
17
27
27
21
20
0
7
7
27
21
1
34
9
27
28
13
4
17
25
1
28
3
39
9
9
0
16
38
40
11
15
26
19
21
10
23
2


C.
macellaria
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0


C.
ruffifaces
0
0
0
0
46
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0


P.
regina
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
2
5
0
0
0
0
0
0
1


C.
megaceph. C. livida


C. vicina
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0













PIG: DATE:
14E 3/7/2003
15A 4/1/2003
15A 4/2/2003
15A 4/4/2003
15A 4/5/2003
15A 4/5/2003
15A 4/6/2003
15A 4/6/2003
15B 4/1/2003
15B 4/2/2003
15B 4/4/2003
15B 4/4/2003
15B 4/5/2003
15B 4/5/2003
15C 4/1/2003
15C 4/2/2003
15C 4/4/2003
15C 4/4/2003
15C 4/5/2003
15C 4/5/2003
15C 4/6/2003
15C 4/6/2003
15E 4/1/2003
15E 4/2/2003
15E 4/4/2003
15E 4/4/2003
15E 4/5/2003
15E 4/5/2003
15E 4/6/2003
15E 4/6/2003
16B 4/26/2003
16B 4/27/2003
16B 4/28/2003
16B 4/28/2003
16B 4/29/2003
16B 4/29/2003
16B 4/30/2003
16C 4/26/2003
16C 4/27/2003
16C 4/28/2003
16C 4/28/2003
16C 4/29/2003
16C 4/29/2003
16C 4/30/2003
16C 4/30/2003
16C 5/1/2003
16E 4/26/2003
16E 4/27/2003
16E 4/27/2003


sample
#
312
320
324
330
340
342
352
353
321
325
331
333
343
345
319
323
327
329
337
339
349
350
322
326
334
336
346
348
355
357
358
365
372
373
381
389
388
360
361
366
367
375
376
383
384
393
359
362
363


#
adults
0
13
21
18
23
12
16
2
9
26
28
13
24
6
10
21
20
52
0
21
0
13
11
10
23
12
28
6
36
2
31
11
13
33
34
6
19
14
21
14
34
11
63
5
85
0
16
13
19


P.
coerul.
0
13
20
6
21
4
7
0
9
25
20
11
19
3
10
21
15
50
0
7
0
3
9
10
23
10
21
5
1
1
31
11
9
28
34
5
19
13
9
1
32
2
0
0
0
0
16
6
19


