Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region
- CJAI #27
- January 22, 2015
Sabrina Rochefort1
,
Marjolaine Giroux2
,
Jade Savage3
,
Terry A. Wheeler1
1Department of Natural Resource Sciences, McGill University, Macdonald Campus, Ste-Anne-de-Bellevue, QC, H9X 3V9, Canada (sabrina.rochefort@mail.mcgill.ca, terry.wheeler@mcgill.ca)
2Montréal Insectarium / Space for life, 4581, rue Sherbrooke Est, Montréal, QC, H1X 2B2, Canada (marjolaine.giroux@ville.montreal.qc.ca)
3Biological Sciences, Bishop’s University, 2600 College Street, Sherbrooke, QC, J1M 1Z7, Canada (jsavage@ubishops.ca)
Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region
Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region
- CJAI #27
- January 22, 2015
Sabrina Rochefort1
,
Marjolaine Giroux2
,
Jade Savage3
,
Terry A. Wheeler1
1Department of Natural Resource Sciences, McGill University, Macdonald Campus, Ste-Anne-de-Bellevue, QC, H9X 3V9, Canada (sabrina.rochefort@mail.mcgill.ca, terry.wheeler@mcgill.ca)
2Montréal Insectarium / Space for life, 4581, rue Sherbrooke Est, Montréal, QC, H1X 2B2, Canada (marjolaine.giroux@ville.montreal.qc.ca)
3Biological Sciences, Bishop’s University, 2600 College Street, Sherbrooke, QC, J1M 1Z7, Canada (jsavage@ubishops.ca)
Abstract
Many species of Piophilidae (Diptera) are relevant to forensic entomology because their presence on a corpse can be helpful in estimating the postmortem interval (PMI) and document insect succession. The aims of this paper are to document the fauna of forensically relevant Piophilidae species worldwide and to present an updated checklist and identification key to the Nearctic species, as existing keys are either outdated, too broad in geographical scope to be user-friendly, and/or contain ambiguous characters. Thirteen species are included in the checklist and key. Information on their biology, taxonomy, character variability, and distribution is provided, supplementing the extensive work of McAlpine (1977).
Piophila casei males engaging in pugilistic territory defense.
Photo by Stephen A. Marshall
Introduction
Forensic entomology is the use of insects and other arthropods as evidence in legal investigations (Catts & Goff 1992). An important aspect of the discipline involves the estimation of the postmortem interval (PMI) based on arthropods associated with a body, an approach that requires extensive knowledge of the local fauna and its association with carrion at different decomposition stages and under different conditions (Goff 2000, Amendt et al. 2011). Necrophagous flies (Diptera) are of forensic importance as they often appear on a body first, consume most of the tissues, and display similar patterns of succession in different regions of the world, at least at the family level (Catts & Goff 1992, Amendt et al. 2011).
The family Piophilidae contains 82 species worldwide (Pape et al. 2009), at least 37 of which are present in the Nearctic region (McAlpine 1977). Several species can be found on dung, bone, garbage, decaying vegetation, fungi, bird nests and discarded antlers (Melander & Spuler 1917, McAlpine 1977, Bonduriansky & Brooks 1999a). Other species are associated with carrion in a range of decomposition stages and are therefore relevant to forensic entomology (Greenberg 1991, Byrd & Castner 2001). Depending on the locality, adult Piophilidae are mostly seen throughout the bloated and decay stages (Johnson 1975, Fiedler et al. 2008, Prado e Castro et al. 2012) while the larvae are more common in the advanced (Martín-Vega et al. 2011) and dry/remain stages (Martinez et al. 2006, Grisales et al. 2010).
Identifying species of forensic importance can sometimes be challenging when using morphological characters alone (Byrd & Castner 2001, Amendt et al. 2011) and alternatives such as DNA markers have been developed to identify problematic specimens (Wells & Stevens 2008). While molecular techniques can be a useful complement to morphology-based specimen identification, they do not always yield correct or unambiguous identification results (e.g. Whitworth et al. 2007) and require access to equipment, facilities and funding for molecular analysis; thus, morphology-based regional keys remain time- and cost-effective tools for the identification of most forensically important species.
