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Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region

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 ([email protected][email protected])
2Montréal Insectarium / Space for life, 4581, rue Sherbrooke Est, Montréal, QC, H1X 2B2, Canada ([email protected])
3Biological Sciences, Bishop’s University, 2600 College Street, Sherbrooke, QC, J1M 1Z7, Canada ([email protected])

Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region

Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region

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 ([email protected][email protected])
2Montréal Insectarium / Space for life, 4581, rue Sherbrooke Est, Montréal, QC, H1X 2B2, Canada ([email protected])
3Biological Sciences, Bishop’s University, 2600 College Street, Sherbrooke, QC, J1M 1Z7, Canada ([email protected])

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and to present an updated checklist and identification key to the Nearctic species, as existing keys are either outdated, too broad in geographical scope to be user-friendly, and/or contain ambiguous characters.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 Pmegastigmata and Pcasei 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 PflavipesBoreopiophila 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)

Protopiophila litigata Bonduriansky 1995

Stearibia nigriceps Meigen, 1826

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.

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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