C.
macellaria
0
0
0
12
0
8
6
0
0
0
1
2
2
1
0
0
3
0
0
10
0
4
2
0
0
1
5
1
17
1
0
0
3
5
0
1
0
1
12
11
2
8
59
5
84
0
0
7
0


C.
ruffifaces
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0


P.
regina
0
0
1
0
2
0
3
2
0
1
7
0
3
2
0
0
2
1
0
4
0
6
0
0
0
1
2
0
18
0
0
0
1
0
0
0
0
0
0
1
0
1
4
0
1
0
0
0
0


C.
megaceph. C. livida


C. vicina
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0








80



sample # P. C. C. P. C.
PIG: DATE: A, R # adults coerul. macellaria ruffifaces regina megaceph. C. livida C. vicina
16E 4/28/2003 A 369 11 4 6 0 1 0 0 0
16E 4/28/2003 R 370 33 33 0 0 0 0 0 0
16E 4/29/2003 A 378 0 0 0 0 0 0 0 0
16E 4/29/2003 R 379 32 32 0 0 0 0 0 0
16E 4/30/2003 R 386 26 4 18 0 4 0 0 0
17B 6/12/2003 A 396 16 8 8 0 0 0 0 0
17B 6/13/2003 A 399 23 22 1 0 0 0 0 0
17B 6/14/2003 A 400 30 0 20 0 10 0 0 0
17B 6/14/2003 R 401 10 1 9 0 0 0 0 0
17B 6/15/2003 R 419 20 0 14 6 0 0 0 0
17B 6/15/2003 A 421 13 0 13 0 0 0 0 0
17C 6/12/2003 A 394 46 27 19 0 0 0 0 0
17C 6/13/2003 A 398 0 0 0 0 0 0 0 0
17C 6/14/2003 R 403 0 0 0 0 0 0 0 0
17C 6/14/2003 A 405 18 6 10 2 0 0 0 0
17C 6/15/2003 R 414 5 0 1 4 0 0 0 0
17C 6/15/2003 A 416 12 0 10 2 0 0 0 0
17C 6/15/2003 R 417 32 0 0 32 0 0 0 0
17E 6/12/2003 A 395 31 19 11 1 0 0 0 0
17E 6/13/2003 A 397 0 0 0 0 0 0 0 0
17E 6/13/2003 R 401 21 7 14 0 0 0 0 0
17E 6/14/2003 R 406 28 0 28 0 0 0 0 0
17E 6/14/2003 A 408 13 0 13 0 0 0 0 0
17E 6/15/2003 R 409 24 0 24 0 0 0 0 0
17E 6/15/2003 A 411 19 0 19 0 0 0 0 0
17E 6/15/2003 R 412 10 1 7 2 0 0 0 0
18A 12/8/2003 A 424 23 23 0 0 0 0 0 0
18A 12/10/2003 A 428 10 10 0 0 0 0 0 0
18A 12/13/2003 R 439 9 9 0 0 0 0 0 0
18A 12/13/2003 A 441 14 14 0 0 0 0 0 0
18A 12/14/2003 R 448 22 22 0 0 0 0 0 0
18A 12/14/2003 A 450 4 4 0 0 0 0 0 0
18A 12/16/2003 R 457 24 24 0 0 0 0 0 0
18A 12/16/2003 A 459 9 7 0 0 2 0 0 0
18A 12/20/2003 R 464 23 21 0 0 2 0 0 0
18B 12/8/2003 A 422 10 10 0 0 0 0 0 0
18B 12/10/2003 A 427 13 13 0 0 0 0 0 0
18B 12/11/2003 A 429 2 0 0 0 0 0 2 0
18B 12/12/2003 A 430 6 5 0 0 1 0 0 0
18B 12/12/2003 R 432 23 23 0 0 0 0 0 0
18B 12/13/2003 A 436 20 17 0 0 3 0 0 0
18B 12/13/2003 R 437 16 16 0 0 0 0 0 0
18B 12/14/2003 R 445 25 25 0 0 0 0 0 0
18B 12/14/2003 A 447 4 2 0 1 1 0 0 0
18B 12/16/2003 A 454 8 5 0 1 2 0 0 0
18B 12/16/2003 R 455 25 25 0 0 0 0 0 0
18C 12/8/2003 A 423 9 9 0 0 0 0 0 0
18C 12/10/2003 A 426 9 9 0 0 0 0 0 0
18C 12/12/2003 A 425 0 0 0 0 0 0 0 0












sample #
PIG: DATE: A, R # adults
18C 12/13/2003 A 433 14
18C 12/13/2003 R 434 20
18C 12/14/2003 R 442 20
18C 12/14/2003 A 444 16
18C 12/16/2003 A 451 15
18C 12/16/2003 R 452 20
18C 12/17/2003 R 460 19
18C 12/17/2003 R 462 20
19A 1/23/2004 A 468 14
19A 1/25/2004 A 469 19
19A 1/26/2004 A 474 16
19A 1/27/2004 R 476 0
19A 1/27/2004 A 477 21
19A 1/28/2004 R 490 31
19A 1/29/2004 R 492 27
19A 1/29/2004 A 494 2
19B 1/24/2004 A 467 7
19B 1/25/2004 A 470 22
19B 1/26/2004 A 473 27
19B 1/27/2004 R 478 45
19B 1/27/2004 A 480 15
19B 1/28/2004 R 487 32
19B 1/28/2004 A 489 1
19B 1/29/2004 R 495 37
19B 1/30/2004 A 502 3
19B 1/31/2004 A 507 1
19C 1/24/2004 A 466 11
19C 1/25/2004 A 471 0
19C 1/26/2004 A 472 42
19C 1/27/2004 R 482 41
19C 1/27/2004 A 483 8
19C 1/28/2004 R 484 55
19C 1/28/2004 A 486 2
19C 1/29/2004 R 497 26
19C 1/30/2004 A 500 8
19C 1/31/2004 A 505 1
20A 3/6/2004 A 512 31
20A 3/7/2004 A 516 21
20A 3/8/2004 A 518 17
20A 3/8/2004 R 519 47
20A 3/9/2004 A 530 4
20A 3/9/2004 R 531 14
20A 3/14/2004 A 548 0
20B 3/5/2004 A 536 1
20B 3/6/2004 A 511 17
20B 3/7/2004 A 515 22
20B 3/8/2004 A 521 14
20B 3/8/2004 R 523 0
20B 3/9/2004 A 527 7


