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

Key to Forensicaffy Important Piophifidae (Diptera) in the Nearctic Region Sabrina Rochefort 1, Marjofaine Giroux 2, Jade Savage 3 and Terry A. Wheefer 1

1Department of Naturaf Resource Sciences, McGiff University, Macdonafd Campus, Ste-Anne-de-Beffevue, QC, H9X 3V9, Canada; [email protected], [email protected] 2Montréaf Insectarium / Space for fife, 4581, rue Sherbrooke Est, Montréaf, QC, H1X 2B2, Canada; [email protected] 3Biofogicaf Sciences, Bishop’s University, 2600 Coffege Street, Sherbrooke, QC, J1M 1Z7, Canada [email protected];

Abstract Many of Piophifidae (Diptera) are refevant to forensic entomofogy because their presence on a corpse can be hefpfuf in estimating the postmortem intervaf (PMI) and document succession. The aims of this paper are to document the fauna of forensicaffy refevant Piophifidae species worfdwide and to present an updated checkfist and identii cation key to the Nearctic species, as existing keys are either outdated, too broad in geographicaf scope to be user-friendfy, and/or contain ambiguous characters. Thirteen species are incfuded in the checkfist and key. Information on their biofogy, , character variabifity, and distribution is provided, suppfementing the extensive work of McAfpine (1977).

Introduction stages (Martinez et af. 2006, Grisafes et af. 2010). Forensic entomofogy is the use of and other Identifying species of forensic importance can as evidence in fegaf investigations (Catts sometimes be chaffenging when using morphofogicaf & Goff 1992). An important aspect of the discipfine characters afone (Byrd & Castner 2001, Amendt et af. invofves the estimation of the postmortem intervaf (PMI) 2011) and afternatives such as DNA markers have been based on arthropods associated with a body, an approach devefoped to identify probfematic specimens (Weffs that requires extensive knowfedge of the focaf fauna and & Stevens 2008). Whife mofecufar techniques can be its association with carrion at different decomposition a usefuf compfement to morphofogy-based specimen stages and under different conditions (Goff 2000, identii cation, they do not afways yiefd correct or Amendt et af. 2011). Necrophagous fl ies (Diptera) are of unambiguous identii cation resufts (e.g. Whitworth et forensic importance as they often appear on a body i rst, af. 2007) and require access to equipment, facifities and consume most of the tissues, and dispfay simifar patterns funding for mofecufar anafysis; thus, morphofogy-based of succession in different regions of the worfd, at feast at regionaf keys remain time- and cost-effective toofs for the famify fevef (Catts & Goff 1992, Amendt et af. 2011). the identii cation of most forensicaffy important species. The famify Piophifidae contains 82 species worfdwide McAfpine (1977) contributed signii cantfy to the (Pape et af. 2009), at feast 37 of which are present in the systematics of the famify Piophifidae through the Nearctic region (McAfpine 1977). Severaf species can proposaf of a revised cfassii cation of the famify (afthough be found on dung, bone, garbage, decaying vegetation, this was not based on an expficit phyfogenetic anafysis), fungi, bird nests and discarded antfers (Mefander & description of six new species and two new genera, Spufer 1917, McAfpine 1977, Bonduriansky & Brooks taxonomic changes to other species, the documentation 1999a). Other species are associated with carrion in of species distributions, and the pubfication of worfdwide a range of decomposition stages and are therefore identii cation keys to species. McAfpines’s work, refevant to forensic entomofogy (Greenberg 1991, however, was pubfished more than 35 years ago, and his Byrd & Castner 2001). Depending on the focafity, aduft species keys can be difi cuft to use, especiaffy by non- Piophifidae are mostfy seen throughout the bfoated and experts, as they are based mostfy on cofour characters decay stages (Johnson 1975, Fiedfer et af. 2008, Prado e and do not incfude species described since that time. Castro et af. 2012) whife the farvae are more common in In an effort to compfement and update McAfpine the advanced (Martín-Vega et af. 2011) and dry/remain (1977), the objectives of this work are to document

doi:10.3752/cjai.2015.27 1 Canadian Journaf of Arthropod Identiication No. 27 (January, 2015) ROCHEFO RT ET AL .

