Diptera: Syrphidae), Including Description of a New Species

216 Review of New World Sericomyia (Diptera: Syrphidae), including description of a new species

Jeffrey H. Skevington,1 F. Christian Thompson

Abstract—The 19 New World species of Sericomyia Meigen are reviewed, including one species new to North America (Sericomyia jakutica (Stackelberg)) and one previously undescribed species (Sericomyia vockerothi Skevington sp. nov. from Alberta, Minnesota, Northwest Territories, Quebec, and Yukon Territory). Mallota powelli Nayar and Cole is recognized as a junior synonym of Sericomyia ﬂagrans (Osten Sacken). A description and illustrations of S. vockerothi and an illustrated key to New World Sericomyia are presented. DNA barcode data are presented for 14 New World species and a cytochrome oxidase subunit I gene tree is presented and discussed. Genetic evidence supports the contention that the subgenera of Sericomyia are not monophyletic. Arctophila Schiner and Conosyprhus Frey are thus proposed as junior synonyms of Sericomyia.

Re´sume´—Nous re´visons les 19 espe`ces de Sericomyia Meigen du Nouveau Monde, y compris une espe`ce signaleé pour la premie`re fois en Ame´rique du Nord (Sericomyia jakutica (Stackelberg)) et une espe`ce encore non dećrite (Sericomyia vockerothi Skevington sp. nov. d’Alberta, du Minnesota, des Territoires du Nord-Ouest, du Que´bec et du territoire du Yukon). Mallota powelli Nayar et Cole devient un synonyme plus rećent de Sericomyia flagrans (Osten Sacken). Nous pre´sentons une description et des illustrations de S. vockerothi, ainsi qu’un cle´ illustreé pour l’identification des Sericomyia du Nouveau Monde. Des donneéssurlescodesa`barresgeńe´tiques sont fournies pour 14 espe`ces du Nouveau Monde, de meˆme qu’un arbre geńique de la CO1 qui fait l’objet d’une discussion. Les donneés geńe´tiques appuient la proposition que les sous-genres de Sericomyia ne sont pas monophyle´tiques. Nous proposons donc Arctophila Schiner et Conosyrphus Frey comme synonymes plus rećents de Sericomyia.

Introduction Sericomyiine flower flies are common in boreal forests across the Holarctic region and We are pleased to be able to contribute to this extend southward at higher elevations into the Festschrift celebrating the impact of the Manual Oriental and Neotropical regions. Thirty species of Nearctic Diptera and its coordinators. Because were known at the inception of this project: 10 this is a paper on flower flies, we intend to focus on restricted to the Palaearctic, 13 restricted to the Richard (Dick) Vockeroth. Dick was one of the Nearctic, two Holarctic, three from the Orient, pre-eminent dipterists of the 20th century and had and two from the Neotropics. tremendous impact on syrphid taxonomy. His In this paper we describe one new species of species concepts (whether published or not) are Sericomyia, highlight the presence of a Holarctic tremendously insightful and his keys covering species not previously known from North America, many families are among the best ever written for recognize a misplaced and junior synonym, provide fly identification. It will be decades before we are new data on the difficult to identify and often finished compiling his collection data and notes Holarctic northern species, illustrate the genitalia of into published papers. Here, we have incorporated some species for the first time, provide habitus his unpublished work on Sericomyia Meigen with illustrations for all of the species, and provide the our own views and some new data. first key to the group since Curran (1934).

Received 28 January 2011. Accepted 4 April 2011. J.H. Skevington,1 Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, 960 Carling Avenue, Ottawa, ON K1A 0C6, Canada F.C. Thompson, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012, United States of America 1Corresponding author (e-mail: [email protected]). doi:10.4039/tce.2012.24

