Protophormia terraenovae

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Protophormia terraenovae
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Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Family: Calliphoridae
Genus: Protophormia
Species:
P. terraenovae
Binomial name
Protophormia terraenovae
Synonyms

Protophormia terraenovae is commonly called northern blowfly, blue-bottle fly or blue-assed fly (blue-arsed fly in British English). It is distinguished by its deep blue coloration and large size and is an important species throughout the Northern Hemisphere. This fly is notable for its economic effect as a myiasis pest of livestock and its antibiotic benefits in maggot therapy. Also of interest is P. terraenovae’s importance in forensic investigations: because of their temperature-dependent development and their prominent presence on corpses, the larvae of this species are useful in minimum post-mortem interval (mPMI) determination.

Contents

Taxonomy

Protophormia terraenovae, of the family Calliphoridae, was named and first described by French entomologist André Jean Baptiste Robineau-Desvoidy in his 1830 “Essai sur les myodaires.” Its genus is shared by one other fly, Protophormia atriceps. Both flies are a dark, undusted, metallic blue-green-black. P. terraenovae is differentiated from P. atriceps by its flat face, plumose arista, and by up to 2 additional pairs of setae along the margin of the scutellum. [3] P. terraenovae’s specific epithet is translated from the Latin as “of the New World.”

The oldest-known specimens of P. terraenovae pupae were identified in 1973 within the fossilized skull of a steppe wisent. The bison skull, estimated to date from the late Eemian period, was excavated from the site of a new sluice for the Brussels-Rupel Canal in Zemst, Belgium. The specimens, although approximately 75,000 years old, are identical in form to the pupae of the modern species. [4]

Morphology

Adult

The adult Protophormia terraenovae, one of the larger species of calliphorids, measures between 7 and 12 mm in length. [5] It is characterized by a black to brown anterior thoracic spiracle, [6] a black basicosta, brown wing veins, and dark calypters. Upper calypters sprout black setae. While the thorax and abdomen of P. terraenovae can range in color from dark purple to dark green, the head and legs of this fly are black. Postocular setae are short but prominent; the palpi are yellow or dusky brown. [7]

Sexes of the species are easily distinguished by the width of the space between the eyes, the frons: that of the female is .386 times the width of the head, while the frons of the male is only .140 times the head width. Chaetotaxy, the study of setae arrangement, is also useful for determination of sex—for example, the male lacks the fronto-orbital, lateroclinate setae found in the female. These setae are located just at the edge of the frons, near the upper portion of the complex eyes. [7]

Larvae

Protophormia terraenovae larvae are tiny and white, with 12 segments. Length varies by larval age, with the first instar growing up to 2.63 mm and the third, by contrast, as long as 11.87 mm. In the third instar, the last segment is adorned with pointed tubercles, and segment 10 exhibits dorsal spines along its posterior margin. The latter characteristic distinguishes P. terraenovae, 3rd instar, from a similar calliphorid, Phormia regina . [8]

Development

Being of the order Diptera, Protophormia terraenovae is holometabolous in its development, meaning it experiences dramatic changes from immature to adult. Protophormia terraenovae undergoes three instars, a pupal stage, and finally an adult stage.

First instar larvae obtain a liquid diet from orifices or wounds of a body. The instars that follow digest the body itself. Among the first to colonize, Calliphoridae species are found on the body almost immediately. As eggs hatch into the first larval stage, P. terraenovae begins feeding and increases in size, limited by its chitinous outer cuticle. As P. terraenovae larvae molt into the second instar, feeding intensifies: with larger and more developed mouthparts, second instars are able to break down tougher body tissues. [9]

The third instar is unique, consisting of several substages. First, the larvae feed until they reach their maximum size. Next, larvae clear their crops. In this prepupal stage, also called the wandering stage, the maggots seek a dry, safe area for pupation. During the pupal stage, the outer cuticle hardens and tans into a dark brown color, gradually darkening with age. Finally, metamorphosis into the adult is completed with the shedding of the pupal casing and the emergence of the fly. [9]