P.
coerul.
13
20
20
13
13
11
19
20
14
19
16
0
19
31
27
1
6
22
25
45
15
30
0
36
2
1
11
0
39
41
4
42
1
25
7
1
29
18
4
47
1
2
0
1
12
6
0
0
0


C.
macellaria
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
1
0
0


C.
ruffifaces
1
0
0
1
0
9
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0


P.
regina
0
0
0
1
2
0
0
0
0
0
0
0
2
0
0
1
0
0
1
0
0
2
1
1
1
0
0
0
2
0
4
13
0
1
1
0
2
3
13
0
3
12
0
0
5
14
13
0
7


C.
megaceph. C. livida


C. vicina
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0













PIG: DATE: A, R
20B 3/9/2004 R
20B 3/10/2004 R
20B 3/13/2004 A
20B 3/14/2004 A
20B 3/14/2004 R


sample
#
528
537
543
545
547


# P. C. C. P. C.
adults coerul. macellaria ruffifaces regina megaceph. C. livida C. vicina
47 1 0 0 46 0 0 0
35 0 0 0 35 0 0 0
8 3 3 0 2 0 0 0
23 1 5 0 17 0 0 0
6 0 0 1 5 0 0 0




















APPENDIX B
RAW DATA: PRESERVED SPECIMENS


sample 1st
PIG: DATE: # instar
2B 11/21/2001 10B 0
3A 12/29/2001 11A 0
3A 1/11/2002 14 0
3B 12/29/2001 12A 0
3B 1/11/2002 18 0
3C 1/11/2002 16 0
4E 2/9/2002 29 19
5A 3/17/2002 34 66
5A 3/18/2002 46 0
5B 3/17/2002 39 82
5C 3/17/2002 37 34
5C 3/18/2002 43 254
5E 3/17/2002 41 82
5E 3/18/2002 45 0
5E 3/19/2002 48 0
6A 5/1/2002 54 17
6A 5/1/2002 58 1
6B 5/1/2002 58 0
6B 5/1/2002 60 174
6C 5/1/2002 62 5
6C 5/1/2002 64 1
6E 5/1/2002 67 12
6E 5/1/2002 69 78
6E 5/1/2002 71 0
7A 5/23/2002 79 1
8A 7/23/2002 88 138
8A 7/24/2002 93 2
8A 7/24/2002 95 51
8A 7/24/2002 96 8
8A 7/25/2002 106 0
8A 7/25/2002 108 0
8C 7/23/3003 90 0
8C 7/24/2002 104 9
8C 7/24/2002 105 15
8E 7/23/2002 89 10
8E 7/24/2002 99 14
8E 7/24/2002 100 400
8E 7/24/2002 101 43
8E 7/25/2002 110 0


1-2
trans.
0
0
0
0
0
0
4
17
0
35
3
52
10
0
2
8
2
0
1
2
0
5
5
0
0
7
0
20
1
0
0
0
1
1
4
1
53
14
0


2nd
instar
0
0
42
0
0
8
64
56
17
80
129
16
24
88
61
73
75
0
0
59
14
73
39
17
19
0
31
50
89
14
47
0
76
41
40
111
49
28
3


2-3
trans.
0
0
11
0
1
2
16
14
19
2
5
0
0
6
12
10
8
0
0
48
7
0
10
4
1
0
6
0
8
12
8
0
40
28
0
6
0
1
2


3rd P.
instar total coerul.
0 0 0
0 0 0
219 272 108
90 90 90
152 153 148
107 117 107
151 254 107
0 153 0
79 115 76
0 199 0
14 185 14
0 322 0
0 116 0
7 101 7
34 109 34
13 121 13
59 145 59
0 0 0
0 175 0
10 124 10
63 85 63
10 100 10
8 140 8
33 54 33
175 196 175
0 145 0
65 104 65
1 122 1
2 108 1
164 190 0
241 296 0
0 0 0
77 203 77
54 139 54
0 54 0
16 148 15
0 502 0
10 96 8
70 75 0