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Sericomyia larvae are rat-tailed maggots and number (see above). Two Palaearctic and 14 New live in ponds rich in decomposing vegetation World species of Sericomyia were successfully where they filter out microorganisms as their sequenced. Data are missing for the following food (Rotheray 1993; van Veen 2004). Adults New World species: Sericomyia arctica Schirmer, are regularly found on flowers and many are also Sericomyia carolinensis (Metcalf), Sericomyia conspicuous members of the hilltopping com- harveyi (Osburn), Sericomyia meyeri (Fluke), and munity (J.H. Skevington, unpublished data). Sericomyia fairmanorum Fairman. Attempts were made to sequence these species but our material Materials and methods was too old to amplify or had been prepared in such a way that the DNA was destroyed (e.g., Specimens examined in this study were dried using ethyl acetate). A single leg was obtained from the following collections: Biodiversity removed from each specimen for sequencing and a Institute of Ontario, Guelph, ON, Canada (BIOUG), 658 base pair fragment of the 50 end of the cyto- Canadian National Collection of Insects, Ottawa, chrome oxidase subunit I (COI) gene (the ‘bar- ON, Canada (CNC), University of Guelph Insect coding’ fragment) was amplified using the primer Collection, Guelph, ON, Canada (DEBU), Royal pair LepF1 (50-ATTCAACCAATCATAAAGAT Ontario Museum, Toronto, ON, Canada (ROME), ATTGG-30)andLepR1(50-TAAACTTCTGGAT University of Alberta, E.H. Strickland Entomologi- GTCCAAAAAATCA-30)(Hebertet al. 2003). cal Museum, Edmonton, AB, Canada (UASM), DNA extraction and sequencing was performed National Museum of Natural History, Washington, at both CNC and at the Canadian Centre for DC, United States of America (USNM). DNA Barcoding following protocols outlined in Specimen preparation follows Skevington (2003). Hajibabaei et al. (2005). The resultant sequences, Photographs were taken with a Leica DM550B as well as images and related data, can be accessed compound microscope (Leica Microsystems Inc., through the Barcode of Life Data Systems (BOLD) Concord, Ontario, Canada) and a Canon EOS 50D (http://www.barcodinglife.org/) in the public project camera equipped with a 100 mm macro lens (Canon ‘Sericomyia – Skevington (SERSK)’ (http://www. Canada Inc., Mississauga, Ontario, Canada). In most boldsystems.org/views/projectmenu.php?&). In cases, Leica Application Suite was used to create a addition, all sequences were deposited in GenBank montage from multiple layers of photographs. (Appendix 1). Measurements were made using a graticule. At least five specimens were used from each species to Data analysis obtain the recorded values. Terminology follows No insertions or deletions occur in the dataset Thompson et al. (2010). All specimens are labelled so alignment was unambiguous. Phenetic and with a unique reference number, typically in the parsimony analyses were performed with PAUP* format J. Skevington Specimen # n, CNC Diptera (Swofford 2001). Neighbour-joining was used to # n,debun, or USNM ENT. The former three have explore species concepts for all 105 ingroup been shortened to follow the format JSSn,CNCDn, taxa. Parsimony and Bayesian methods were and debun, respectively, throughout the text. These used with a reduced dataset (one exemplar per numbers are used in the CNC Diptera specimen species) for creating phylogenetic hypotheses. database (available upon request). Standard two- Character polarity was based on outgroup com- letter postal acronyms are used for distribution data. parison (Nixon and Carpenter 1994). Several Figures are presented in the order that they species of Cheilosia Meigen were defined as appear in the key. Maps include points for all outgroup for all analyses (but not constrained as specimens examined and were produced using such) because they are genetically closest to SimpleMappr (Shorthouse 2010). When addi- Sericomyia from all of the available COI data tional range data are known from the literature it available on BOLD and in GenBank. Parsimony is mentioned in the key along with a citation. analysis with a heuristic search procedure was used with stepwise-addition and 100 random Molecular methods replications. The heuristic search option was All molecular specimens (Appendix 1) were used with tree bisection–reconnection branch dried, pinned, labelled, and labelled with a unique swapping, MULPARS, and random addition of

