Canadian Journal of Arthropod Identifi cation No. 27 (January, 2015) ROCHEFORT ET AL.

Key to Forensically Important Piophilidae (Diptera) in the Nearctic Region Sabrina Rochefort1, Marjolaine Giroux2, Jade Savage3 and 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and to present an updated checklist and identifi cation key to the Nearctic species, as existing keys are either outdated, too broad in geographical scope to be user-friendly, and/or contain ambiguous characters.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).

Introduction stages (Martinez et al. 2006, Grisales et al. 2010). Forensic entomology is the use of insects and other Identifying species of forensic importance can arthropods as evidence in legal investigations (Catts sometimes be challenging when using morphological & Goff 1992). An important aspect of the discipline characters alone (Byrd & Castner 2001, Amendt et al. involves the estimation of the postmortem interval (PMI) 2011) and alternatives such as DNA markers have been based on arthropods associated with a body, an approach developed to identify problematic specimens (Wells that requires extensive knowledge of the local fauna and & Stevens 2008). While molecular techniques can be its association with carrion at different decomposition a useful complement to morphology-based specimen stages and under different conditions (Goff 2000, identifi cation, they do not always yield correct or Amendt et al. 2011). Necrophagous fl ies (Diptera) are of unambiguous identifi cation results (e.g. Whitworth et forensic importance as they often appear on a body fi rst, al. 2007) and require access to equipment, facilities and consume most of the tissues, and display similar patterns funding for molecular analysis; thus, morphology-based of succession in different regions of the world, at least at regional keys remain time- and cost-effective tools for the family level (Catts & Goff 1992, Amendt et al. 2011). the identifi cation of most forensically important species. The family Piophilidae contains 82 species worldwide McAlpine (1977) contributed signifi cantly to the (Pape et al. 2009), at least 37 of which are present in the systematics of the family Piophilidae through the Nearctic region (McAlpine 1977). Several species can proposal of a revised classifi cation of the family (although be found on dung, bone, garbage, decaying vegetation, this was not based on an explicit phylogenetic analysis), fungi, bird nests and discarded antlers (Melander & description of six new species and two new genera, Spuler 1917, McAlpine 1977, Bonduriansky & Brooks taxonomic changes to other species, the documentation 1999a). Other species are associated with carrion in of species distributions, and the publication of worldwide a range of decomposition stages and are therefore identifi cation keys to species. McAlpines’s work, relevant to forensic entomology (Greenberg 1991, however, was published more than 35 years ago, and his Byrd & Castner 2001). Depending on the locality, adult species keys can be diffi cult to use, especially by non- Piophilidae are mostly seen throughout the bloated and experts, as they are based mostly on colour characters decay stages (Johnson 1975, Fiedler et al. 2008, Prado e and do not include species described since that time. Castro et al. 2012) while the larvae are more common in In an effort to complement and update McAlpine the advanced (Martín-Vega et al. 2011) and dry/remain (1977), the objectives of this work are to document

doi:10.3752/cjai.2015.27 1 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Piophilidae species of known or potential forensic revised classification of the Piophilidae is required. importance worldwide, to compile a checklist of Nearctic Photographs for the key were taken with an Piophilidae species relevant to forensic entomology Olympus DP71 camera mounted on an Olympus SZX16 based on published records in the literature and new data stereoscope. Images were captured and stacked using from Quebec (Canada), to update the distribution records DP Controller and Combine ZP (Hadley 2008) before and document the intraspecific variations of all species being enhanced using Adobe Photoshop [CS3] (Adobe in the Nearctic checklist, and to develop a user-friendly Systems, Mountain view, CA). key to forensically relevant Piophilidae species in the Nearctic. Identification challenges in the Piophilidae The key to the world species of Piophilidae by Materials and Methods McAlpine (1977) remains the most complete work for The checklist of Nearctic Piophilidae of forensic the identification of adults even though several new importance was compiled from the literature and from species have since been described (Soós 1977, McAlpine specimens collected in the context of a study of the 1978, McAlpine 1989, Ozerov 1989, Ozerov & Barták insect fauna associated with nine pig carcasses in three 1993, Bonduriansky 1995, Merz 1996, Ozerov 2000, semi-urban sites located along a latitudinal gradient in 2002, 2004, 2007, Martín-Vega 2014, Rochefort & Quebec (Canada) in summer 2011 by Giroux, Savage Wheeler 2015) and additional distribution records are and collaborators. The sampling sites were Ste-Anne- known based on museum specimens. McAlpine’s (1977) de-Bellevue (45.436°, -73.909°) (22 June to 14 August), key is not ideal to identify piophilids in forensic studies Sherbrooke (45.361°, -71.844°) (6 July to 26 August) as it includes many non-forensically important species and Saguenay (La Baie) (48.350°, -70.967°) (4 July to and is mostly based on colour differences which can be 19 August). troublesome in this family due to intraspecific colour Approximately 1065 piophilid specimens from these variations. Such polymorphisms have been documented three sites were examined. They were identified to species in several species of forensic interest such as Parapiophila using McAlpine (1977) and compared with reference atrifrons (Melander & Spuler) (Rochefort & Wheeler specimens in the Canadian National Collection of Insects, 2015), Parapiophila vulgaris (Fallén) (McAlpine 1977) Arachnids and Nematodes, Ottawa, ON (CNC) and the and Prochyliza nigrimana (Meigen) (Martín-Vega & Lyman Entomological Museum, McGill University, Ste- Baz 2011). Some species are also morphologically very Anne-de-Bellevue, QC (LEM). Voucher specimens were similar. In southern Europe, for example, the very similar deposited in the Insectarium de Montréal’s scientific Piophila casei (L.) and Piophila megastigmata McAlpine collections (IMQC) and the Bishop’s University Insect co-occur; this can cause identification errors that may Collection (BUIC). lead to erroneous PMI estimations (Martín-Vega 2011). In addition, approximately 5600 specimens of Confusion between P. megastigmata and P. casei has Piophilidae included in the key were examined to occurred in forensic research (e.g., Prado e Castro 2010), document geographic distribution and intraspecific a mistake that might actually be widespread in the field variability. These specimens are deposited in CNC; LEM; (Prado e Castro et al. 2012), and we suspect that such the Biodiversity Institute of Ontario, Guelph, ON, (BIO); confusion may also apply to other species pairs in the the University of Guelph Insect Collection, Guelph, Nearctic. ON, (DEBU); the Ouellet-Robert Collection, Université Some forensic studies limit Piophilidae identification de Montréal, Montréal, QC (UMIC); the Spencer to the family or generic levels (e.g., Schoenly et al. 2007, Entomological Collection, Beaty Biodiversity Museum, Voss et al. 2008, Velasquez et al. 2010, Bygarski & Vancouver, BC (UBCZ); the Strickland Museum, Leblanc 2013) or identify the material as Piophila casei, University of Alberta, Edmonton, AB (UASM); and despite the fact that several species have been documented the United States National Museum of Natural History, in forensic literature worldwide (Table 1). Piophila casei Washington DC, (USNM). Additional published records is a well-known, cosmopolitan, synanthropic species and were added where necessary. was the first piophilid documented in forensic studies Classification of genera and species follows (Megnin 1894). This may account for the possibility that McAlpine (1977) for consistency with the Nearctic Piophilidae collected in forensic studies are frequently literature. Ozerov (2004) proposed a different generic identified (or misidentified) as Piophila casei (see classification, followed by some authors, especially in Martín-Vega 2011). 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