C. C.
mac. ruf.
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 1
0 164
0 241
0 0
0 0
0 0
0 0
0 1
0 0
0 2
0 70


P. ?
regina 3rd
0 0
0 0
98 13
0 0
4 0
0 0
44 0
0 0
0 3
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0












sample
PIG: DATE: #
9A 8/21/2002 120
9A 8/21/2002 120B
9A 8/21/2002 121
9A 8/21/2002 122
9A 8/22/2002 155
9A 8/22/2002 156
9A 8/22/2002 158
9B 8/21/2002 126
9B 8/21/2002 126B
9B 8/21/2002 127
9B 8/21/2002 128
9B 8/22/2002 150
9B 8/22/2003 152
9C 8/22/2002 136
9C 8/22/2002 138
9C 8/22/2002 139
9C 8/22/2002 140
9C 8/22/2002 141
9E 8/21/2002 131
9E 8/21/2002 131 B
9E 8/21/2002 132
9E 8/21/2002 133
9E 8/22/2002 144
9E 8/22/2002 146
9E 8/22/2002 147
10A 9/25/2002 164
10A 9/26/2002 179
10A 9/26/2002 181
10B 9/25/2002 168
10B 9/25/2002 169
10B 9/26/2002 182
10B 9/26/2002 184
10B 9/26/2002 186
10C 9/25/2002 176
10C 9/27/2002 189
10E 9/25/2002 172
10E 9/25/2002 173
10E 9/27/2002 190
10E 9/27/2002 192
11A 10/26/2002 194
11A 10/27/2002 202
11A 10/27/2002 204
11A 10/28/2002 211
11A 10/28/2002 213
11B 10/26/2002 197
11C 10/28/2002 207
12A 12/7/2002 220
12A 12/8/2002 229
12A 12/12/2002 254


1st
instar
2
6
102
108
4
8
0
29
16
183
12
0
13
3
0
9
116
0
0
0
0
34
3
1
0
0
0
33
1
3
15
0
89
0
0
0
3
0
0
0
0
94
0
0
170
0
1
0
0


1-2
trans.
1
2
23
3
6
0
0
2
2
0
0
0
2
0
0
2
17
0
0
0
0
4
3
0
0
0
0
8
3
11
1
0
20
0
0
0
2
0
0
0
1
28
0
0
13
0
3
0
4


2nd
instar
126
115
55
45
52
1
7
39
42
0
49
6
36
8
2
3
23
11
8
15
0
111
42
61
19
20
0
186
50
189
8
0
87
45
29
35
32
0
30
0
7
31
14
28
56
1
111
86
52


2-3
trans.
7
15
0
0
5
3
1
24
18
0
12
6
22
10
1
3
9
4
3
1
0
21
7
9
8
49
0
30
74
17
0
0
0
20
24
17
7
0
7
0
24
30
14
48
2
0
45
5
48


3rd P.
instar total coerul.
17 153 17
11 149 11
0 180 0
2 158 2
34 101 34
14 26 1
49 57 48
17 111 17
10 88 9
0 183 0
2 75 2
54 66 53
1 74 0
32 53 5
40 43 33
9 26 4
18 183 18
40 55 38
31 42 31
29 45 24
0 0 0
11 181 11
17 72 13
20 91 2
41 68 41
23 92 23
137 137 135
32 289 32
25 153 25
0 220 0
73 97 71
8 8 0
1 197 1
9 74 9
88 141 43
35 87 35
86 130 86
72 72 0
108 145 1
0 0 0
82 114 54
14 197 13
98 126 98
22 98 0
9 250 9
69 70 0
62 222 62
33 124 33
52 156 52


C. C.
mac. ruf.
0 0
0 0
0 0
0 0
0 0
0 13
0 0
0 0
0 0
0 0
0 0
0 0
0 1
0 27
0 0
0 5
0 0
0 1
0 0
0 3
0 0
0 0
2 0
0 18
0 0
0 0
2 0
0 0
0 0
0 0
0 2
3 5
0 0
0 0
43 0
0 0
0 0
0 72
102 0
0 0
4 24
0 1
0 0
22 0
0 0
0 69
0 0
0 0
0 0