᭧ 2012 Her Majesty the Queen in Right of Canada 218 Can. Entomol. Vol. 144, 2012 taxa. Multistate characters were treated as non- consensus tree, posterior probabilities for each additive. Evidential support for different clades node, and branch length estimates. was assessed using the nonparametric bootstrap (BS – 1000 replicates) (Felsenstein 1985). For Results and discussion Bayesian analysis, models of evolution were determined based on the Akaike Information Sericomyia Meigen Criterion (AIC) using ModelTest 3.7 (Posada and Crandall 1998). Bayesian analyses were Cinxia Meigen, 1800: 35. Type-species, Musca conducted using MrBayes 3.1.2 (Ronquist and lappona L. by subsequent designation of Huelsenbeck 2003) with a Markov Chain Monte Coquillett (1910: 524). Name suppressed by Carlo (MCMC) method. Four chains (three hot, International Commission on Zoological one cold) were run simultaneously for 400,000 Nomenclature (1963). generations. Trees were sampled every 10 gen- Sericomyia Meigen, 1803: 274. Type-species, erations and each simulation was run twice. The Musca lappona L. by subsequent designation MCMC chains achieved stationarity (standard of Latreille (1810: 443). deviation of split frequencies ,0.01; all para- Arctophila Schiner, 1860: 215. Type-species, meter estimates asymptotic) at 370,000 genera- Syrphus bombiformis Falle´n by subsequent tions. Following the discard of the ﬁrst 10,000 designation of Williston (1887: 158). Syn. nov. samples as burn-in, 30,000 samples were used Condidea Coquillett, 1907: 75. Type-species, for each simulation to generate a majority-rule lata Coquillett by original designation.

Fig. 1. Sericomyia tolli. A, Male habitus, dorsal (CNCD30733); B, female habitus, dorsal (CNCD30743); C, male habitus, lateral (CNCD30733); D, female habitus, lateral (CNCD30743). Scale bars 5 2 mm.

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Conosyrphus Frey, 1915: 17. Type-species, tolli setose patch; metatibia transverse apically, rounded Frey by original designation. Syn. nov. basoventrally. Bulboscrobia Gaunitz, 1937: 91. Type-species, undulans Gaunitz by original designation 5 Wing lappona L. Cell R1 open; stigmatic crossvein absent; cell Tapetomyia Fluke, 1939: 370. Type-species, R4 1 5 with long petiole, longer than humeral meyeri Fluke by original designation. crossvein; vein R415 straight to moderately sinuate; vein M2 absent; vein A1 1 CuA2 short, oblique. Description Head Abdomen Face variable, black to yellow, with or without Oval, slightly longer than broad. black medial vitta, broad, about as broad as long, occupying about one-third head width, concave Classification beneath antenna, usually with large low medial Within the current classification of flower tubercle, pollinose and pilose laterally, shiny and flies, Sericomyia falls into the tribe Sericomyini bare medially; gena narrow, about half as broad of the subfamily Eristalinae. This tribe now as long; anterior tentorial pit short, extending contains only four genera. Besides Sericomyia, along ventral one-third of eye; facial stipes these are: (1) Pyritis Hunter, with a single spe- narrow; frontal prominence low, at dorsal half of cies restricted to the Pacific Northwest of the head; frontal lunule small, narrow; frons broad, Nearctic region characterized by pilose eye and about as long as broad at antenna, with slightly face. (2) Pseudovolucella Shiraki, an Oriental convergent sides dorsally, half as broad at vertex and Far Eastern Palaearctic group of 10 species as at antenna, pollinose and pilose; vertex tri- characterized by a distinctive enlarged flattened angular, as long as broad, pollinose and pilose; head and enlarged metafemur (Reemer and ocellar triangle small; eye bare; narrowly to Hippa 2008). (3) Pararctophila Herve´-Bazin, a broadly holoptic in male; antenna short, at most small group of two to three species ranging from about one-third as long as face; basoflagellomere the Transbailkalia to the Himalayas character- oval to quadrate; arista pilose, about one and a ized by a strongly sinuate vein R4 1 5. half times as long as antenna. The taxonomic concept of Sericomyia is old, dating back to Meigen (1800), and based on two Thorax species (M. lappona and Syrphus mussitans F.). Slightly broader than long, long to short The essential characters were the pilose arista on pilose, without setae; postpronotum pilose; an oval basoflagellomere. Meigen (1800) first proanepisternum pilose, prokatepisternum bare; named the taxon Cinxia but the International proanepimeron pilose; mesonotum without Commission on Zoological Nomenclature (1963) pollinose pattern; katepisternum discontinuously suppressed that name, so his subsequent name, pilose, with broadly separated patches; meta- Sericomyia,isvalid. sternum pilose; anepimeron with dorsomedial This original concept included two kinds of portion bare, posterior portion usually bare, rarely flies: short pilose flies with transverse yellow with some pile anteroventrally; katepimeron bare; fasciate maculae on the abdomen and long pilose postalar pile tuft absent, but with pile along ante- flies without yellow maculae on the abdomen. rodorsal edge of postalar area; metathoracic The latter are bumble bee (Bombus Latreille pleuron bare; metathoracic spiracle about same (Hymenoptera: Apidae)) mimics. Schiner (1860) size in height as length of basoflagellomere, but proposed the name Arctophila for the bumble narrower in width; plumula simple, elongate, short, bee mimics. He also noted that these species are not reaching calypteral margin; scutellum without more robust and their faces are produced much apical sulcus, with ventral pile fringe. more ventrally. Next, Coquillett (1907) proposed Condidea for a Legs new species which has large circular yellow macu- Mesocoxa bare posteriorly; metafemur usually lae on the abdomen and is similar to Sericomyia. narrow, rarely slightly swollen, without basoventral Frey (1915) described Conosyrphus for a new high