doi:10.3752/cjai.2015.27 2 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

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 Prado e Castro et al. 2012

Germany 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

doi:10.3752/cjai.2015.27 3 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Table 1 cont’d. Piophilidae species documented in the forensic entomology literature.

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

doi:10.3752/cjai.2015.27 4 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Table 1 cont’d. Piophilidae species documented in the forensic entomology literature.

Stearibia nigriceps Meigen, 1826 cont’d 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.

Checklist of Nearctic Piophilidae of forensic interest Parapiophila atrifrons (Melander & Spuler, 1917) Thirteen species are included in our Nearctic checklist. Parapiophila flavipes (Zetterstedt 1847) Nine of these were compiled from the literature (Table 1), Parapiophila vulgaris (Fallén, 1820) two from museum records, and two, Mycetaulus subdolus Piophila casei (Linnaeus, 1758) (Johnson 1922) and Parapiophila atrifrons (Melander & Prochyliza brevicornis Melander, 1924 Spuler 1917), were recorded for the first time from pig Prochyliza nigrimana (Meigen, 1826) carcasses in Quebec (along with four other species whose Prochyliza xanthostoma Walker, 1849 forensic relevance was already known). Parapiophila Protopiophila latipes (Meigen, 1838) flavipes (Zetterstedt 1847) and Protopiophila litigata Protopiophila litigata Bonduriansky 1995 Bonduriansky 1995 have been added to the Nearctic Stearibia nigriceps Meigen, 1826 checklist since many specimens from DEBU have been collected on carcasses, an indication that they may be Key to the Nearctic Piophilidae of forensic interest of forensic relevance. Parapiophila vulgaris has also Morphological terminology follows Cumming & been added to the Nearctic checklist because it is widely Wood (2009). All measurements were taken on dry distributed in the region and is a species of forensic specimens; body length was measured from the front relevance in Europe (Fiedler et al. 2008, Matuszewski of the head (excluding the antenna) to the end of the et al. 2008). Parapiophila specimens identified only to abdomen; wing length was measured from base to apex. genus in several studies (e.g. Sharanowski et al. 2008, Sepsidae and Heleomyzidae are two other acalyptrate Michaud et al. 2010) could belong to this species, Diptera families often collected in large numbers although they may also correspond to other species such on carcasses. They are sometimes misidentified as as Parapiophila atrifrons and P. flavipes. Boreopiophila a Piophilidae because of similarity in the overall tomentosa was excluded from the checklist based on its appearance of some species. Characters to distinguish dubious record in Gill (2005) (see footnote in Table 1). these two families from Piophilidae are included in the Liopiophila varipes Meigen, 1830 identification key. Mycetaulus bipunctatus (Fallén, 1823) complex Mycetaulus subdolus (Johnson, 1922)

doi:10.3752/cjai.2015.27 5 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Major family-level characters (Fig. 1–5)

PIOPHILIDAE Differentiating Piophilidae from Arista inserted Sepsidae and Heleomyzidae dorsally IDENTIFICATION KEY (Click here) (Click here)

Subcostal break present

Vibrissa present 1 2 3 Postocellars divergent

Majority of species glossy black and yellow

Some species glossy yellow and brown 4 5 doi:10.3752/cjai.2015.27 6 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Differentiating Sepsidae (Fig. 6–9) from Piophilidae

Ant-like body

7 Vibrissa absent, but some 8 specimens have a short seta 6 at the same location Strong seta on margin of posterior thoracic spiracle Features shared with Piophilidae - Similar size - Often glossy black and yellow Subcostal break absent - Apical wing spot present in some species (Fig. 10)

9 Differentiating Heleomyzidae 10 from Piophilidae (Click here) doi:10.3752/cjai.2015.27 7 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Differentiating Heleomyzidae from Piophilidae (Fig. 11–13)

12 Postocellars convergent

11

Features shared with Piophilidae Strong costal spines along costal vein - Subcostal break present - Vibrissa present - Some species yellow and brown