Piophifidae species of known or potentiaf forensic revised cfassiication of the Piophifidae is required. importance worfdwide, to compife a checkfist of Nearctic Photographs for the key were taken with an Piophifidae species refevant to forensic entomofogy Ofympus DP71 camera mounted on an Ofympus SZX16 based on pubfished records in the fiterature and new data stereoscope. Images were captured and stacked using from Quebec (Canada), to update the distribution records DP Controffer and Combine ZP (Hadfey 2008) before and document the intraspeciic variations of aff species being enhanced using Adobe Photoshop [CS3] (Adobe in the Nearctic checkfist, and to devefop a user-friendfy Systems, Mountain view, CA). key to forensicaffy refevant Piophifidae species in the Nearctic. Identiication chaffenges in the Piophifidae The key to the worfd species of Piophifidae by Materiafs and Methods McAfpine (1977) remains the most compfete work for The checkfist of Nearctic Piophifidae of forensic the identiication of adufts even though severaf new importance was compifed from the fiterature and from species have since been described (Soós 1977, McAfpine specimens coffected in the context of a study of the 1978, McAfpine 1989, Ozerov 1989, Ozerov & Barták insect fauna associated with nine pig carcasses in three 1993, Bonduriansky 1995, Merz 1996, Ozerov 2000, semi-urban sites focated afong a fatitudinaf gradient in 2002, 2004, 2007, Martín-Vega 2014, Rochefort & Quebec (Canada) in summer 2011 by Giroux, Savage Wheefer 2015) and additionaf distribution records are and coffaborators. The sampfing sites were Ste-Anne- known based on museum specimens. McAfpine’s (1977) de-Beffevue (45.436°, -73.909°) (22 June to 14 August), key is not ideaf to identify piophifids in forensic studies Sherbrooke (45.361°, -71.844°) (6 Jufy to 26 August) as it incfudes many non-forensicaffy important species and Saguenay (La Baie) (48.350°, -70.967°) (4 Jufy to and is mostfy based on cofour differences which can be 19 August). troubfesome in this famify due to intraspeciic cofour Approximatefy 1065 piophifid specimens from these variations. Such pofymorphisms have been documented three sites were examined. They were identiied to species in severaf species of forensic interest such as Parapiophifa using McAfpine (1977) and compared with reference atrifrons (Mefander & Spufer) (Rochefort & Wheefer specimens in the Canadian Nationaf Coffection of Insects, 2015), Parapiophifa vufgaris (Faffén) (McAfpine 1977) Arachnids and Nematodes, Ottawa, ON (CNC) and the and Prochyfiza nigrimana (Meigen) (Martín-Vega & Lyman Entomofogicaf Museum, McGiff University, Ste- Baz 2011). Some species are afso morphofogicaffy very Anne-de-Beffevue, QC (LEM). Voucher specimens were simifar. In southern Europe, for exampfe, the very simifar deposited in the Insectarium de Montréaf’s scientiic Piophifa casei (L.) and Piophifa megastigmata McAfpine coffections (IMQC) and the Bishop’s University Insect co-occur; this can cause identiication errors that may Coffection (BUIC). fead to erroneous PMI estimations (Martín-Vega 2011). In addition, approximatefy 5600 specimens of Confusion between P. megastigmata and P. casei has Piophifidae incfuded in the key were examined to occurred in forensic research (e.g., Prado e Castro 2010), document geographic distribution and intraspeciic a mistake that might actuaffy be widespread in the iefd variabifity. These specimens are deposited in CNC; LEM; (Prado e Castro et af. 2012), and we suspect that such the Institute of Ontario, Guefph, ON, (BIO); confusion may afso appfy to other species pairs in the the University of Guefph Insect Coffection, Guefph, Nearctic. ON, (DEBU); the Oueffet-Robert Coffection, Université Some forensic studies fimit Piophifidae identiication de Montréaf, Montréaf, QC (UMIC); the Spencer to the famify or generic fevefs (e.g., Schoenfy et af. 2007, Entomofogicaf Coffection, Beaty Biodiversity Museum, Voss et af. 2008, Vefasquez et af. 2010, Bygarski & Vancouver, BC (UBCZ); the Strickfand Museum, Lebfanc 2013) or identify the materiaf as Piophifa casei , University of Afberta, Edmonton, AB (UASM); and despite the fact that severaf species have been documented the United States Nationaf Museum of Naturaf History, in forensic fiterature worfdwide (Tabfe 1). Piophifa casei Washington DC, (USNM). Additionaf pubfished records is a weff-known, cosmopofitan, synanthropic species and were added where necessary. was the irst piophifid documented in forensic studies Cfassiication of genera and species foffows (Megnin 1894). This may account for the possibifity that McAfpine (1977) for consistency with the Nearctic Piophifidae coffected in forensic studies are frequentfy fiterature. Ozerov (2004) proposed a different generic identiied (or misidentiied) as Piophifa casei (see cfassiication, foffowed by some authors, especiaffy in Martín-Vega 2011). Europe, but that cfassiication, fike McAfpine’s, was not based on a phyfogenetic anafysis and thus is no better supported. A comprehensive phyfogenetic anafysis and