McAlpine (1977) contributed significantly to the systematics of the family Piophilidae through the proposal of a revised classification of the family (although this was not based on an explicit phylogenetic analysis), description of six new species and two new genera, taxonomic changes to other species, the documentation of species distributions, and the publication of worldwide identification keys to species. McAlpines’s work, however, was published more than 35 years ago, and his species keys can be difficult to use, especially by non-experts, as they are based mostly on colour characters and do not include species described since that time.
In an effort to complement and update McAlpine (1977), the objectives of this work are to document Piophilidae species of known or potential forensic importance worldwide, to compile a checklist of Nearctic Piophilidae species relevant to forensic entomology based on published records in the literature and new data from Quebec (Canada), to update the distribution records and document the intraspecific variations of all species in the Nearctic checklist, and to develop a user-friendly key to forensically relevant Piophilidae species in the Nearctic.
Table 1. Piophilidae species documented in the forensic entomology literature.
| Species | Locality | References |
| Boreopiophila tomentosa Frey, 1930* | Canada: Manitoba | Gill 2005 |
| Liopiophila varipes Meigen, 1830 | Canada: New Brunswick | Michaud et al. 2010 |
| Germany
|
Fiedler et al. 2008
Baumjohann & Rudzinski 2013 |
|
| Mycetaulus bipunctatus (Fallén, 1823) | USA: Louisiana | Watson & Carlton 2008 |
| Parapiophila vulgaris (Fallén, 1820) | Germany | Fiedler et al. 2008
Baumjohann & Rudzinski 2013 |
| Poland | Matuszewski et al. 2008 | |
| Piophila casei (Linnaeus, 1758) | Canada: Saskatchewan | Sharanowski et al. 2008 |
| USA: Tennessee | Reed 1958** | |
| USA: Hawaii | Early & Goff 1986 | |
| USA: Colorado | de Jong & Chadwick 1999*** | |
| USA: Louisiana | Watson & Carlton 2008 | |
| Costa Rica | Carvalho et al. 2000 | |
| Argentina | Battán Horenstein et al. 2010 | |
| Spain | Martín-Vega et al. 2011 | |
| Portugal
Germany |
Prado e Castro et al. 2012
Baumjohann & Rudzinski 2013 |
|
| South Africa | Braack 1986 | |
| Thailand | Sukontason et al. 2001 | |
| Malaysia | Kumara et al. 2012 | |
| Piophila megastigmata McAlpine, 1978 | Portugal | Prado e Castro et al. 2012 |
| Spain | Martín-Vega et al. 2011 | |
| Paños et al. 2013 | ||
| South Africa | Braack 1986 | |
| Prochyliza azteca McAlpine, 1977 | Costa Rica | Jirón & Cartín 1981 |
| Prochyliza brevicornis Melander, 1924 | Canada: British Columbia | Anderson 1995 |
| Prochyliza nigrimana (Meigen, 1826) | USA: Tennessee | Reed 1958** |
| Portugal | Prado e Castro et al. 2012 | |
| Spain | Martín-Vega et al. 2011 | |
| Martín-Vega & Baz 2013 | ||
| Prochyliza xanthostoma Walker, 1849 | Canada: Manitoba | Gill 2005 |
| USA: Tennessee | Reed 1958** | |
| USA: Illinois | Johnson 1975**** | |
| USA: Louisiana | Watson & Carlton 2003 | |
| USA: Virginia | Tabor et al. 2005 | |
| Protopiophila latipes (Meigen, 1838) | Canada: New Brunswick | Michaud et al. 2010 |
| USA: Tennessee | Reed 1958** | |
| USA: Illinois | Johnson 1975**** | |
| Germany
|
Fiedler et al. 2008
Baumjohann & Rudzinski 2013 |
|
| Portugal | Prado e Castro et al. 2012 | |
| Stearibia nigriceps Meigen, 1826 | Canada: British Columbia | Anderson 1995 |
| Canada: Manitoba | Gill 2005 | |
| Canada: New Brunswick | Michaud et al. 2010 | |
| USA: Tennessee | Reed 1958** | |
| USA: Louisiana | Watson & Carlton 2003 | |
| USA: Virginia | Tabor et al. 2005 | |
| France | Leclercq 1996 | |
| Germany
|
Fiedler et al. 2008
Baumjohann & Rudzinski 2013 |
|
| Poland | Matuszewski et al. 2008 | |
| Spain | Martín-Vega et al. 2011 | |
| Portugal | Prado e Castro et al. 2012 | |
| Martinez et al. 2006 | ||
| Colombia | Grisales et al. 2010 | |
| India | Sathe et al. 2013 |
* Gill (2005) recorded B. tomentosa from southwestern Manitoba. All verified records of this species are from the arctic (Rochefort & Wheeler 2015) and vouchers of Gill’s specimens are unavailable. Thus we consider this record questionable.