P. ?
regina 3rd
0 0
0 0
0 0
0 0
0 0
0 0
1 0
0 0
1 0
0 0
0 0
1 0
0 0
0 0
7 0
0 0
0 0
1 0
0 0
2 0
0 0
0 0
1 1
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
2 0
0 0
0 0
0 0
5 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0












sample
PIG: DATE: #
12A 12/13/2002 256
12A 12/13/2002 257
12A 12/14/2002 259
12B 11/30/2002 219
12B 12/7/2002 223
12B 12/8/2002 232
12B 12/10/2002 242
12B 12/10/2002 243
12B 12/10/2002 244
12B 12/12/2002 250
12B 12/12/2002 252
12C 12/7/2002 227
12C 12/10/2002 236
12C 12/10/2002 237
12C 12/10/2002 238
12C 12/12/2002 246
12E 12/7/2002 225
12E 12/10/2002 241
12E 12/12/2002 248
13B 1/3/2003 267
13B 1/4/2003 277
13B 1/5/2003 284
13B 1/9/2003 293
13C 1/3/2003 271
13C 1/4/2003 275
13C 1/5/2003 283
13C 1/8/2003 291
13C 1/9/2003 299
13C 1/11/2003 301
13E 1/3/2003 269
13E 1/4/2003 273
13E 1/5/2003 280
13E 1/6/2003 287
13E 1/8/2003 289
13E 1/9/2003 296
14A 3/6/2003 305
14A 3/6/2003 307
14A 3/8/2003 315
14A 3/8/2003 317
14B 3/5/2003 303
14E 3/7/2003 310
14E 3/7/2003 313
15A 4/5/2003 341
15A 4/6/2003 354
15B 4/4/2003 332
15B 4/5/2003 344
15C 4/4/2003 328
15C 4/4/2003 329
15C 4/5/2003 338


1st
instar
0
0
0
0
3
0
7
0
1
3
0
0
22
0
0
2
5
89
0
0
0
2
2
12
0
0
0
0
3
2
21
0
0
0
0
0
44
0
0
0
2
5
1
0
32
105
60
0
15


1-2
trans.
0
0
0
0
2
0
4
0
0
3
6
0
5
0
0
1
8
5
0
0
0
1
1
5
1
0
0
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
6
0
5
0
1


2nd
instar
4
10
10
0
126
59
46
1
3
47
109
55
53
9
6
10
161
24
13
0
0
109
19
76
29
72
6
38
84
69
75
12
0
23
3
2
263
8
2
10
73
60
0
1
130
89
92
0
102


2-3
trans.
0
6
5
0
23
2
14
1
4
12
50
17
16
1
2
0
11
17
1
0
0
2
6
6
25
16
1
5
7
15
46
3
1
37
0
1
4
0
0
2
4
6
11
1
13
8
0
0
1


3rd P.
instar total coerul.
85 89 85
61 77 61
99 114 99
0 0 0
6 160 6
25 86 25
149 220 146
111 113 111
135 143 135
22 87 22
24 189 24
33 105 33
27 123 27
56 66 56
103 111 103
24 37 24
5 190 5
34 169 34
102 116 102
0 0 0
0 0 0
0 114 0
69 97 69
1 100 0
51 106 51
33 121 33
101 108 101
42 85 42
3 97 2
8 94 8
28 173 28
61 76 61
83 84 83
47 107 42
54 57 54
71 74 0
7 318 6
97 105 97
47 49 47
52 64 27
37 116 37
29 100 29
27 39 27
66 68 49
0 181 0
4 206 4
0 157 0
0 0 0
7 126 7


C. C.
mac. ruf.
0 0
0 0
0 0
0 0
0 0
0 0
3 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
1 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0


P. ?
regina 3rd
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 1
0 0
0 0
0 0
0 0
0 5
0 0
71 0
0 1
0 0
0 0
25 0
0 0
0 0
0 0
17 0
0 0
0 0
0 0
0 0
0 0












sample
PIG: DATE: #
15C 4/6/2003 351
15E 4/4/2003 335
15E 4/5/2003 347
15E 4/6/2003 356
16B 4/28/2003 374
16B 4/29/2003 382
16C 4/28/2003 368
16C 4/29/2003 377
16C 4/30/2003 385
16C 5/1/2003 390
16C 5/1/2003 391
16C 5/1/2003 392
16E 4/27/2003 364
16E 4/28/2003 371
16E 4/29/2003 380
16E 4/30/2003 387
17B 6/14/2003 402
17B 6/15/2003 420
17C 6/14/2003 404
17C 6/15/2003 415
17C 6/15/2003 418
17E 6/13/2003 400A
17E 6/14/2003 407
17E 6/15/2003 410
17E 6/15/2003 413
18A 12/13/2003 440
18A 12/14/2003 449
18A 12/16/2003 458
18A 12/20/2003 465
18B 12/12/2003 431
18B 12/13/2003 438
18B 12/14/2003 446
18B 12/16/2003 456
18C 12/13/2003 435
18C 12/14/2003 443
18C 12/16/2003 453
18C 12/17/2003 461
18C 12/17/2004 463
19A 1/27/2004 475
19A 1/28/2004 491
19A 1/29/2004 493
19A 1/30/2004 503
19A 1/31/2004 508
19B 1/27/2004 479
19B 1/28/2004 488
19B 1/29/2004 496
19B 1/30/2004 501
19B 1/31/2004 506
19C 1/27/2004 481