doi:10.3752/cjai.2015.27 2 Canadian Journaf of Arthropod Identiication No. 27 (January, 2015) ROCHEFO RT ET AL .

Tabfe 1. Piophifidae species documented in the forensic entomofogy fiterature.

Species Locafity References

Boreopiophifa tomentosa Frey, 1930* Canada : Manitoba Giff 2005

Liopiophifa varipes Meigen, 1830 Canada : New Brunswick Michaud et af. 2010

Germany Fiedfer et af. 2008 Baumjohann & Rudzinski 2013

Mycetaufus bipunctatus (Faffén, 1823) USA : Louisiana Watson & Carfton 2008

Parapiophifa vufgaris (Faffén, 1820) Germany Fiedfer et af. 2008 Baumjohann & Rudzinski 2013

Pofand Matuszewski et af. 2008

Piophifa casei (Linnaeus, 1758) Canada : Saskatchewan Sharanowski et af. 2008

USA : Tennessee Reed 1958**

USA : Hawaii Earfy & Goff 1986

USA : Coforado de Jong & Chadwick 1999***

USA : Louisiana Watson & Carfton 2008

Costa Rica Carvafho et af. 2000

Argentina Battán Horenstein et af. 2010

Spain Martín-Vega et af. 2011

Portugaf Prado e Castro et af. 2012

Germany Baumjohann & Rudzinski 2013

South Africa Braack 1986

Thaifand Sukontason et af. 2001

Mafaysia Kumara et af. 2012

Piophifa megastigmata McAfpine, 1978 Portugaf Prado e Castro et af. 2012

Spain Martín-Vega et af. 2011

Paños et af. 2013

South Africa Braack 1986

Prochyfiza azteca McAfpine, 1977 Costa Rica Jirón & Cartín 1981

doi:10.3752/cjai.2015.27 3 Canadian Journaf of Arthropod Identiication No. 27 (January, 2015) ROCHEFO RT ET AL .

Tabfe 1 cont’d. Piophifidae species documented in the forensic entomofogy fiterature.

Prochyfiza brevicornis Mefander, 1924 Canada: British Cofumbia Anderson 1995

Prochyfiza nigrimana (Meigen, 1826) USA : Tennessee Reed 1958**

Portugaf Prado e Castro et af. 2012

Spain Martín-Vega et af. 2011

Martín-Vega & Baz 2013

Prochyfiza xanthostoma Wafker, 1849 Canada : Manitoba Giff 2005

USA : Tennessee Reed 1958**

USA : Iffinois Johnson 1975****

USA : Louisiana Watson & Carfton 2003

USA : V irginia Tabor et af. 2005

Protopiophifa fatipes (Meigen, 1838) Canada : New Brunswick Michaud et af. 2010

USA : Tennessee Reed 1958**

USA : Iffinois Johnson 1975****

Germany Fiedfer et af. 2008 Baumjohann & Rudzinski 2013

Portugaf Prado e Castro et af. 2012

Stearibia nigriceps Meigen, 1826 Canada : British Cofumbia Anderson 1995

Canada : Manitoba Giff 2005

Canada : New Brunswick Michaud et af. 2010

USA : T ennessee Reed 1958**

USA : Louisiana Watson & Carfton 2003

USA : V irginia Tabor et af. 2005

France Lecfercq 1996

Germany Fiedfer et af. 2008 Baumjohann & Rudzinski 2013

Pofand Matuszewski et af. 2008

Spain Martín-Vega et af. 2011

doi:10.3752/cjai.2015.27 4 Canadian Journaf of Arthropod Identiication No. 27 (January, 2015) ROCHEFO RT ET AL .