13 PIOPHILIDAE IDENTIFICATION KEY (Click here) doi:10.3752/cjai.2015.27 8 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

14 16

15 17 1 Wing with apical spot present (Fig. 14). Fore femur and tibia entirely yellow (Fig. 15). 2

1’ Wing with apical spot absent (Fig. 16). Fore femur and/or tibia partially to entirely black (Fig. 3 17).

doi:10.3752/cjai.2015.27 9 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Cell r2+3 18 Cell r2+3 21

19 20 22 23

2 (1) Apical spot positioned at wing apex and filling tip of cell r2+3 (Fig. 18). Mycetaulus bipunctatus Two postsutural dorsocentrals present (Fig. 19). Abdomen entirely black (Fig. 20) complex

2’ Apical spot positioned before wing apex, not filling tip of cell r2+3 (Fig. Mycetaulus subdolus 21). Three postsutural dorsocentrals present, anterior one reduced (Fig. 22). Syntergite 1+2 and sometimes tergite 3 partially to entirely (Johnson) doi:10.3752/cjai.2015.27yellow, remaining tergites black or brown (Fig. 23). 10 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

29 30 31

3 (1) Postpronotum with 1–2 postpronotals (socket visible if bristle broken) (Fig. 29–30). 4

3’ Postpronotum without bristles (Fig. 31). 8

doi:10.3752/cjai.2015.27 11 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

32 34

33 35

4(3) Two postpronotals present (Fig. 32). Scutum with 1+3 dorsocentrals (Fig. 33). 5

4’ One postpronotal (posterior) present (Fig. 34). Scutum with 0+1 dorsocentrals (Fig. 35). 6

doi:10.3752/cjai.2015.27 12 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

36 38

37 39 5(4) Anepisternum, anepimeron, and katepisternum entirely glossy (Fig. 36). Mid Protopiophila and hind femora and tibiae mainly yellow (Fig. 37). latipes (Meigen)

5’ Anepisternum partly dull, anepimeron and katepisternum entirely dull (Fig. Protopiophila 38). Mid and hind femora and tibiae mainly black (Fig. 39). litigata doi:10.3752/cjai.2015.27 Bonduriansky 13 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

42 44 45

43 46 6 (4) First tarsomere of fore leg white (Fig. 42). Fore coxa black (Fig. 43). Parapiophila atrifrons (Melander & Spuler)

6’ First tarsomere of fore leg yellow or black (Fig. 44–45). Fore coxa partially 7 to entirely yellow (Fig. 46).

doi:10.3752/cjai.2015.27 14 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. 52

48 49 51

50 53

7 (6) Frons mostly yellow (black on posterior fronto-orbital plate and ocellar Parapiophila vulgaris triangle) (Fig. 48). Apical tarsomere black (Fig. 49). Meron at least partly dull (Fig. 50). (Fallén)

7’ Frons entirely black (Fig. 51). Apical tarsomere always yellow (Fig. 52). Parapiophila flavipes Meron entirely glossy (Fig. 53). (Zetterstedt) doi:10.3752/cjai.2015.27 15 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

56 58

57 59 8 (4) Anepisternum bare (Fig. 56). Frons and gena entirely black (Fig. 57). Stearibia nigriceps Meigen

8’ Anepisternum setulose (Fig. 58). Frons and/or gena at least partially yellow 9 (Fig. 59).

doi:10.3752/cjai.2015.27 16 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

61 63

62 64

10 (9) Scutum with setulae forming three distinct rows (1 acrostichal, 2 Piophila casei dorsocentral) (Fig. 61). Male with long pale ventral setulae on hind tronchanter (Fig. 62). (Linnaeus)

10’ Scutum without distinct rows of setulae (Fig. 63). Male without long pale 11 ventral setulae on hind trochanter (Fig. 64).

doi:10.3752/cjai.2015.27 17 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

66 67 69

68 70

11(10) Pedicel at least twice as long as wide (longer in males than in females) (Fig. Prochyliza 66–67). Frons entirely black from lunule to occiput (Fig. 68). xanthostoma Walker 11’ Pedicel not elongated, as long as wide in both males and females (Fig. 69). 12 Frons yellow at least on anterior margin (Fig. 70).

doi:10.3752/cjai.2015.27 18 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

72 73

12 (11) Frons entirely yellow except for black ocellar triangle and posterior fronto- Prochyliza orbital plates (Fig. 72). nigrimana (Meigen) 12’ Frons with yellow portion restricted to anterior margin, never extending to 13 ocellar triangle and fronto-orbital plates (Fig. 73).

doi:10.3752/cjai.2015.27 19 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

75 77

76 78 13 (12) Anepimeron sparsely setulose (Fig. 75). Sternite 8 of male enlarged and Liopiophila convex (Fig. 76). Male sternites 1–4 with dense, coarse black setae (Fig. 76). varipes Meigen

13’ Anepimeron bare (Fig. 77). Sternite 8 of male not enlarged (Fig. 78). Male Prochyliza sternites 1–4 only with sparse setulae (Fig.78). brevicornis doi:10.3752/cjai.2015.27 Melander 20 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Liopiophila varipes Meigen

Body length: 2.3 – 4.4mm Wing length: 2.7 – 3.8mm

Variation: Colour of the mid tibia ranges from mostly black to mostly yellow.

Biology: In addition to its forensic relevance, this species is also considered a pest in the food industry (Zuska & Laštovka 1965). Bonduriansky & Brooks (1999b) and Bonduriansky (2003) studied sexual selection mechanisms and reproductive allocations in this species. Immature stages were described by Martín-Vega et al. (2014).