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Fig. 2. A, Sericomyia vockerothi female ﬁeld photo, dorsal (Canada: AB: Blackfoot Lake, 53.5369448N, 112.7797228W, 14.v.2006, J.J. Dombroski (specimen not collected)); B, S. vockerothi male habitus, lateral (CNCD35754); C, S. vockerothi surstylus and associated structures, lateral (CNCD35755); D, S. vockerothi surstylus and associated structures, dorsal (CNCD35755); E, S. vockerothi hypandrium and associated structures, lateral (CNCD35755); F, S. vockerothi hypandrium, dorsal (CNCD35755); G, S. harveyi male habitus, dorsal (JSS22100); H, S. harveyi male habitus, lateral (JSS22100). Scale bars for colour illustrations 5 2 mm; for genitalia 5 0.2 mm.

A B

C surstylus

cercus G

epandrium D

distal lobe of surstylus

proximal lobe of surstylus hypandrium postgonite E H

phallus

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Map 1. Range map for Sericomyia slossonae (triangles), Sericomyia tolli (circles), and Sericomyia vockerothi (stars).

Map 1 arctic species, which is sexually dimorphic (male they were bumble bee mimics with robust meta- with an entirely black abdomen, the female with the femora and long faces. Later, however, when Osten standard yellow fasciate maculae on the abdomen) Sacken (1875) described flagrans from North and has a greatly ventrally produced face. Later, America, which has neither a long face nor robust Gaunitz (1937) encountered an aberrant specimen of metafemur, the only distinction was the mimetic Sericomyia lappona and, because of the unusual appearance. Likewise, when Fluke (1939) descri- appearance of furrows on the abdomen, proposed bed Tapetomyia, he was comparing his species Bulboscrobia for it. Finally, Fluke (1939) described with flagrans, not the European species, which Tapetomyia for a bumble bee mimic from Mexico. have longer faces. As for the dichoptic male, there The ‘‘truncated’’ basoflagellomere and dichoptic is a Chinese bumble bee mimic which is also male along with the ‘‘longer and narrower’’ face dichoptic, but has a short face like flagrans. So, the were declared as distinctive. only characteristic that defines and separates Hull (1949), in the last comprehensive generic ‘‘Arctophila’’ from Sericomyia is the mimetic treatment of the family, recognized Sericomyia appearance; hence the declaration that Arctophila and Tapetomyia as distinct with Arctophila, is only a phenetic group. Bulboscrobia, Condidea, and Conosyrphus as As for Conosyrphus, again when the species subgenera of Sericomyia. Curran (1934) declared tolli was described (Frey 1915), it was atypical Condidea as ‘‘hardly tenable’’, and Wirth et al. of the genus Sericomyia due to it long face and (1965) formally synonymized the genus with dimorphic sexes. However, Portschinsky (1881) Sericomyia. Gaunitz (1963) later synonymized his described a species from Caucasia with a long face own genus Bulboscrobia under Sericomyia (as just like tolli that was later placed in Conosyrphus Cinxia). Thompson et al. (1976) in their Neotropical (as a new species by Coe 1966). This species is not flower fly catalogue synonymized Tapetomyia with sexually dimorphic but does have a face even Arctophila. Later Thompson et al. (2000) declared longer than tolli!Also,Sericomyia himalayensis Arctophila to be only a phenetic group. Brunetti (1907, 1908) has a long face, but otherwise When these new off-shots from Sericomyia were is typical of Sericomyia. proposed, they seemed reasonable. For Schiner In summary, the characters originally proposed (1860), the two European Arctophila species were to define the taxonomic concepts of Arctophila, quite different from the other Sericomyia species as Conosyrphus,andTapetomyia vary independently