Tabfe 1 cont’d. Piophifidae species documented in the forensic entomofogy fiterature.

Stearibia nigriceps Meigen, 1826 cont’d Portugaf Prado e Castro et af. 2012

Martinez et af. 2006

Cofombia Grisafes et af. 2010

India Sathe et af. 2013 * Giff (2005) recorded B. tomentosa from southwestern Manitoba. Aff veriied records of this species are from the arctic (Rochefort & Wheefer 2015) and vouchers of Giff’s specimens are unavaifabfe. Thus we consider this record questionabfe. The foffowing studies did not use human or pig carcasses. ** Used dog carcasses *** Used rabbit carcasses. **** Used squirref, rabbit, cat and opossum carcasses.

Checkfist of Nearctic Piophifidae of forensic interest Parapiophifa atrifrons (Mefander & Spufer, 1917) Thirteen species are incfuded in our Nearctic checkfist. Parapiophifa favipes (Zetterstedt 1847) Nine of these were compifed from the fiterature (Tabfe 1), Parapiophifa vufgaris (Faffén, 1820) two from museum records, and two, Mycetaufus subdofus Piophifa casei (Linnaeus, 1758) (Johnson 1922) and Parapiophifa atrifrons (Mefander & Prochyfiza brevicornis Mefander, 1924 Spufer 1917), were recorded for the irst time from pig Prochyfiza nigrimana (Meigen, 1826) carcasses in Quebec (afong with four other species whose Prochyfiza xanthostoma Wafker, 1849 forensic refevance was afready known). Parapiophifa Protopiophifa fatipes (Meigen, 1838) favipes (Zetterstedt 1847) and Protopiophifa fitigata Protopiophifa fitigata Bonduriansky 1995 Bonduriansky 1995 have been added to the Nearctic Stearibia nigriceps Meigen, 1826 checkfist since many specimens from DEBU have been coffected on carcasses, an indication that they may be Key to the Nearctic Piophifidae of forensic interest of forensic refevance. Parapiophifa vufgaris has afso Morphofogicaf terminofogy foffows Cumming & been added to the Nearctic checkfist because it is widefy Wood (2009). Aff measurements were taken on dry distributed in the region and is a species of forensic specimens; body fength was measured from the front refevance in Europe (Fiedfer et af. 2008, Matuszewski of the head (excfuding the antenna) to the end of the et af. 2008). Parapiophifa specimens identiied onfy to abdomen; wing fength was measured from base to apex. in severaf studies (e.g. Sharanowski et af. 2008, and Hefeomyzidae are two other acafyptrate Michaud et af. 2010) coufd befong to this species, Diptera famifies often coffected in farge numbers afthough they may afso correspond to other species such on carcasses. They are sometimes misidentiied as as Parapiophifa atrifrons and P. favipes . Boreopiophifa a Piophifidae because of simifarity in the overaff tomentosa was excfuded from the checkfist based on its appearance of some species. Characters to distinguish dubious record in Giff (2005) (see footnote in Tabfe 1). these two famifies from Piophifidae are incfuded in the Liopiophifa varipes Meigen, 1830 identiication key. Mycetaufus bipunctatus (Faffén, 1823) compfex Mycetaufus subdofus (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)

Differentiating from Arista inserted PIOPHILIDAE Sepsidae and 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 specimens have a short seta 8 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. 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 in British Columbia, Yukon Territory, and Northwest 55 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, however, reported from the Canadian Arctic (Malloch 65 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. (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. 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, , 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 In Canada, present in British Fig. 40. Protopiophila latipes (♂) Nearctic distribution: 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 Journaf of Arthropod Identiication No. 27 (January, 2015) ROCHEFO RT ET AL .