The time of development between instars to pupae to adult differs depending on temperatures. The mean duration of development, at constant pressure and 25 °C, between oviposition and hatching is 1.83 days. From first to second instar, approximately 2.92 days pass. About 6.17 days pass between the second and third instar, and 8.0 days between the third instar and the post-feeding stage. The wandering maggot takes around 9.67 days to reach the pupal stage. Lastly, emergence of this blowfly occurs about 15.83 days after pupation. [9] In fluctuating conditions such as those around crime scenes, there is a tendency for development to slow down. Under-approximation of age is therefore a common mistake. [10]

The development of P. terraenovae is inversely related to temperature. Studies at (15, 20, 25, 30, 35)°C were performed, and the minimal duration of development from oviposition to adult ranges from 9.19 ± .3 days at 35 °C, to 37.78 ± 2.96 days at 15 °C. The minimum development threshold for total immature development is 8.95 °C. The overall thermal constant (K) for P. terraenovae is 240.2 ± 9.3 day-degrees above threshold. The inverse relationship and regression of development from oviposition to pupation resulted in a 9.8 °C threshold for the species. [11]

Pupation occurs on or within .5 meters of the decaying body. The duration of the pupal stage is relatively long compared to the rest of development, inactively occupying 43% of P. terraenovae’s total cycle. The third instar larva spends 13% of its cycle in feeding stage and 22% in post-feeding stages. [12]

Geographic distribution

Protophormia terraenovae has a Holarctic distribution, meaning the species is found throughout the northern hemisphere. The fly is common in cool regions and, being "the most cold tolerant of all calliphorid species," can withstand extreme temperatures. P. terraenovae can be found as close as 550 miles from the North Pole and is abundantly found in the Arctic. [13]

In North America, P. terraenovae can be found from Mexico to Canada. Appearance of this species is rare and mostly confined to the winter months in warm regions such as Texas and Florida. P. terraenovae is a spring or summer species in high latitudes and elevations. [5]

Importance

Medical

Maggot debridement therapy (MDT) is commonly used for the cleansing and disinfecting of chronic wounds containing necrotic flesh. Various studies have shown that MDT is effective in treating wounds that have failed to heal. Effective MDT species consume necrotic tissue while cleansing the wound. [14] Protophormia terraenovae is among the few blowfly species that fit these criteria. The larvae of most blowflies are necrophagous, meaning they develop in the bodies of dead vertebrates where they consume necrotic tissues. Because the majority of other blowfly larvae consume both necrotic and healthy tissues, P. terraenovae is an important species. [14] P. terraenovae is also known to produce antibiotics during feeding: the secretions of P. terraenovae larvae are effective in fighting infections involving Streptococcus pyogenes and Streptococcus pneumoniae . [15]

The main concern with the use of MDT is sepsis. P. terraenovae is blamed for five reported bloodstream infections. [16] To prevent infection, P. terraenovae maggots must be raised in vitro under sterile conditions. Provided these precautions are taken, the use of P. terraenovae on chronic wounds appears to remain safe and efficient in wound treatment.

Economic

As a species of the blowfly family Calliphoridae, Protophormia terraenovae causes economically important myiasis in livestock and, occasionally, in humans. It also affects other populations of Diptera due to the predacious nature of its larvae when competing for necrotic tissue. P. terraenovae larvae have also been known to behave cannibalistically. [17]

Myiasis due to Protophormia terraenovae has been reported in both wild and domestic animals, but it most commonly affects livestock. Species usually oviposit on carrion, but are capable of oviposition in the wounds of livestock and wild animals. Thus, P. terraenovae acts as a secondary invader in causing myiasis. [18] Commonly referred to as ‘wound strike’, lesions are foul smelling ulcers that are filled with larvae. Animals infected with myiasis may die from toxemia and sepsis. [19]

In particular, Protophormia terraenovae causes facultative, cutaneous myiasis of cattle, sheep and reindeer in the northern Holarctic region. [6] Larvae feeding on the skin of sheep causes distress to the animal and loss of wool. [20] This skin damage occurs to a lesser extent in horses, goats and pigs. Adults of this species are capable of causing myiasis but are found only in the Palaearctic and Nearctic regions. [6]

Also of economic significance are manifestations of the species in slaughter-houses and poultry houses. Populations of P. terraenovae are maintained by refuse and the carcasses of dead animals. Due to their predacious nature, large numbers of this species can populate in the presence of other Dipteran species. Blowfly bred at these locations cause considerable local nuisance and may spread disease to both humans and animals by contaminating meat and foodstuffs. [17]