The following studies did not use human or pig carcasses.
** Used dog carcasses
*** Used rabbit carcasses.
**** Used squirrel, rabbit, cat and opossum carcasses.
Methods & Materials
The checklist of Nearctic Piophilidae of forensic importance was compiled from the literature and from specimens collected in the context of a study of the insect fauna associated with nine pig carcasses in three semi-urban sites located along a latitudinal gradient in Quebec (Canada) in summer 2011 by Giroux, Savage and collaborators. The sampling sites were Ste-Anne-de-Bellevue (45.436°, -73.909°) (22 June to 14 August), Sherbrooke (45.361°, -71.844°) (6 July to 26 August) and Saguenay (La Baie) (48.350°, -70.967°) (4 July to 19 August).
Approximately 1065 piophilid specimens from these three sites were examined. They were identified to species using McAlpine (1977) and compared with reference specimens in the Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, ON (CNC) and the Lyman Entomological Museum, McGill University, Ste-Anne-de-Bellevue, QC (LEM). Voucher specimens were deposited in the Insectarium de Montréal’s scientific collections (IMQC) and the Bishop’s University Insect Collection (BUIC).
In addition, approximately 5600 specimens of Piophilidae included in the key were examined to document geographic distribution and intraspecific variability. These specimens are deposited in CNC; LEM; the Biodiversity Institute of Ontario, Guelph, ON, (BIO); the University of Guelph Insect Collection, Guelph, ON, (DEBU); the Ouellet-Robert Collection, Université de Montréal, Montréal, QC (UMIC); the Spencer Entomological Collection, Beaty Biodiversity Museum, Vancouver, BC (UBCZ); the Strickland Museum, University of Alberta, Edmonton, AB (UASM); and the United States National Museum of Natural History, Washington DC, (USNM). Additional published records were added where necessary.
Classification of genera and species follows McAlpine (1977) for consistency with the Nearctic literature. Ozerov (2004) proposed a different generic classification, followed by some authors, especially in Europe, but that classification, like McAlpine’s, was not based on a phylogenetic analysis and thus is no better supported. A comprehensive phylogenetic analysis and revised classification of the Piophilidae is required.
Photographs for the key were taken with an Olympus DP71 camera mounted on an Olympus SZX16 stereoscope. Images were captured and stacked using DP Controller and Combine ZP (Hadley 2008) before being enhanced using Adobe Photoshop [CS3] (Adobe Systems, Mountain view, CA).
Identification challenges in the Piophilidae
The key to the world species of Piophilidae by McAlpine (1977) remains the most complete work for the identification of adults even though several new species have since been described (Soós 1977, McAlpine 1978, McAlpine 1989, Ozerov 1989, Ozerov & Barták 1993, Bonduriansky 1995, Merz 1996, Ozerov 2000, 2002, 2004, 2007, Martín-Vega 2014, Rochefort & Wheeler 2015) and additional distribution records are known based on museum specimens. McAlpine’s (1977) key is not ideal to identify piophilids in forensic studies as it includes many non-forensically important species and is mostly based on colour differences which can be troublesome in this family due to intraspecific colour variations. Such polymorphisms have been documented in several species of forensic interest such as Parapiophila atrifrons (Melander & Spuler) (Rochefort & Wheeler 2015), Parapiophila vulgaris (Fallén) (McAlpine 1977) and Prochyliza nigrimana (Meigen) (Martín-Vega & Baz 2011). Some species are also morphologically very similar. In southern Europe, for example, the very similar Piophila casei (L.) and Piophila megastigmata McAlpine co-occur; this can cause identification errors that may lead to erroneous PMI estimations (Martín-Vega 2011). Confusion between P. megastigmata and P. casei has occurred in forensic research (e.g., Prado e Castro 2010), a mistake that might actually be widespread in the field (Prado e Castro et al. 2012), and we suspect that such confusion may also apply to other species pairs in the Nearctic.