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Fig. 3. A, Sericomyia flagrans male habitus, dorsal (CNCD1976); B, S. flagrans male habitus, dorsal (to show variation in abdominal colouration) (USNMENT00035368); C, S. flagrans male habitus, lateral (CNCD1976); D, Sericomyia meyeri female habitus, dorsal (USNMENT00022495); E, S. meyeri female habitus, lateral (USNMENT00022495); F, Sericomyia fairmanorum female habitus, dorsal (USNMENT00037868); G, S. fairmanorum female habitus, lateral (USNMENT00037868). Scale bars 5 2 mm.

A B

G F

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Map 2. Range map for Sericomyia fairmanorum (circles), Sericomyia harveyi (triangles), and Sericomyia meyeri (stars).

Map 2

within the clade that Meigen originally recognized Conosyrphus as new synonyms of it. Molecular as Sericomyia. Hence, we here formally return to support for this is discussed in the phylogeny Meigen’s concept and ofﬁcially treat Arctophila and section below.

Key to New World Sericomyia

1 Face yellow, medially and greatly produced ventrally, projecting more than half eye length below eye (Fig. 1A–1D) ...... tolli (Frey) Arctic (AK, NT, NU, YT, Eurasia) (Map 1); Frey (1915: 18, Conosyrphus). – Face usually projecting less than half eye length below eye (cf. Fig. 10F), if longer then with black medial vitta (Fig. 5D–5E) ...... 2 2 Abdomen with yellow markings at leastontergum2;bodywithshorthairs;fliesnotmimickingbumblebees....7 – Abdomen entirely black, with some or all of terga 2–4 red- or yellow-haired; body with long pile; flies mimicking bumble bees ...... 3 3 Scutellum yellow pilose (Fig. 3A, 3D, 3F) ...... 5 – Scutellum black pilose (Fig. 2A, 2G) ...... 4 4 Faceblack;postalarcallusyellowpilose;scutumentirelyyellowpilose(Fig.2A–2F) ...... vockerothi sp. nov. North central and northeastern North America (AB, MN, NT, QC, YT) (Map 1); here described. – Face brownish-yellow, with brown medial vitta; postalar callus black pilose; scutum broadly black pilose anterior to scutellum (Figs. 2G–2H, 5) ...... harveyi (Osburn) Pacific coast (BC to OR, CA) (Map 2); Osburn (1908: 9, Arctophila).