Forensic

Medicocriminal entomology is the branch of forensic entomology dealing with the use of arthropod evidence in criminal investigations. Because they make up the first wave of fauna to colonize a corpse, blowflies are among the most accurate forensic indicators of time elapsed since death, technically referred to as the post-mortem interval (PMI). This estimation is made by determining the developmental stages of the insects present on a body. Protophormia terraenovae is forensically important because of its extremely specific developmental time. [11] Two methods are used to determine the PMI of a P. terraenovae-infested body. One approach utilizes information about the developmental stages of larvae located on or within the body, and the second utilizes facts about the larval stage as it emerges from the body to pupate.

The first, second, and third instar larvae present on a body are considered to be feeding stages. The first and second molting are characterized by rapid increases in size and generally occur within a certain range of larval length. Because the development of these flies is dependent on temperature, entomologists can use past weather data to estimate the ages of larvae and even pupae discovered on a corpse. [11]

When post-feeding larvae venture away from the body to pupate, as is common for most blowflies, displacement behavior becomes the determining factor of age. [21] When a body is discovered, if the larvae are leaving, it is possible to measure the distance they have traveled and, with knowledge of the species' crawling speed, to calculate when they left the body. Crawling speed is affected by terrain, temperature, and maggot length. These considerations allow forensic entomologists to make precise age estimations. [21]

Another noteworthy forensic consideration is the use of blowfly species, in this case Protophormia terraenovae, in entomotoxicology. Evidence of drug use can be found in blowflies feeding upon cadavers with post-mortem drugs in their systems. Studies have shown that unlike other species of necrophagous Diptera, blowfly species can indicate the presence of morphine in the cuticle during larval growth, and, definitively, in pupa casings. Because puparial cases decay at an extremely slow rate and can be recovered years after the death of an individual, they can become very important to the examination of a corpse. [22]

The "Pig Farm Case"

In 2007, entomological evidence was used in the infamous “Pig Farm Case”. An eighteen-month-long search of several acres of farmland and property in British Columbia resulted in the discovery of trace DNA evidence linked to twenty-six missing women. The heads, hands, and feet of two of the missing women were eventually found. Protophormia terraenovae larvae were prominent on these remains. Forensic entomologists determined developmental rates based on locally collected specimens of P. terraenovae, and with this information, concluded that both women had been exposed for weeks prior to being frozen. The pig farmer and co-owner of the grounds, Robert “Willie” Pickton, was charged with the murders of all twenty-six women. In December 2007 Pickton was convicted of second-degree murder in the deaths of six women and stands accused of first-degree murder in the deaths of twenty other women. [23]

Current and future research

Investigation of the species is still ongoing: specifically, researchers are examining PMI and ADH/ADD for P. terraenovae, ecdysteroid levels and molting, and adult diapause with reactions to certain temperatures. [24]

Microlesions were made inside the brain of Protophormia terraenovae to study the diapause factor. Results show that secretory neurons are necessary for entering the reproductive diapause. Other research based on the haemolymph circulation rate suggests that sensory-induced changes can result in a series of gustatory stimulations that shorten the cardiac cycle of P. terraenovae.

Related Research Articles

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The Calliphoridae are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing bait, are known as gentles. The family is known to be polyphyletic, but much remains disputed regarding proper treatment of the constituent taxa, some of which are occasionally accorded family status.

<span class="mw-page-title-main">Common green bottle fly</span> Species of insect

The common green bottle fly is a blowfly found in most areas of the world and is the most well-known of the numerous green bottle fly species. Its body is 10–14 mm (0.39–0.55 in) in length – slightly larger than a house fly – and has brilliant, metallic, blue-green or golden coloration with black markings. It has short, sparse, black bristles (setae) and three cross-grooves on the thorax. The wings are clear with light brown veins, and the legs and antennae are black. The larvae of the fly may be used for maggot therapy, are commonly used in forensic entomology, and can be the cause of myiasis in livestock and pets. The common green bottle fly emerges in the spring for mating.