Some forensic studies limit Piophilidae identification to the family or generic levels (e.g., Schoenly et al. 2007, Voss et al. 2008, Velasquez et al. 2010, Bygarski & Leblanc 2013) or identify the material as Piophila casei, despite the fact that several species have been documented in forensic literature worldwide (Table 1). Piophila casei is a well-known, cosmopolitan, synanthropic species and was the first piophilid documented in forensic studies (Megnin 1894). This may account for the possibility that Piophilidae collected in forensic studies are frequently identified (or misidentified) as Piophila casei (see Martín-Vega 2011).
Checklist
Thirteen species are included in our Nearctic checklist. Nine of these were compiled from the literature (Table 1), two from museum records, and two, Mycetaulus subdolus (Johnson 1922) and Parapiophila atrifrons (Melander & Spuler 1917), were recorded for the first time from pig carcasses in Quebec (along with four other species whose forensic relevance was already known). Parapiophila flavipes (Zetterstedt 1847) and Protopiophila litigata Bonduriansky 1995 have been added to the Nearctic checklist since many specimens from DEBU have been collected on carcasses, an indication that they may be of forensic relevance. Parapiophila vulgaris has also been added to the Nearctic checklist because it is widely distributed in the region and is a species of forensic relevance in Europe (Fiedler et al. 2008, Matuszewski et al. 2008). Parapiophila specimens identified only to genus in several studies (e.g. Sharanowski et al. 2008, Michaud et al. 2010) could belong to this species, although they may also correspond to other species such as Parapiophila atrifrons and P. flavipes. Boreopiophila tomentosa was excluded from the checklist based on its dubious record in Gill (2005) (see footnote in Table 1).
Liopiophila varipes Meigen, 1830
Mycetaulus bipunctatus (Fallén, 1823) complex
Mycetaulus subdolus (Johnson, 1922)
Parapiophila atrifrons (Melander & Spuler, 1917)
Parapiophila flavipes (Zetterstedt 1847)
Parapiophila vulgaris (Fallén, 1820)
Piophila casei (Linnaeus, 1758)
Prochyliza brevicornis Melander, 1924
Prochyliza nigrimana (Meigen, 1826)
Prochyliza xanthostoma Walker, 1849
Protopiophila latipes (Meigen, 1838)
Species Key
Acknowledgments
We thank Robert Loiselle (Université du Québec à Chicoutimi) and Eric Lucas (Université du Québec à Montréal) who coordinated the Chicoutimi and Montreal field work, Jeff Skevington and Brad Sinclair (CNC), Valérie Lévesque-Beaudin (BIO), Steve Marshall and Steve Paiero (DEBU), Louise Cloutier (UMIC), Karen Needham (UBCZ), Danny Shpeley (UASM) and Torsten Dikow (USNM) for access to specimens, Stéphane LeTirant (IMQC) for logistic support and access to specimens, and Karine Thivierge and Dominique St-Pierre (Laboratoire de Santé Publique du Québec) for training and access to the stereomicroscope and photography equipment. We thank G. Anderson, R. Bonduriansky and S. Marshall for helpful comments on the manuscript. This research was funded by Natural Sciences and Engineering Research Council of Canada Discovery Grants to JS and TAW, a Bishop’s University Senate Research Grant to JS, and by support from the Insectarium de Montréal to MG.
References
Amendt, J., C.S. Richards, C.P. Campobasso, R. Zehner, and M.J.R. Hall. 2011. Forensic entomology: Applications and limitations. Forensic Science, Medicine, and Pathology, 7: 379–392.