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5 Face yellow medially; antenna orange; anepimeron pilose posteriorly (Figs. 3A–3C, 6) . . . ﬂagrans (Osten Sacken) Western North America (AB, AK, BC, CA, CO, ID, MT, NM, NV, OR, UT, WA, WY, YT) (Map 3); Osten Sacken (1875: 54, Arctophila) – Face and antenna dark, reddish-brown to black (Fig. 3E, 3G); anepimeron bare posteriorly ...... 6 6 Abdomen extensively yellow to reddish pilose on 3rd and 4th terga; wing bare basomedially; face produced ventrally; male dichoptic; male metatibia without apical spur (Figs. 3D–3E, 5) ...... meyeri (Fluke) Central Mexico (Map 2); Fluke (1939: 370, Tapetomyia). – Abdomen black pilose on 3rd and 4th terga; wing microtrichose; face not produced ventrally; male holoptic; male metatibia with apical spur (Figs. 3F–3G, 5) ...... fairmanorum (Fairman) Costa Rica (Map 2); Fairman (Thompson et al. 2000: 40, Arctophila). 7 Face yellow with medial black vitta (Fig. 5D) ...... 9 – Face completely yellow without medial black vitta (Fig. 4C) ...... 8 8 Abdomen with only one pair of maculae; maculae fasciate, narrow, on 2nd tergum; scutellum yellow pilose; all basitarsi orange to reddish-brown (Fig. 4A–4B) ...... carolinensis (Metcalf) Southeastern United States of America (MD to MS) (Map 3); Metcalf (1917: 209, Cinxia). – Abdomen with three pairs of maculae; maculae rotund on 2nd tergum, arcuate on 3rd and 4th terga; scutellum extensively black pilose; all tarsi black (Fig. 4C–4F) ...... lata (Coquillett) BC to NB south to NE, WV (Map 4; Pape and Thompson 2010); Coquillett (1907: 75, Condidea). 9 Abdomen with two pairs of fasciate maculae; maculae absent on 4th tergum (male only) (Fig. 5A–5B) ...... bifasciata Williston MI and ON to NL south to PA (Map 3; Pape and Thompson 2010); Williston (1887: 154). – Abdomen usually with three or more pairs of fasciate maculae (cf. Fig. 6A, 6E); if only two, then none on 2nd tergum and a pair present on 4th tergum (Fig. 6F) ...... 10 10 Abdomen with maculae on 2nd tergum twice as large as rest of maculae; legs all black except femoral-tibial joints reddish;scutellumwithdiscwithappressedblacksetae(Fig.6A–6C) ...... transversa (Osburn) ND and MB east to NL (Map 5; Pape and Thompson 2010); Osburn (1926: 51, Condidea). – Abdomen with maculae on 2nd tergum subequal to those on other terga; legs with at least pro- and mesotibiae reddish-brown to yellow; scutellum with erect normal pile ...... 11 11 Maculae on 2nd tergum small, often absent; all maculae small, oblique, dashlike, widest medially; scutellum usually reddish; male metacoxa with spur; anepimeron black pilose; postalar callus yellow pilose, rarely with a few black pili intermixed;abdomenusuallyyellowpilose,with4thtergumyellow(Fig.6D–6G) ...... militaris Walker AK to NL south to NM (Map 6; Pape and Thompson 2010); Walker (1849: 595). – Maculae on 2nd tergum very distinct, elongate and oblique, those of following terga wider laterally ...... 12 12 Face produced strongly ventrally by a distance more than three-fourth height of eye; metafemur black except narrowly yellowonapex;fronsallblackpilose(femalesonly)(Fig.5C–5F) ...... bifasciata Williston See above. – Face not strongly produced ventrally, by a distance less than half of eye height (cf. Fig. 10F); metafemur of female all yellow; frons with ventral one-third yellow pilose (males and females) ...... 13 13 Abdominal fasciae with concave margins, narrowest submedially, broadly separated medially (cf. Fig. 10A) ...... 16 – Abdominal fasciae with straight margins narrowest medially, continuous or only narrowly separated medially (cf. Fig. 8A) ...... 14 14 All femora black on basal two-thirds or more; hypandrium very similar to that of S. woodi with two to three small apical protuberances (Fig. 7A–7B) ...... slossonae Curran ON, NY and NH (Map 1); Curran (1934: 4). – At least metafemur reddish-orange to reddish-brown (Figs. 8B, 9B); hypandrium as illustrated in Figs. 8D, 8F, 9D–9E ...... 15 15 Proepimeral and procoxal setae entirely yellow (Fig. 8C); hypandrium knife-like with large dorsal proximal protuberance and two smaller distal protuberances (Fig. 8D, 8F) ...... nigra Portschinsky Arctic, AK to NL, Eurasia (Map 7); Portschinsky (1873: 291). – Proepimeral and procoxal setae usually entirely black, occasionally yellow with some black bristles (Fig. 9C); hypandrium with two to three small distal protuberances (Fig. 9D, 9F) ...... woodi Nielsen and Vockeroth Arctic, AK, NT, YT (Map 5); Nielsen and Vockeroth (2000: 137).