Nearctic distribution: Widespread in Canada from Yukon Territory and British Columbia to Nova Scotia, and Newfoundland and Labrador, and the United 79 States. Also present in Greenland (Duda 1924). Fig. 79. Liopiophila varipes (♂) (Examined specimens: 755 ♂, 378 ♀, 3?)

doi:10.3752/cjai.2015.27 21 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Mycetaulus bipunctatus complex

Body length: 2.3 – 3.5mm Wing length: 2.4 – 3.7mm

Variation: Colour of the head and thorax ranges from brown (Fig. 24–25) to yellow (Fig. 26–27) in this group.

Taxonomy: Several species (including some undescribed) will key to the Mycetaulus bipunctatus 24 25 complex (McAlpine 1977). These taxa mostly differ Fig. 24–25. Mycetaulus bipunctatus complex (♀) in genitalic characters of the males and/or females

with darker colour. 24. full view 25. thorax and are in need of revision. The Nearctic species M. costalis and M. longipennis are part of this group and will key out to this couplet, but these two species have not been associated with vertebrate carrion.

Nearctic distribution: Widespread in the Nearctic. Because several undescribed species are present in this complex, specific distribution information is not given. (Examined specimens: 234 ♂, 283 ♀, 8?) 26 27 Fig. 26–27. Mycetaulus bipunctatus complex (♂) doi:10.3752/cjai.2015.27with paler colour. 26. full view 27. thorax 22 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Mycetaulus subdolus (Johnson)

Body length: 2.0 – 3.3mm Wing length: 2.1 – 3.1mm

Variation: The anteriormost dorsocentral bristle is reduced and can occasionally be hard to distinguish from surrounding hairs. The apical spot is usually

separated into a spot covering R2+3 up to the costal vein and a smaller spot on R4+5. In some specimens, there is only one large apical spot which covers both

R2+3 and R4+5.

Nearctic distribution: Present in the following Canadian provinces: Ontario, Quebec. According to Johnson (1922), it is also present in Massachusetts and Vermont, USA. (Examined specimens: 41 ♂, 16 ♀)

28 Fig. 28. Mycetaulus subdolus (♂)

doi:10.3752/cjai.2015.27 23 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Parapiophila atrifrons (Melander & Spuler)

Body length: 1.9 – 2.8mm Wing length: 2.2 – 3.1mm

Variation: The palps, face and middle portion of the fore tibia may all range in colour from yellow to black in this species.

Taxonomy: Allopiophila calceata Duda, 1924 has recently been synonymized with P. atrifrons (Rochefort & Wheeler 2015). Specimens of “Parapiophila sp.1” recorded by Bonduriansky and Brooks (1999b) as occurring on carcasses have been examined and identified as P. atrifrons.

Nearctic distribution: Widespread throughout Canada from Yukon Territory and British Columbia to Nova Scotia, and Newfoundland and Labrador, and western United States in Alaska and from 47 Washington to New Mexico. Fig. 47. Parapiophila atrifrons (♀) (Examined specimens: 105 ♂, 175 ♀, 3?)

doi:10.3752/cjai.2015.27 24 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Parapiophila flavipes (Zetterstedt)

Body length: 1.9 – 2.7mm Wing length: 2.5 – 2.8mm

Variation: Antenna, face, lunule and palp are usually black, but can be partly yellow in some specimens. All femora are usually yellow but may be partly black.

Taxonomy: Specimens of “Parapiophila sp.2” recorded by Bonduriansky and Brooks (1999b) as occurring on carcasses and discarded antlers have been examined and identified as P. flavipes.

Nearctic distribution: In western Canada, present 55 in British Columbia, Yukon Territory, and Northwest Territories. In eastern Canada, present in Ontario, Fig. 55. Parapiophila flavipes (♀) Quebec, and Newfoundland and Labrador. In the United States, present in Alaska. (Examined specimens: 10 ♂, 74 ♀,1 ?)

doi:10.3752/cjai.2015.27 25 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Parapiophila vulgaris (Fallén)

Body length: 2.4 – 3.4mm Wing length: 2.5 – 3.3mm

Variation: Colour variation may occur on the head (i.e. gena, palp, antenna) and legs. Aberrations in chaetotaxy of the head such as an additional pair of vibrissae and/or verticals are common.

Biology: In addition to its forensic relevance, this species is also considered a pest species in the food industry (Zuska & Laštovka 1965).

Nearctic distribution: Widespread in Canada from Yukon Territory and British Columbia to Nova Scotia, and Newfoundland and Labrador, and the northern 54 United States (south to Colorado and West Virginia). Fig. 54. Parapiophila vulgaris (♂) Also present in Greenland according to Duda (1924) and specimens examined in USNM and CNC. (Examined specimens: 370 ♂, 374 ♀, 5?)

doi:10.3752/cjai.2015.27 26 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Piophila casei (Linnaeus)

Body length: 2.5 – 4.0mm Wing length: 2.5 – 3.9mm

Variation: The first flagellomere and the legs vary in colour in this species from yellow to black.

Biology: In addition to its forensic relevance, this cosmopolitan species is also considered a pest species in the food industry (Zuska & Laštovka 1965). The immature stages were described by Liu & Greenberg (1989) and Sukontason et al. (2001).

Nearctic distribution: Widespread in the United States and Canada but no arctic records were found in museum material examined in this study. It was, 65 however, reported from the Canadian Arctic (Malloch 1932), Alaska (Melander & Spuler 1917) and Fig. 65. Piophila casei (♂) Greenland (Melander & Spuler 1917, Duda 1924). (Examined specimens: 180 ♂, 88 ♀, 1?)

doi:10.3752/cjai.2015.27 27 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Prochyliza brevicornis Melander

Body length: 2.5 – 4.4mm Wing length: 2.7 – 3.8mm

Variation: This species is morphologically similar to Piophila casei. It can, however, be easily distinguished by the absence of three distinct rows of setulae on the scutum. Colour variation may occur on the legs.