Anderson, G.S. 1995. The use of insects in death investigations: An analysis of cases in British Columbia over a five year period. Canadian Society of Forensic Science Journal, 28: 277–292.
Baumjohann, K., and Rudzinski, H.-G. 2013. Bemerkenswerte Fliegen-Funde (Diptera) im Rahmen einer forensisch entomologischen Freilandstudie in Nordrhein-Westfalen (Deutschland). Studia Dipterologica, 19: 9–16.
Battán Horenstein, M., A.X. Linhares, B. Rosso de Ferradas, and D. García. 2010. Decomposition and dipteran succession in pig carrion in central Argentina: ecological aspects and their importance in forensic science. Medical and Veterinary Entomology, 24: 16–25.
Bonduriansky, R. 1995. A new Nearctic species of Protopiophila Duda (Diptera: Piophilidae), with notes on its behaviour and comparison with P. latipes (Meigen). The Canadian Entomologist, 127: 859–863.
Bonduriansky, R. 2002. Leaping behaviour and responses to moisture and sound in larvae of piophilid carrion flies. The Canadian Entomologist, 134: 647–656.
Bonduriansky, R. 2003. Layered sexual selection: a comparative analysis of sexual behaviour within an assemblage of piophilid flies. Canadian Journal of Zoology, 81: 479–491.
Bonduriansky, R. 2006. Convergent evolution of sexual shape dimorphism in Diptera. Journal of Morphology, 267: 602–611.
Bonduriansky, R., and R.J. Brooks. 1998a. Copulation and oviposition behaviour of Protopiophila litigata (Diptera: Piophilidae). The Canadian Entomologist, 130: 399–405.
Bonduriansky, R., and R.J. Brooks. 1998b. Male antler flies (Protopiophila litigata; Diptera: Piophilidae) are more selective than females in mate choice. Canadian Journal of Zoology, 76: 1277–1285.
Bonduriansky, R., and R.J. Brooks. 1999a. Why do male antler flies (Protopiophila litigata) fight? The role of male combat in the structure of mating aggregations on moose antlers. Ethology, Ecology & Evolution 11: 287–301.
Bonduriansky, R., and R.J. Brooks. 1999b. Reproduction allocation and reproductive ecology of seven species of Diptera. Ecological Entomology, 24: 389–395.
Bonduriansky, R., and L. Rowe. 2005. Sexual selection, genetic architecture, and the condition dependence of body shape in sexually dimorphic fly Prochyliza xanthostoma (Piophilidae). Evolution, 59: 138–151.
Bonduriansky, R., E.J. Wheeler, and L. Rowe. 2005. Ejaculate feeding and female fitness in the sexually dimorphic fly Prochyliza xanthostoma (Diptera: Piophilidae). Animal Behaviour, 69: 489–497.
Braack, L.E.O. 1986. Arthropod associated with carcasses in the northern Kruger National Park. South African Journal of Wildlife Research 16: 91–98.
Bucheli, S.R., J.A. Bytheway, S.M. Pustilnik, and J. Florence. 2009. Insect successional pattern of a corpse in cooler months of subtropical southeastern Texas. Journal of Forensic Sciences, 54: 452–455.
Bygarski, K., and H.N. LeBlanc. 2013. Decomposition and arthropod succession in Whitehorse, Yukon Territory, Canada. Journal of Forensic Sciences, 58: 413–418.
Byrd, J.H., and J.L. Castner. 2001. Insects of forensic importance In: J.H. Byrd, and J.L. Castner (Eds). Forensic entomology: the utility of arthropods in legal investigation. CRC, Boca Raton, FL. pp. 43–79.
Carvalho, L.M.L., P.J. Thyssen, A.X. Linhares, and F.A.B. Palhares. 2000. A checklist of arthropods associated with pig carrion and human corpses in Southeastern Brazil. Memórias do Instituto Oswaldo Cruz, Rio de Janeiro, 95: 135–138.
Catts, E.P., and M.L. Goff. 1992. Forensic entomology in criminal investigations. Annual Review of Entomology, 37: 253–272.