1st
instar
1
44
5
0
54
0
1
0
0
0
0
0
28
0
0
0
99
0
192
0
0
153
8
0
0
50
28
4
1
129
0
4
0
10
2
0
0
0
122
12
4
0
0
224
107
46
10
0
88


1-2
trans.
0
10
0
0
10
0
0
0
0
0
0
0
9
0
2
0
17
1
0
0
0
6
3
0
0
10
0
0
8
21
0
0
0
2
0
0
0
0
12
0
0
0
0
26
0
2
0
0
0


2nd
instar
20
122
30
0
15
106
59
63
4
0
0
0
13
34
2
1
113
12
9
1
6
3
73
77
14
88
270
121
31
9
68
84
92
178
0
19
0
15
110
138
68
11
2
44
3
128
70
23
0


2-3
trans.
2
8
4
0
0
0
6
0
0
0
1
0
0
0
1
0
13
3
0
1
0
0
3
1
4
2
2
6
0
3
0
20
8
4
8
2
0
2
0
6
0
0
1
0
0
6
12
1
0


3rd P.
instar total coerul.
70 93 70
2 186 0
67 106 67
58 58 10
0 79 0
104 210 104
10 76 6
32 95 23
105 109 0
67 67 0
61 62 2
41 41 0
0 50 0
43 77 43
134 139 134
54 55 17
22 264 22
44 60 44
40 241 40
29 31 7
75 81 11
0 162 0
73 160 73
45 123 35
60 78 54
0 150 0
4 304 2
14 145 10
52 92 50
0 162 0
2 70 2
36 144 12
85 185 77
0 194 0
192 202 176
94 115 88
70 70 70
99 116 99
0 244 0
18 174 4
38 110 24
57 68 57
58 61 58
0 294 0
0 110 0
12 194 0
38 130 20
61 85 61
0 88 0


C. C.
mac. ruf.
0 0
0 0
0 0
25 0
0 0
0 0
3 0
9 0
105 0
67 0
58 0
41 0
0 0
0 0
0 0
29 0
0 0
0 0
0 0
4 18
64 0
0 0
0 0
2 3
5 0
0 0
0 0
0 0
0 2
0 0
0 0
0 0
0 0
0 0
0 0
0 6
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0


P. ?
regina 3rd
0 0
2 0
0 0
23 0
0 0
0 0
1 0
0 0
0 0
0 0
1 0
0 0
0 0
0 0
0 0
7 1
0 0
0 0
0 0
0 0
0 0
0 0
0 0
5 0
0 1
0 0
2 0
4 0
0 0
0 0
0 0
24 0
8 0
0 0
16 0
0 0
0 0
0 0
0 0
14 0
14 0
0 0
0 0
0 0
0 0
12 0
18 0
0 0
0 0








87


sample 1st 1-2 2nd 2-3 3rd P. C. C. P. ?
PIG: DATE: # instar trans. instar trans. instar total coerul. mac. ruf. regina 3rd
19C 1/28/2004 485 76 0 74 0 0 150 0 0 0 0 0
19C 1/29/2004 498 48 0 150 10 2 210 0 0 0 2 0
19C 1/30/2004 499 0 0 56 1 17 74 17 0 0 0 0
19C 1/31/2004 504 0 0 70 4 46 120 46 0 0 0 0
20A 3/8/2004 520 0 0 138 4 16 158 14 0 0 2 0
20A 3/9/2004 532 13 0 67 2 45 127 38 0 0 7 0
20B 3/8/2004 522 0 0 280 20 28 328 28 0 0 0 0
20B 3/9/2004 529 24 0 70 8 22 124 22 0 0 0 0
20B 3/10/2004 538 0 0 20 0 68 90 64 2 0 2 0
20B 3/13/2004 544 0 0 10 0 61 71 0 4 0 57 0
20B 3/14/2004 546 0 0 0 1 65 66 1 3 0 61 0
















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