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16 Supra-alar pile yellow (Figs. 11A–11C, 12A–12B, 12F–12G) ...... 18 – Supra-alar pile extensively black (Fig. 10A, 10E) ...... 17 17 Abdominal bands oblique (Fig. 10E, 10G); hypandrium narrow with a few scattered lateral spines (Fig. 10H) ...... chrysotoxoides Macquart AB to NL south to TN and SC (Map 8); Macquart (1842: 79). – Abdominal bands transverse (Fig. 10A, 10C); hypandrium wide, ﬂared out medially, with a row of rather uniformly spaced lateral spines (Fig. 10D) ...... chalcopyga Loew AK to SK south to CA and UT (Map 9); Loew (1863: 12). 18 Tergum 3 with yellow pile; scutellum black, same colour or slightly paler than thoracic dorsum; tibiae black; markings on terga 2–4 almost identical, constricted medially; hypandrium tip ﬂared with three hook-like teeth (Fig. 11A–11G) ...... sexfasciata Walker AK to NL and ME (Map 10; Pape and Thompson 2010); Walker (1849: 596). – Tergum 3 with black pile in male, yellow or mixed yellow and black pile in female; scutellum reddish brown, contrasting with black thoracic dorsum; tibiae usually brownish red, rarely black; band on tergum 4 narrower than those on terga 2–3, only slightly constricted medially; hypandrium not as above ...... 19 19 Male hypandrium with saw-like edges; females indistinguishable from S. jakutica (Fig. 12A–12E) ...... arctica Schirmer AK to NU, Eurasia (Map 4); Schirmer (1913: 221). – Male hypandrium with smooth edges, and sharp divergent tips; females indistinguishable from S. arctica (Fig. 12F–12J) ...... jakutica (Stackelberg) AK to MB, Russia (Map 8); Stackelberg (1927: 21, Cinxia).

Sericomyia vockerothi Skevington pilose (Fig. 2A); proepisternum with sparse sp. nov. yellow pile; proepimeron with black pile; anterior anepisternum bare; posterior anepisternum and (Fig. 2A–2F, Map 1) anterior anepimeron with long, dense yellow pile; Diagnosis katepisternum with black pile ventrally, bare and Sericomyia vockerothi is superﬁcially similar shining medially and with yellow pile dorsally; to the common and widespread Sericomyia ﬂa- meron, posterior anepimeron and katetergum bare; grans (Fig. 3) but differs in having an entirely metasternum black pilose; plumula black with black pilose scutellum (similar to the uncommon short black pile; calypter black with black fringe. S. harveyi). The face is entirely black, the postalar callus is yellow pilose, and the scutum is Legs entirely yellow pilose. Femora and tibiae shape cylindrical with no protuberances, black, black pilose; tarsomeres Description 1–3 reddish brown with dorsal setae black, ventral Body length: 12.6 to 14.0 mm; wing length: pile reddish brown; tarsomeres 4–5 blackish with 9.5 to 10.5 mm. black setae. Head Wing Frons and face entirely dark brown to black (Fig. Wing completely densely microtrichose; dark 2B); face bare and shiny except for golden vestiture brown at base and on pterostigma, light brown in beneath antennae and a mixture of pale yellow and costal cell, middle of r1, basal half of r2 1 3, distal black pile along the lateral fringe; frons and ocellar third of br and around r-m and bm-cu; remainder triangle covered with long, pale yellow pile; occiput of wing hyaline. silvery-white pollinose, yellow pilose dorsally; Abdomen antenna light brown, black pilose; arista pilose; Black; terga 1–4 shiny, black pilose; tergum 5 male holoptic with area of contact somewhat shorter mostly to entirely black, sometimes yellow along than length of ocellar triangle, female dichoptic. posterior margin, shiny to weakly grey pollinose, Thorax yellow pilose; sterna 2–4 shiny black with black Black; postpronotum, mesonotum and postalar pile; sternum 5 black to brown with yellow pile; callus entirely yellow pilose; scutellum black terminalia black to brown and yellow pilose.

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Fig. 4. A, Sericomyia carolinensis male habitus, lateral (CNCD29611); B, S. carolinensis male habitus, dorsal (CNCD29611); C, S. lata female, frontal of head (CNCD601); D, S. lata male habitus, lateral (CNCD2222); E, Sericomyia lata male habitus, dorsal (CNCD2222); F, S. lata female ﬁeld photo (S.A. Marshall). Scale bars 5 2mm.