Cumming, J.M., and D.M Wood. 2009. Adult morphology and terminology. In: Brown, B.V., A. Borkent, J.M. Cumming, D.M. Wood, N.E. Woodley & M. Zumbado (Eds). Manual of Central American Diptera. Volume 1. Ottawa, Canada: National Research Council Press. Pp. 2–50.
Duda, O. 1924. Revision der eropäischen u. Grönländischen sowie einiger südostasiat. Arten der Gattung Piophila Fallén (Dipteren). Konowia 3: 97–113, 153–203.
Early, M., and M.I. Goff. 1986. Arthropod succession patterns in exposed carrion on the island of O’ahu, Hawaiian Islands, USA. Journal of Medical Entomology, 24: 520–531.
Fiedler, A., M. Halbach, B. Sinclair, and M. Benecke. 2008. What is the edge of a forest? A diversity analysis of adult Diptera found on decomposing piglets inside and on the edge of a western German Woodland inspired by a courtroom question. Entomologie Heute, 20: 173–191.
Gill, G.J. 2005. Decomposition and arthropod succession on above ground pig carrion in rural Manitoba. Technical Report (TR-06-2005). Canadian Police Research Centre. 178pp.
Goff, M.L. 2000. A fly for the prosecution: How insect evidence helps solve crimes. Harvard University Press, Cambridge.
Greenberg, B. 1991. Flies as forensic indicators. Journal of Medical Entomology, 28: 565–577.
Grisales, D., M. Ruiz, and S. Villegas. 2010. Insects associated with exposed decomposing bodies in the Colombian Andean Coffee Region. Revista Brasileira de Entomologia, 54: 637–644.
Hadley, A. 2008. CombineZP Image Stacking Software. Retrieved July 10, 2014, from http://www.hadleyweb.pwp.blueyonder.co.uk/
Jirón, L.F., and V.M. Cartín. 1981. Insect succession in the decomposition of a mammal in Costa Rica. Journal of the New York Entomological Society, 89: 158–165.
de Jong, G.D., and J.W. Chadwick. 1999. Decomposition and arthropod succession on exposed rabbit carrion during summer at high altitudes in Colorado, USA. Journal of Medical Entomology, 36: 833–845.
Johnson, C.W. 1922. New genera and species of Diptera. Occasional Papers of the Bostom Society of Natural History.Johnson, M.D. 1975. Seasonal and microseral variations in the insect populations on carrion. American Midland Naturalist, 93: 79–90.
Keh, B. 1985. Scope and applications of forensic entomology. Annual Review of Entomology, 30: 137–154.
Kumara, T.K., R.H.L. Disney, A.A. Hassan, M. Flores, T.S. Hwa, Z. Mohamed, M.R. Chesalmah, and S. Bhupinder. 2012. Occurrence of oriental flies associated with indoor and outdoor human remains in the tropical climate of north Malaysia. Journal of Vector Ecology, 37: 62–68.
Leclercq, M. 1996. A propos de l’entomofaune d’un cadavre de sanglier. Bulletin et Annales de la Societé Royale Entomologique de Belgique, 132: 417–422.
Liu, D., and B. Greenberg. 1989. Immature stages of some flies of forensic importance. Annals of the Entomological Society of America, 82: 80–93.
Malloch, J.R. 1932. New species and other records of Otitidae (Ortalidae), Piophilidae, Clusiidae, Chloropidae, and Drosophilidae from the Marquesas. Bernice P. Bishop Museum-Bulletin, 98: 205–223.
Martinez, E., P. Duque, and M. Wolff. 2006. Succession pattern of carrion-feeding insects in Paramo, Colombia. Forensic Science International, 166: 182–189.
Martín-Vega, D. 2011. Skipping clues: Forensic importance of the family Piophilidae (Diptera). Forensic Science International, 212: 1–5.
Martín-Vega, D. 2014. On the identity of Prochyliza nigrimana (Meigen) and Prochyliza nigricornis (Meigen) (Diptera: Piophilidae), with a synopsis of Prochyliza Walker and description of a new species. Zootaxa, 3893: 277–292.
Martín-Vega, D., and A. Baz. 2011. Variation in the colour of the necrophagous fly, Prochyliza nigrimana (Diptera: Piophilidae): A case of seasonal polymorphism. European Journal of Entomology, 108: 231–234.