Nearctic distribution: Widespread in Canada from Yukon Territory and British Columbia to Newfoundland and Labrador, and the United States. Also present in El Salto, Durango, Mexico. (Examined specimens: 218 ♂, 198 ♀, 1?)

80

Fig. 80. Prochyliza brevicornis (♂)

doi:10.3752/cjai.2015.27 28 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Prochyliza nigrimana (Meigen)

Body length: 2.7 – 3.7mm Wing length: 2.6 – 3.4mm

Variation: Colour variation in the gena, legs and abdomen occurs in this species. This variation, influenced by seasonality, separates the species into a pale morph (Fig. 74) and a dark morph which has a black gena, black on the front coxa, and mainly black mid and hind femora and tibiae (Martín-Vega & Baz 2011). Some specimens of the pale morph have also been observed with black colour on all femora.

Biology: In addition to its forensic relevance, this species is also considered a pest in the food industry (Zuska & Laštovka 1965). The immature 74 stages were described by Martín-Vega et al. (2012). Fig. 74. Prochyliza nigrimana (♂) Nearctic distribution: Widespread in Canada from Yukon Territory and British Columbia to Newfoundland and Labrador, and the United States. Also present in Greenland (Duda 1924). (Examined specimens: 58 ♂, 55 ♀ ) doi:10.3752/cjai.2015.27 29 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Prochyliza xanthostoma Walker

Body length: 2.9 – 4.4mm Wing length: 2.8 – 3.9mm

Variation: Colour of the fore and hind femora and tibiae ranges from partially to extensively black.

Biology: This species has been used in studies of sexual selection mechanisms, sexual dimorphism, mating behaviour and reproductive allocations (Bonduriansky & Brooks 1999b, Bonduriansky 2003, 2006, Bonduriansky & Rowe 2005, Bonduriansky et al. 2005.).The leaping behaviour of the larvae was described by Bonduriansky (2002).

71 Nearctic distribution: In Canada, present in British Columbia, Manitoba, New Brunswick, Ontario and Fig. 71. Prochyliza xanthostoma (♂) Quebec. In the United States, widespread. (Examined specimens: 322 ♂, 226 ♀, 2?)

doi:10.3752/cjai.2015.27 30 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Protopiophila latipes (Meigen)

Body length: 2.0 – 3.2mm Wing length: 2.0 – 2.8mm

Variation: All dorsocentral bristles are weak except for the posteriormost. In some specimens, the presutural dorsocentral bristle can be hard to distinguish from other hairs.

Biology: In addition to its forensic relevance, this species is also considered a pest species in the food industry (Zuska & Laštovka 1965). This species has also been used in studies of sexual selection mechanisms and reproductive strategies (Bonduriansky & Brooks 1999b, Bonduriansky 2003). 40 Fig. 40. Protopiophila latipes (♂) Nearctic distribution: In Canada, present in British Columbia and from Ontario to Nova Scotia (excluding Newfoundland and Labrador, and Prince Edward Island). In the United States, from Minnesota to New York, south to South Carolina. Also present in Greenland (Duda 1924). (Examined specimens: 733 ♂, 351 ♀, 5?) doi:10.3752/cjai.2015.27 31 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Protopiophila litigata Bonduriansky

Body length: 2.0 – 2.8mm Wing length: 1.9 – 2.6mm

Variation: All dorsocentral bristles are weak except for the posteriormost. Antenna colour varies from yellow to black.

Biology: This species has been used in studies of sexual selection mechanisms, as well as reproductive strategies and behaviors (Bonduriansky & Brooks 1998a, 1998b, 1999a, 1999b, Bonduriansky 2003).

41 Additional note: Two females of Protopiophila latipes in copula with males of P. litigata were found Fig 41. Protopiophila litigata (dissected ♂) in the University of Guelph Insect Collection, indicating that species limits between these two species should be further investigated.

Nearctic distribution: In Canada, present in Alberta, Saskatchewan, Ontario, Quebec Newfoundland and Labrador, and Nova Scotia. (Examined specimens: 234 ♂, 255 ♀) doi:10.3752/cjai.2015.27 32 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al. Stearibia nigriceps Meigen Body length: 2.5 – 4.1mm Wing length: 2.5 – 3.4 mm

Variation: This species is mostly glossy black, with a metallic blue tinge in some specimens. Colour variation in legs may occur; a specimen from Costa Rica is known to have entirely yellow mid and hind legs (Ozerov & Norrbom 2010).

Biology: In addition to its forensic relevance, this species is also considered a pest species in the food industry (Zuska & Laštovka 1965). It has also been used in studies of sexual selection mechanisms and reproductive strategies (Bonduriansky & Brooks 1999b, Bonduriansky 2003).