<i>Chrysomya putoria</i> Species of fly

Chrysomya putoria, also known as the tropical African latrine blowfly, is a fly species belonging to the blowfly family, Calliphoridae.C. putoria is native to Africa and has recently spread to the Americas. These flies pose significant health risks, especially due to their close association with human settlements. Adult flies can carry pathogens, while larvae may cause myiasis by growing and feeding on the flesh of domestic animals and humans. Other myiasis-causing flies in the same genus are C. bezziana and C. megacephala. C. putoria and other flies that feed on decomposing tissue are used as important tools in forensic entomology to establish the post-mortem interval, or the time elapsed since death.

<i>Cynomya mortuorum</i> Species of fly

Cynomya mortuorum belongs to the order Diptera, sometimes referred to as "true flies". In English, the only common name occasionally used is "fly of the dead". It has a bluish-green appearance, similar to other Calliphoridae and is found in multiple geographic locations with a preference for colder regions. Belonging to the family Calliphoridae, it has been shown to have forensically relevant implications due to its appearance on carrion. Current research is being done to determine C. mortuorum's level of importance and usage within forensic entomology.

<i>Calliphora vomitoria</i> Species of fly

Calliphora vomitoria, known as the blue bottle fly, orange-bearded blue bottle, or bottlebee is a species of blow fly, a species in the family Calliphoridae. Calliphora vomitoria is the type species of the genus Calliphora. It is common throughout many continents including Europe, Americas, and Africa. They are fairly large flies, nearly twice the size of the housefly, with a metallic blue abdomen and long orange setae on the gena.

<i>Chrysomya</i> Genus of flies

Chrysomya is an Old World blow fly genus of the family Calliphoridae. The genus Chrysomya contains a number of species including Chrysomya rufifacies and Chrysomya megacephala. The term “Old World blow fly” is a derivative of both the associated family, Calliphoridae, and the belief that the genus Chrysomya originated in Asia and migrated to North America only relatively recently. Chrysomya’s primary importance to the field of medico-criminal forensic entomology is due to the genus’ reliable life cycle, allowing investigators to accurately develop a postmortem interval. Chrysomya adults are typically metallic colored with thick setae on the meron and plumose arista. The name comes from the word chrysos, meaning “golden” in reference to the metallic sheen of the genus’ species, and -mya, a derivation from the word myia, meaning “fly”.

<i>Chrysomya rufifacies</i> Species of fly

Chrysomya rufifacies is a species belonging to the blow fly family, Calliphoridae, and is most significant in the field of forensic entomology due to its use in establishing or altering post mortem intervals. The common name for the species is the hairy maggot blow fly, and it belongs to the genus Chrysomya, which is commonly referred to as the Old World screwworms. This genus includes other species such as Chrysomya putoria and Chrysomya bezziana, which are agents of myiasis. C. rufifacies prefers very warm weather and has a relatively short lifecycle. It is widely distributed geographically and prefers to colonize large carcasses over small ones. The species commonly has a greenish metallic appearance and is important medically, economically, and forensically.

<i>Lucilia illustris</i> Species of insect

Lucilia illustris is a member of the fly family Calliphoridae, commonly known as a blow fly. Along with several other species, L. illustris is commonly referred to as a green bottle fly. Lucilia illustris is typically 6–9 mm in length and has a metallic blue-green thorax. The larvae develop in three instars, each with unique developmental properties. The adult fly typically will feed on flowers, but the females need some sort of carrion protein in order to breed and lay eggs.

Insect development during storage requires special consideration when further criminal investigation is necessary to solve a crime. Decomposition is a natural process of the body, dissipating slowly over time. This process is aided by insects, making the rate of decomposition faster. For forensic entomologists, it is important to carefully collect, preserve and analyze insects found near or on a victim. By doing that, they can provide an estimated time of death as well as the manner of death and the movement of the corpse from one site to another. The role of a forensic entomologist adjunction to the pathologist is to “collect and identify the arthropods associated with such cases and to analyze entomological data for interpreting insect evidence.”

<i>Calliphora vicina</i> Species of insect

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<i>Phormia regina</i> Species of fly

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<i>Chrysomya megacephala</i> Species of fly

Chrysomya megacephala, more commonly known as the oriental latrine fly or oriental blue fly, is a member of the family Calliphoridae (blowflies). It is a warm-weather fly with a greenish-blue metallic box-like body. The fly infests corpses soon after death, making it important to forensic science. This fly is implicated in some public health issues; it can cause accidental myiasis, and also infects fish and livestock.