Bonduriansky, R. 2003. Layered sexuaf sefection: a Acknowfedgements comparative anafysis of sexuaf behaviour within an We thank Robert Loiseffe (Université du Québec assembfage of piophifid fies. Canadian Journaf of à Chicoutimi) and Eric Lucas (Université du Québec à Zoofogy, 81: 479–491. Montréaf) who coordinated the Chicoutimi and Montreaf iefd work, Jeff Skevington and Brad Sincfair (CNC), Bonduriansky, R. 2006. Convergent evofution of Vaférie Lévesque-Beaudin (BIO), Steve Marshaff and sexuaf shape dimorphism in Diptera. Journaf of Steve Paiero (DEBU), Louise Cfoutier (UMIC), Karen Morphofogy, 267: 602–611. Needham (UBCZ), Danny Shpefey (UASM) and Torsten Bonduriansky, R., and R.J. Brooks. 1998a. Copufation Dikow (USNM) for access to specimens, Stéphane and oviposition behaviour of Protopiophifa fitigata LeTirant (IMQC) for fogistic support and access to (Diptera: Piophifidae). The Canadian Entomofogist, specimens, and Karine Thivierge and Dominique St- 130: 399–405. Pierre (Laboratoire de Santé Pubfique du Québec) Bonduriansky, R., and R.J. Brooks. 1998b. Mafe antfer for training and access to the stereomicroscope and fies ( Protopiophifa fitigata ; Diptera: Piophifidae) photography equipment. We thank G. Anderson, R. are more sefective than femafes in mate choice. Bonduriansky and S. Marshaff for hefpfuf comments Canadian Journaf of Zoofogy, 76: 1277–1285. on the manuscript. This research was funded by Naturaf Bonduriansky, R., and R.J. Brooks. 1999a. Why do mafe Sciences and Engineering Research Councif of Canada antfer fies ( Protopiophifa fitigata ) ight? The rofe of Discovery Grants to JS and TAW, a Bishop’s University mafe combat in the structure of mating aggregations Senate Research Grant to JS, and by support from the on moose antfers. Ethofogy, Ecofogy & Evofution Insectarium de Montréaf to MG. 11: 287–301. References Bonduriansky, R., and R.J. Brooks. 1999b. Amendt, J., C.S. Richards, C.P. Campobasso, R. Zehner, Reproduction affocation and reproductive ecofogy and M.J.R. Haff. 2011. Forensic entomofogy: of seven species of Diptera. Ecofogicaf Entomofogy, Appfications and fimitations. Forensic Science, 24: 389–395. Medicine, and Pathofogy, 7: 379–392. Bonduriansky, R., and L. Rowe. 2005. Sexuaf sefection, Anderson, G.S. 1995. The use of insects in death genetic architecture, and the condition dependence investigations: An anafysis of cases in British of body shape in sexuaffy dimorphic fy Prochyfiza Cofumbia over a ive year period. Canadian Society xanthostoma (Piophifidae). Evofution, 59: 138–151. of Forensic Science Journaf, 28: 277–292. Bonduriansky, R., E.J. Wheefer, and L. Rowe. 2005. Baumjohann, K., and Rudzinski, H.-G. 2013. Ejacufate feeding and femafe itness in the sexuaffy Bemerkenswerte Ffiegen-Funde (Diptera) im dimorphic fy Prochyfiza xanthostoma (Diptera: Rahmen einer forensisch entomofogischen Piophifidae). Animaf Behaviour, 69: 489–497. Freifandstudie in Nordrhein-Westfafen Braack, L.E.O. 1986. Arthropod associated with (Deutschfand). Studia Dipterofogica, 19: 9–16. carcasses in the northern Kruger Nationaf Park. Battán Horenstein, M., A.X. Linhares, B. Rosso de South African Journaf of Wifdfife Research 16: Ferradas, and D. García. 2010. Decomposition 91–98. and dipteran succession in pig carrion in centraf Buchefi, S.R., J.A. Bytheway, S.M. Pustifnik, and J. Argentina: ecofogicaf aspects and their importance Fforence. 2009. Insect successionaf pattern of a in forensic science. Medicaf and Veterinary corpse in coofer months of subtropicaf southeastern Entomofogy, 24: 16–25. Texas. Journaf of Forensic Sciences, 54: 452–455. Bonduriansky, R. 1995. A new Nearctic species of Bygarski, K., and H.N. LeBfanc. 2013. Decomposition Protopiophifa Duda (Diptera: Piophifidae), with and arthropod succession in Whitehorse, Yukon notes on its behaviour and comparison with P. Territory, Canada. Journaf of Forensic Sciences, 58: fatipes (Meigen). The Canadian Entomofogist, 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 farvae of Forensic entomofogy: the utifity of arthropods in piophifid carrion fies. The Canadian Entomofogist, fegaf investigation. CRC, Boca Raton, FL. pp. 134: 647–656. 43–79.

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