Martín-Vega, D., and A. Baz. 2013. Sarcosaprophagous Diptera assemblages in natural habitats in Central Spain: Spatial and seasonal changes in composition. Medical and Veterinary Entomology, 27: 64–76.
Martín-Vega, D., A. Baz, and L.M. Díaz-Aranda. 2012. The immature stages of the necrophagous fly, Prochyliza nigrimana: comparison with Piophila casei and medicolegal considerations (Diptera: Piophilidae). Parasitology Research, 111: 1127–1135.
Martín-Vega, D., A. Gómez-Gómez, A. Baz, and L.M. Díaz-Aranda. 2011. New piophilid in town: the first Palaearctic record of Piophila megastigmata and its coexistence with Piophila casei in Central Spain. Medical and Veterinary Entomology, 25: 64–69.
Martín-Vega, D., L.M. Diaz-Aranda., and A. Baz. 2014. The immature stages of the necrophagous fly Liopiophila varipes and considerations on the genus Liopiophila (Diptera: Piophilidae). Deutsche Entomologische Zeitschrift, 61: 37–42.
Matuszewski, S., D. Bajerlein, S. Konwerski, and K. Szpila. 2008. An initial study of insect succession and carrion decomposition in various forest habitats of Central Europe. Forensic Science International, 180: 61–69.
Merz, B. 1996. Die Piophilidae (Diptera) der Schweiz mit Beschreibung einer neuen Art. Bulletin de la Société Entomologique Suisse, 69: 345–360.
McAlpine, D.K. 1989. A synopsis of the Australian Piophilidae (Diptera: Schizophora). General and Applied Entomology, 21: 17–24.
McAlpine, J.F. 1977. A revised classification of the Piophilidae, including ‘Neottiophilidae’ and ‘Thyreophoridae’ (Diptera: Schizophora). Memoirs of the Entomological Society of Canada, 103: 1–66.
McAlpine, J.F. 1978. A new species of Piophila from South Africa (Diptera: Piophilidae). Annals of the Natal Museum, 23: 455–459.
Megnin, P. 1894. La faune des cadavres – Application de l’entomologie à la médecine légale. Encyclopedie Scientifique des Aide-Mémoire, 214 pp. G. Masson, Gauthier-Villars et fils, Paris.
Melander, A.L., and A. Spuler. 1917. The dipterous families Sepsidae and Piophilidae. Washington Agricultural Experiment Station Bulletin: 97.
Michaud, J.-P., C.G. Majka, J.-P. Privé, and G. Moreau. 2010. Natural and anthropogenic changes in the insect fauna associated with carcasses in the North American Maritime lowlands. Forensic Science International, 202: 64–70.
Ozerov, A.L. 1989. Sepsidae and Piophilidae (Diptera) of the Zeya State Reserve. Translated from Entomologicheskoye Obozreniye, 4: 839–849.
Ozerov, A.L. 2000. A new species of the genus Actenoptera (Diptera, Piophilidae) from the Russian Far East. Entomological Review, 80: 244–245.
Ozerov, A.L. 2002. First record of Neopiophila McAlpine, 1977 (Diptera: Piophilidae) in Palaearctic, with description of a new species. Russian Entomological Journal, 11: 223–224.
Ozerov, A.L. 2004. On the classification of the family Piophilidae (Diptera). Entomological Review, 84: 600–608.
Ozerov, A.L. 2007. Three new species of Carnidae and Piophilidae (Diptera) from Turkey and Russia. Russian Entomological Journal, 16: 491–493.
Ozerov, A.L. and M. Barták. 1993. Two new Palaearctic species of Piophilidae (Diptera). Russian Entomological Journal, 2: 73-76.
Ozerov, A.L. and A.L. Norrbom. 2010. Piophilidae. In: Brown, B.V., A. Borkent, J.M. Cumming, D.M. Wood, N.E. Woodley & M. Zumbado (Eds). Manual of Central American Diptera. Volume 2. Ottawa, Canada: National Research Council Press. Pp. 865–869.