Nearctic distribution: Widespread in Canada from Yukon Territory and British Columbia to Nova Scotia, 60 and Newfoundland and Labrador, and the United Fig. 60. Stearibia nigriceps (♀) States. Also present in Greenland (Duda 1924). (Examined specimens: 499 ♂, 385 ♀, 4?)

doi:10.3752/cjai.2015.27 33 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Bonduriansky, R. 2003. Layered sexual selection: a Acknowledgements comparative analysis of sexual behaviour within an We thank Robert Loiselle (Université du Québec assemblage of piophilid flies. Canadian Journal of à Chicoutimi) and Eric Lucas (Université du Québec à Zoology, 81: 479–491. Montréal) who coordinated the Chicoutimi and Montreal field work, Jeff Skevington and Brad Sinclair (CNC), Bonduriansky, R. 2006. Convergent evolution of Valérie Lévesque-Beaudin (BIO), Steve Marshall and sexual shape dimorphism in Diptera. Journal of Steve Paiero (DEBU), Louise Cloutier (UMIC), Karen Morphology, 267: 602–611. Needham (UBCZ), Danny Shpeley (UASM) and Torsten Bonduriansky, R., and R.J. Brooks. 1998a. Copulation Dikow (USNM) for access to specimens, Stéphane and oviposition behaviour of Protopiophila litigata LeTirant (IMQC) for logistic support and access to (Diptera: Piophilidae). The Canadian Entomologist, specimens, and Karine Thivierge and Dominique St- 130: 399–405. Pierre (Laboratoire de Santé Publique du Québec) Bonduriansky, R., and R.J. Brooks. 1998b. Male antler for training and access to the stereomicroscope and flies (Protopiophila litigata; Diptera: Piophilidae) photography equipment. We thank G. Anderson, R. are more selective than females in mate choice. Bonduriansky and S. Marshall for helpful comments Canadian Journal of Zoology, 76: 1277–1285. on the manuscript. This research was funded by Natural Bonduriansky, R., and R.J. Brooks. 1999a. Why do male Sciences and Engineering Research Council of Canada antler flies Protopiophila( litigata) fight? The role of Discovery Grants to JS and TAW, a Bishop’s University male combat in the structure of mating aggregations Senate Research Grant to JS, and by support from the on moose antlers. Ethology, Ecology & Evolution Insectarium de Montréal to MG. 11: 287–301. References Bonduriansky, R., and R.J. Brooks. 1999b. Amendt, J., C.S. Richards, C.P. Campobasso, R. Zehner, Reproduction allocation and reproductive ecology and M.J.R. Hall. 2011. Forensic entomology: of seven species of Diptera. Ecological Entomology, Applications and limitations. Forensic Science, 24: 389–395. Medicine, and Pathology, 7: 379–392. Bonduriansky, R., and L. Rowe. 2005. Sexual selection, Anderson, G.S. 1995. The use of insects in death genetic architecture, and the condition dependence investigations: An analysis of cases in British of body shape in sexually dimorphic flyProchyliza Columbia over a five year period. Canadian Society xanthostoma (Piophilidae). Evolution, 59: 138–151. of Forensic Science Journal, 28: 277–292. Bonduriansky, R., E.J. Wheeler, and L. Rowe. 2005. Baumjohann, K., and Rudzinski, H.-G. 2013. Ejaculate feeding and female fitness in the sexually Bemerkenswerte Fliegen-Funde (Diptera) im dimorphic flyProchyliza xanthostoma (Diptera: Rahmen einer forensisch entomologischen Piophilidae). Animal Behaviour, 69: 489–497. Freilandstudie in Nordrhein-Westfalen Braack, L.E.O. 1986. Arthropod associated with (Deutschland). Studia Dipterologica, 19: 9–16. carcasses in the northern Kruger National Park. Battán Horenstein, M., A.X. Linhares, B. Rosso de South African Journal of Wildlife Research 16: Ferradas, and D. García. 2010. Decomposition 91–98. and dipteran succession in pig carrion in central Bucheli, S.R., J.A. Bytheway, S.M. Pustilnik, and J. Argentina: ecological aspects and their importance Florence. 2009. Insect successional pattern of a in forensic science. Medical and Veterinary corpse in cooler months of subtropical southeastern Entomology, 24: 16–25. Texas. Journal of Forensic Sciences, 54: 452–455. Bonduriansky, R. 1995. A new Nearctic species of Bygarski, K., and H.N. LeBlanc. 2013. Decomposition Protopiophila Duda (Diptera: Piophilidae), with and arthropod succession in Whitehorse, Yukon notes on its behaviour and comparison with P. Territory, Canada. Journal of Forensic Sciences, 58: latipes (Meigen). The Canadian Entomologist, 127: 413–418. 859–863. Byrd, J.H., and J.L. Castner. 2001. Insects of forensic Bonduriansky, R. 2002. Leaping behaviour and importance In: J.H. Byrd, and J.L. Castner (Eds). responses to moisture and sound in larvae of Forensic entomology: the utility of arthropods in piophilid carrion flies. The Canadian Entomologist, legal investigation. CRC, Boca Raton, FL. pp. 134: 647–656. 43–79.