<i>Lucilia silvarum</i> Species of fly

The common toad fly, Lucilia silvarum, is a member of the fly family Calliphoridae. This fly was first discovered by Johann Wilhelm Meigen in 1826 and is found most notably in European and Western Countries.

<i>Sarcophaga pernix</i> Species of fly

Sarcophaga pernix, also known as the red-tailed flesh fly, is a fly in the Sarcophagidae family. This fly often breeds in carrion and feces, making it a possible vector for disease. The larvae of this species can cause myiasis, as well as accidental myiasis. It is potentially useful in forensic entomology.

Lucilia thatuna belongs to the family Calliphoridae, the species most commonly referred to as the blowflies, and the genus Lucilia. Along with several other species of Lucilia, L. thatuna is commonly referred to as a green bottle fly. L. thatuna is very scarce and not much is known about this particular fly. It has been noted to reside in mountainous regions of the northwestern United States.

Compsomyiops callipes, previously known as Paraluclia wheeleri, is a member of the blowfly family Calliphoridae. It is a warm weather fly that can be found in southwestern parts of the United States and parts of South America. This species can be identified by its chaeotaxy, metallic blue color, club-shaped palp, and brown calypters.

<i>Cynomya cadaverina</i> Species of fly

Cynomya cadaverina, also known as the shiny blue bottle fly, is a member of the family Calliphoridae, which includes blow flies as well as bottle flies. In recent years, this family has become a forensically important facet in many medicocriminal investigations in the growing field of forensic entomology. C. cadaverina is specifically important in determining a post-mortem interval, as well as other important factors.

<i>Calliphora livida</i> Species of fly

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<i>Lucilia coeruleiviridis</i> Species of fly

Lucilia coeruleiviridis, formerly Phaenecia coeruleiviridis, is commonly known as a green bottle fly, because of its metallic blue-green thorax and abdomen. L. coeruleiviridis was first discovered by French entomologist Pierre-Justin-Marie Macquart in 1855. It belongs to the family Calliphoridae and is one of many forensically important Diptera, as it is often found on decaying substances. L. coeruleiviridis is one of the most ubiquitous blow fly species in the southeastern United States, particularly in the spring and fall months.

Calliphora loewi is part of the family Calliphoridae, bottle flies and blowflies, and in the genus Calliphora, blue bottle flies. The genus can be deceiving since C. loewi is not blue. Though this species is rare, it can play an important part in forensic entomology, spreading disease, and decomposing carrion. The life cycle of C. loewi is similar to the life cycle of the genus Calliphora. Since this species is rare there has not been very much research done with this species.