Prado e Castro, C., and M.D. García. 2010. Additions to the Piophilidae (Diptera) fauna from Portugal, with new records. Graellsia, 66: 101–105.
Prado e Castro, C., E. Cunha, A. Serrano, and M.D. García. 2012. Piophila megastigmata (Diptera: Piophilidae): First records on human corpses. Forensic Science International, 214: 23–26.
Paños, A., M.I. Arnaldos, M.D. García, and N. Ubero-Pascal. 2013. Ultrastructure of preimaginal stages of Piophila megastigmata McAlpine, 1978 (Diptera, Piophilidae): A fly of forensic importance. Parasitology Research, 112: 3771–3788.
Pape, T., D. Bickel, and R. Meier (Eds). 2009. Appendix A: Species of Diptera per family for all regions. In: Diptera Diversity: Status, challenges and tools. Brill, Leiden: Boston. Pp. 439–444.
Reed, H.B. 1958. A study of dog carcass communities in Tennessee, with special reference to the insects. American Midland Naturalist, 59: 213–245.
Rochefort, S., and T.A. Wheeler. 2015. Diversity of Piophilidae (Diptera) in northern Canada and description of a new Holarctic species of Parapiophila McAlpine. Zootaxa (manuscript in revision).
Sathe, T.V., A. Sathe, and N.T. Sathe. 2013. Diversity of dipterous forensic insects from Western Maharashtra, India. International Journal of Pharma and Bio Sciences, 4(2): P173–P179.
Schoenly, K.G., N.H. Haskell, R.D. Hall, and J.R. Gbur. 2007. Comparative performance and complementarity of four sampling methods and arthropod preference tests from human and porcine remains at the Forensic Anthropology Center in Knoxville, Tennessee. Journal of Medical Entomology, 44: 881–894.
Sharanowski, B.J., E.G. Walker, and G.S. Anderson. 2008. Insect succession and decomposition patterns on shaded and sunlit carrion in Saskatchewan in three different seasons. Forensic Science International, 179: 219–240.
Soós, A. 1977. Acalyptrata (Diptera) from Afghanistan II. Folia Entomologica Hungarica, XXX: 133–137.
Sukontason, K.L., K. Sukontason, S. Piangjai, W. Choochote, R.C. Vogtsberger, and J.K. Olson. 2001. Scanning electron microscopy of the third-instar Piophila casei (Diptera: Piophilidae), a fly species of forensic importance. Journal of Medical Entomology, 38: 756–759.
Tabor, K.L., R.D. Fell, and C.C. Brewster. 2005. Insect fauna visiting carrion in southwest Virginia. Forensic Science International, 150: 73–80.
Valásquez, T., C. Magaña, A. Martinez-Sánchez, and S. Rojo. 2010. Diptera of forensic importance in the Iberian Peninsula: Larval identification key. Medical and Veterinary Entomology, 24: 293–308.
Voss, S.C., S.L. Forbes, and I.R. Dadour. 2008. Decomposition and insect succession on cadavers inside a vehicle environment. Forensic Science, Medicine and Pathology, 4: 22–32.
Watson, E.J., and C.E. Carlton. 2003. Spring succession of necrophilous insects on wildlife carcasses in Louisiana. Journal of Medical Entomology, 40: 338–347.
Wells, J.D., and R.S. Stevens. 2008. Application of DNA-based methods in forensic entomology. Annual Review of Entomology, 53: 103–120.
Whitworth, T.L., R.D. Dawson, H. Magalon, and E. Baudry. 2007. DNA barcoding cannot reliably identify species of the blowfly genus Protocalliphora (Diptera: Calliphoridae). Proceedings of the Royal Society, B, 274: 1731–1739.
Zuska, J., and P. Laštovka. 1965. A review of the Czechoslovak species of the family Piophilidae with special reference to their importance to food industry (Diptera, Acalyptrata). Acta Entomologica Bohemoslovaca, 62: 141–157.
Cite
Rochefort, S., Giroux, M., Savage, J., Wheeler, T.A. 2015. Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region. Canadian Journal of Arthropod Identification No. 27: January 22, 2015. Available online at doi: 10.3752/cjai.2015.27