doi:10.3752/cjai.2015.27 34 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Carvalho, L.M.L., P.J. Thyssen, A.X. Linhares, and de Jong, G.D., and J.W. Chadwick. 1999. F.A.B. Palhares. 2000. A checklist of arthropods Decomposition and arthropod succession on associated with pig carrion and human corpses in exposed rabbit carrion during summer at high Southeastern Brazil. Memórias do Instituto Oswaldo altitudes in Colorado, USA. Journal of Medical Cruz, Rio de Janeiro, 95: 135–138. Entomology, 36: 833–845. Catts, E.P., and M.L. Goff. 1992. Forensic entomology Johnson, C.W. 1922. New genera and species of in criminal investigations. Annual Review of Diptera. Occasional Papers of the Bostom Society of Entomology, 37: 253–272. Natural History. Cumming, J.M., and D.M Wood. 2009. Adult Johnson, M.D. 1975. Seasonal and microseral variations morphology and terminology. In: Brown, B.V., A. in the insect populations on carrion. American Borkent, J.M. Cumming, D.M. Wood, N.E. Woodley Midland Naturalist, 93: 79–90. & M. Zumbado (Eds). Manual of Central American Keh, B. 1985. Scope and applications of forensic Diptera. Volume 1. Ottawa, Canada: National entomology. Annual Review of Entomology, 30: Research Council Press. Pp. 2–50. 137–154. Duda, O. 1924. Revision der eropäischen u. Kumara, T.K., R.H.L. Disney, A.A. Hassan, M. Flores, Grönländischen sowie einiger südostasiat. Arten T.S. Hwa, Z. Mohamed, M.R. Chesalmah, and der Gattung Piophila Fallén (Dipteren). Konowia 3: S. Bhupinder. 2012. Occurrence of oriental flies 97–113, 153–203. associated with indoor and outdoor human remains Early, M., and M.I. Goff. 1986. Arthropod succession in the tropical climate of north Malaysia. Journal of patterns in exposed carrion on the island of O’ahu, Vector Ecology, 37: 62–68. Hawaiian Islands, USA. Journal of Medical Leclercq, M. 1996. A propos de l’entomofaune d’un Entomology, 24: 520–531. cadavre de sanglier. Bulletin et Annales de la Societé Fiedler, A., M. Halbach, B. Sinclair, and M. Benecke. Royale Entomologique de Belgique, 132: 417–422. 2008. What is the edge of a forest? A diversity Liu, D., and B. Greenberg. 1989. Immature stages of analysis of adult Diptera found on decomposing some flies of forensic importance. Annals of the piglets inside and on the edge of a western German Entomological Society of America, 82: 80–93. Woodland inspired by a courtroom question. Malloch, J.R. 1932. New species and other records Entomologie Heute, 20: 173–191. of Otitidae (Ortalidae), Piophilidae, Clusiidae, Gill, G.J. 2005. Decomposition and arthropod Chloropidae, and Drosophilidae from the succession on above ground pig carrion in rural Marquesas. Bernice P. Bishop Museum-Bulletin, 98: Manitoba. Technical Report (TR-06-2005). 205–223. Canadian Police Research Centre. 178pp. Martinez, E., P. Duque, and M. Wolff. 2006. Succession Goff, M.L. 2000. A fly for the prosecution: How insect pattern of carrion-feeding insects in Paramo, evidence helps solve crimes. Harvard University Colombia. Forensic Science International, 166: Press, Cambridge. 182–189. Greenberg, B. 1991. Flies as forensic indicators. Journal Martín-Vega, D. 2011. Skipping clues: Forensic of Medical Entomology, 28: 565–577. importance of the family Piophilidae (Diptera). Grisales, D., M. Ruiz, and S. Villegas. 2010. Insects Forensic Science International, 212: 1–5. associated with exposed decomposing bodies in Martín-Vega, D. 2014. On the identity of Prochyliza the Colombian Andean Coffee Region. Revista nigrimana (Meigen) and Prochyliza nigricornis Brasileira de Entomologia, 54: 637–644. (Meigen) (Diptera: Piophilidae), with a synopsis of Hadley, A. 2008. CombineZP Image Stacking Software. Prochyliza Walker and description of a new species. Retrieved July 10, 2014, from http://www. Zootaxa, 3893: 277–292. hadleyweb.pwp.blueyonder.co.uk/ Martín-Vega, D., and A. Baz. 2011. Variation in Jirón, L.F., and V.M. Cartín. 1981. Insect succession the colour of the necrophagous fly, Prochyliza in the decomposition of a mammal in Costa Rica. nigrimana (Diptera: Piophilidae): A case of seasonal Journal of the New York Entomological Society, 89: polymorphism. European Journal of Entomology, 158–165. 108: 231–234.