References

  1. 1 2 3 "Protophormia terraenovae". Integrated Taxonomic Information System . Retrieved 22 March 2009.
  2. 1 2 Chandler, Peter J. (1998). "Checklists of Insects of the British Isles (New Series) Part 1: Diptera". Handbooks for the Identification of British Insects . 2. London: Royal Entomological Society of London. 12 (1): 1–234.
  3. ↑ Whitworth, Terry. "Keys to the Genera and Species of Blow Flies (Diptera: Calliphoridae) of America North of Mexico. Proc. Entomol. Soc. Wash. 108(3). 2006.
  4. ↑ Gautier, A. & H. Schumann. “Puparia of the subarctic or black blowfly Protophormia terraenovae (Robineau-Desvoidy, 1830) in a skull of a late Eemian (?) bison at Zemst, Brabant (Belgium).” Palaeogeogr., Palaeoclimatol., Palaeoecol. 14 (1973): 119-125.
  5. 1 2 Byrd, Jason, and James Castner (2001). Forensic Entomology: The Utility of Arthropods in Legal Investigations, CRC Press, New York. ISBN   0-8493-8120-7.
  6. 1 2 3 Wall, Richard, and David Shearer. Veterinary Entomology: Arthropod ectoparasites of veterinary importance. London: Chapman & Hall, 1997.
  7. 1 2 Rognes, Knut. "Blowflies (Diptera, Calliphoridae) of Fennoscandia and Denmark." Fauna Entomologica Scandinavia 24. Leiden: E.J. Brill/Scandinavian Science Press Ltd. 1991.
  8. ↑ Erzinçliolu, Y. Z. “The larvae of the species of Phormia and Boreellus: Northern, cold-adapted blowflies (Diptera: Calliphoridae).” Journal of Natural History, 22.1(1988):11-16.
  9. 1 2 3 Warren, Jodie-Ann. "The Development of Protophormia terraenovae at Constant and Fluctuating Temperatures. . Simon Fraser University, 1999."
  10. ↑ Clarkson, Colin A., Niki Hobischak, and Gail Anderson. “Developmental rate of Protophormia terraenovae (R-D) raised under constant and fluctuating temperatures, for use in determining time since death in natural outdoor conditions, in the early postmortem interval.” Archived 2011-07-06 at the Wayback Machine 2005.
  11. 1 2 3 Grassberger, Martin, and Christian Reiter. "Effect of Temperature on Development of the Forensically Important Holarctic Blow Fly Protophormia terraenovae (Robineau-Desvoidy) (Diptera: Calliphoridae)." Forensic Science International 128 (2002): 177-182.
  12. ↑ E. Gaudry, C. Blais, A. Maria, C. Dauphin-Villemant. Study of steroidogenesis in pupae of the forensically important blow fly Protophormia terraenovae (Robineau-Desvoidy) (Diptera: Calliphoridae). Forensic Science International. 160.1(2006): 27-34.
  13. ↑ Smith, K.G.V. A Manual of Forensic Entomology. London, British Museum (Natural History). 1986.
  14. 1 2 Kočišová, A., J. Pistl, R. Link, E. Čonková, and M. Goldová. "Maggot Debridement Therapy in the Treatment of Footrot and Foot Scald in Sheep." Acta Vet. Brno 75 (2006): 277-81.
  15. ↑ Sherman, R. A., M. J. R. Hall, and S. Thomas. “Medicinal Maggots: an Ancient Remedy for Some Contemporary Afflictions.” Annu. Rev. Entomol. 45(2000):55-81.
  16. ↑ Dimarcq, J.L., D. Zachary, J.A. Hoffmann, D. Hoffmann, and J.M. Reichhart. "Insect immunity: expression of the two major inducible antibacterial peptides, defensin and diptericin, in Phormia terranovae." EMBO J. 9: 2507-515.
  17. 1 2 Green, A. A. "The Control of Blowflies Infesting Slaughter-Houses I. Field observations of the habits of blowflies." Annals of Applied Biology 38 (1951): 475-494.
  18. ↑ Samuel, William M., A. A. Kocan, Margo J. Pybus, and John W. Davis, eds. Parasitic Diseases of Wild Mammals. 2nd ed. Grand Rapids: Blackwell Professional, 2001.
  19. ↑ Scott, Danny W., and William H. Miller. Equine Dermatology. Philadelphia: Saunders, 2003.
  20. ↑ Pedigo, Larry P. Economic Thresholds for Integrated Pest Management. Ed. Leon G. Higley. Lincoln: University of Nebraska P, 1996.
  21. 1 2 Charabidze, Damien, Benoit Bourel, Helene LeBlanc, Valery Hedouin, and Didier Gosset. "Effect of Body Length and Temperature on the Crawling Speed of Protophormia terraenovae Larvae (Robineau-Desvoidy) (Diptera Calliphoridae)." Journal of Insect Physiology 54 (2008): 529-533.
  22. ↑ Bourel, Benoit, Gilles Tournel, Valery Hedouin, Mark Deveaux, M. L. Goff, and Didier Gosset. "Morphine Extraction in Necrophagous Insects Remains for Determining Ante-Mortem Opiate Intoxication." Forensic Science International 120 (2001): 127-31.
  23. ↑ Warren, J. A., and G. S. Anderson. "Use of Protophormia terraenovae (R-D) Developmental Data in a High Profile Homicide Case." [ permanent dead link ]Proceedings of the sixth meeting of the European Association For Forensic Entomology. EAFE 2008, Orthodox Academy of Crete, Kolymbari/Crete. 21 May 2008. Centre for Forensic Research.
  24. ↑ Numata, Hideharu and Sakiko Shiga. “Induction of Adult Diapause by Photoperiod and Temperature in Protophormia terraenovae (Diptera: Calliphoridae) in Japan” Environmental Entomology 24.6 (1995): 1633-1636.
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