doi:10.3752/cjai.2015.27 35 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Martín-Vega, D., and A. Baz. 2013. Sarcosaprophagous Ozerov, A.L. 1989. Sepsidae and Piophilidae (Diptera) Diptera assemblages in natural habitats in Central of the Zeya State Reserve. Translated from Spain: Spatial and seasonal changes in composition. Entomologicheskoye Obozreniye, 4: 839–849. Medical and Veterinary Entomology, 27: 64–76. Ozerov, A.L. 2000. A new species of the genus Martín-Vega, D., A. Baz, and L.M. Díaz-Aranda. 2012. Actenoptera (Diptera, Piophilidae) from the Russian The immature stages of the necrophagous fly, Far East. Entomological Review, 80: 244–245. Prochyliza nigrimana: comparison with Piophila Ozerov, A.L. 2002. First record of Neopiophila casei and medicolegal considerations (Diptera: McAlpine, 1977 (Diptera: Piophilidae) in Piophilidae). Parasitology Research, 111: 1127– Palaearctic, with description of a new species. 1135. Russian Entomological Journal, 11: 223–224. Martín-Vega, D., A. Gómez-Gómez, A. Baz, and L.M. Ozerov, A.L. 2004. On the classification of the family Díaz-Aranda. 2011. New piophilid in town: the first Piophilidae (Diptera). Entomological Review, 84: Palaearctic record of Piophila megastigmata and its 600–608. coexistence with Piophila casei in Central Spain. Ozerov, A.L. 2007. Three new species of Carnidae Medical and Veterinary Entomology, 25: 64–69. and Piophilidae (Diptera) from Turkey and Russia. Martín-Vega, D., L.M. Diaz-Aranda., and A. Baz. Russian Entomological Journal, 16: 491–493. 2014. The immature stages of the necrophagous Ozerov, A.L. and M. Barták. 1993. Two new Palaearctic fly Liopiophila varipes and considerations on the species of Piophilidae (Diptera). Russian genus Liopiophila (Diptera: Piophilidae). Deutsche Entomological Journal, 2: 73-76. Entomologische Zeitschrift, 61: 37–42. Ozerov, A.L. and A.L. Norrbom. 2010. Piophilidae. Matuszewski, S., D. Bajerlein, S. Konwerski, and K. In: Brown, B.V., A. Borkent, J.M. Cumming, D.M. Szpila. 2008. An initial study of insect succession Wood, N.E. Woodley & M. Zumbado (Eds). Manual and carrion decomposition in various forest habitats of Central American Diptera. Volume 2. Ottawa, of Central Europe. Forensic Science International, Canada: National Research Council Press. Pp. 180: 61–69. 865–869. Merz, B. 1996. Die Piophilidae (Diptera) der Schweiz Prado e Castro, C., and M.D. García. 2010. Additions to mit Beschreibung einer neuen Art. Bulletin de la the Piophilidae (Diptera) fauna from Portugal, with Société Entomologique Suisse, 69: 345–360. new records. Graellsia, 66: 101–105. McAlpine, D.K. 1989. A synopsis of the Australian Prado e Castro, C., E. Cunha, A. Serrano, and M.D. Piophilidae (Diptera: Schizophora). General and García. 2012. Piophila megastigmata (Diptera: Applied Entomology, 21: 17–24. Piophilidae): First records on human corpses. McAlpine, J.F. 1977. A revised classification of the Forensic Science International, 214: 23–26. Piophilidae, including ‘Neottiophilidae’ and Paños, A., M.I. Arnaldos, M.D. García, and N. Ubero- ‘Thyreophoridae’ (Diptera: Schizophora). Memoirs Pascal. 2013. Ultrastructure of preimaginal of the Entomological Society of Canada, 103: 1–66. stages of Piophila megastigmata McAlpine, 1978 McAlpine, J.F. 1978. A new species of Piophila from (Diptera, Piophilidae): A fly of forensic importance. South Africa (Diptera: Piophilidae). Annals of the Parasitology Research, 112: 3771–3788. Natal Museum, 23: 455–459. Pape, T., D. Bickel, and R. Meier (Eds). 2009. Appendix Megnin, P. 1894. La faune des cadavres – Application A: Species of Diptera per family for all regions. In: de l’entomologie à la médecine légale. Encyclopedie Diptera Diversity: Status, challenges and tools. Brill, Scientifique des Aide-Mémoire, 214 pp. G. Masson, Leiden: Boston. Pp. 439–444. Gauthier-Villars et fils, Paris. Reed, H.B. 1958. A study of dog carcass communities Melander, A.L., and A. Spuler. 1917. The dipterous in Tennessee, with special reference to the insects. families Sepsidae and Piophilidae. Washington American Midland Naturalist, 59: 213–245. Agricultural Experiment Station Bulletin: 97. Rochefort, S., and T.A. Wheeler. 2015. Diversity Michaud, J.-P., C.G. Majka, J.-P. Privé, and G. Moreau. of Piophilidae (Diptera) in northern Canada 2010. Natural and anthropogenic changes in the and description of a new Holarctic species of insect fauna associated with carcasses in the North Parapiophila McAlpine. Zootaxa (manuscript in American Maritime lowlands. Forensic Science revision). International, 202: 64–70.

doi:10.3752/cjai.2015.27 36 Canadian Journal of Arthropod Identification No. 27 (January, 2015) Rochefort et al.

Sathe, T.V., A. Sathe, and N.T. Sathe. 2013. Diversity Valásquez, T., C. Magaña, A. Martinez-Sánchez, and S. of dipterous forensic insects from Western Rojo. 2010. Diptera of forensic importance in the Maharashtra, India. International Journal of Pharma Iberian Peninsula: Larval identification key. Medical and Bio Sciences, 4(2): P173–P179. and Veterinary Entomology, 24: 293–308. Schoenly, K.G., N.H. Haskell, R.D. Hall, and J.R. Voss, S.C., S.L. Forbes, and I.R. Dadour. 2008. Gbur. 2007. Comparative performance and Decomposition and insect succession on cadavers complementarity of four sampling methods and inside a vehicle environment. Forensic Science, arthropod preference tests from human and porcine Medicine and Pathology, 4: 22–32. remains at the Forensic Anthropology Center Watson, E.J., and C.E. Carlton. 2003. Spring succession in Knoxville, Tennessee. Journal of Medical of necrophilous insects on wildlife carcasses in Entomology, 44: 881–894. Louisiana. Journal of Medical Entomology, 40: Sharanowski, B.J., E.G. Walker, and G.S. Anderson. 338–347. 2008. Insect succession and decomposition patterns Wells, J.D., and R.S. Stevens. 2008. Application of on shaded and sunlit carrion in Saskatchewan DNA-based methods in forensic entomology. in three different seasons. Forensic Science Annual Review of Entomology, 53: 103–120. International, 179: 219–240. Whitworth, T.L., R.D. Dawson, H. Magalon, and E. Soós, A. 1977. Acalyptrata (Diptera) from Afghanistan Baudry. 2007. DNA barcoding cannot reliably II. Folia Entomologica Hungarica, XXX: 133–137. identify species of the blowfly genus Protocalliphora Sukontason, K.L., K. Sukontason, S. Piangjai, W. (Diptera: Calliphoridae). Proceedings of the Royal Choochote, R.C. Vogtsberger, and J.K. Olson. Society, B, 274: 1731–1739. 2001. Scanning electron microscopy of the third- Zuska, J., and P. Laštovka. 1965. A review of the instar Piophila casei (Diptera: Piophilidae), a fly Czechoslovak species of the family Piophilidae species of forensic importance. Journal of Medical with special reference to their importance to food Entomology, 38: 756–759. industry (Diptera, Acalyptrata). Acta Entomologica Tabor, K.L., R.D. Fell, and C.C. Brewster. 2005. Bohemoslovaca, 62: 141–157. Insect fauna visiting carrion in southwest Virginia. Forensic Science International, 150: 73–80.

doi:10.3752/cjai.2015.27 37