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Map 3. Range map for Sericomyia bifasciata (circles), Sericomyia carolinensis (stars), and Sericomyia ﬂagrans (triangles).

Map 3

Map 4. Range map for Sericomyia lata (circles) and Sericomyia arctica (triangles).

Map 4

Genitalia with large holes dorsally giving it a skull-like Surstylus short, with pointed, upturned term- appearance from above (Fig. 2E); large tooth at inal finger deflected left (Fig. 2C–2D); proximal distal end of hypandrium (Fig. 2E–2F); ejacu- lobe of surstylus bare on outer surface, heavily latory apodeme bulb-shaped; phallapodeme setose on medial surface; distal lobe of surstylus broad, axe-shaped proximally; postgonite with covered with setae on most surfaces, bare only four to five dorsal teeth on outer lobe (Fig. 2E), on finger-like tips; cercus somewhat rectangular inner lobe smooth; phallus smooth, hook-shaped, with longer setae than surstylus; hypandrium with hook projecting dorsally (Fig. 2E).

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Fig. 5. Sericomyia bifasciata. A, Male habitus, dorsal (CNCD29545); B, male habitus, lateral (CNCD2711); C, female habitus, lateral (CNCD2712); D, female head, frontal (CNCD600); E, female head, lateral (CNCD2712); F, female habitus, dorsal (CNCD31287). Scale bars 5 2 mm.

A B

D E

Material examined Paratypes: CANADA: ALBERTA: Doussal, Holotype # labelled: CANADA, YUKON 55.61698N, 116.83158W, 632 m, 1.vi.1961, A.R. TERRITORY: Sheldon Lake, 62.66778N, Brooks, CNCD35754 (1 #, CNC). NORTH- 131.10008W, 914 m, 21.vii.1960, J.E.H. Martin, WEST TERRITORIES: Martin River, 10 miles CNCD35752 (CNC). NW of Fort Simpson, 61.89348N, 121.61328W,

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Fig. 6. A, Sericomyia transversa female habitus, dorsal (CNCD509); B, S. transversa female habitus, lateral (CNCD510); C, S. transversa male scutellum, dorsal (CNCD509); D, Sericomyi militaris male scutellum, dorsal (CNCD111); E, S. militaris female ﬁeld photo (S.A. Marshall); F, S. militaris male habitus, dorsal (CNCD111); G, S. militaris male habitus, lateral (CNCD111). Scale bars 5 2 mm.

A B

C F

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Map 5. Range map for Sericomyia transversa (triangles) and Sericomyia woodi (circles).

Map 5

Map 6. Range map for Sericomyia militaris.

Map 6

15.vi.1972, B.V. Peterson, CNCD35755 (1 #, J. Moisan-De Serres, JSS19210 (1 #,USNM). CNC). QUEBEC: Albanel, 48.82948N, 72.35128W, YUKON TERRITORY: Swim Lakes, 62.21678N, 13.vi.2009, blueberry field, J. Moisan-De Serres, 133.00008W, 975 m, 23.vi.1960, F.W. Rockburne, JSS19703 (1 ~, CNC); Dolbeau, 48.86108N, CNCD35753 (1 ~, USNM); La Force Lake, 72.23458W, 17.vi.2009, blueberry field, J. Moisan- 62.68338N, 132.33338W, 1006 m, 5.vii.1960, F.W. De Serres, JSS19236 (1 ~, CNC); St.-Euge`ne, Rockburne, CNCD35751 (1 ~,CNC).UNITED 49.00038N, 72.34768W, 16.vi.2009, blueberry field, STATES: MINNESOTA: West Cook, Saganaga

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Fig. 7. Sericomyia slossonae male habitus (CNCD508). A, Dorsal; B, lateral. Scale bars 5 2 mm.

A B

Lake, Conners Island, 48.240168N, 90.849558W, Nearctic Region: USA: AK: King Salmon, 3.vi.1964, W.E. LaBerge & D.W. Ribble, USN- Naknek River, 58.68338N, 156.658W, 1.viii.1952, MENT00036891 (1 #,USNM). 13.vii.1952, 23.vii.1952, 31.vii.1952, J.B. Hartley, CNCD30105, 30108, 30670-6, 30683 (10 ##, Etymology CNC); Oumalik, 69.83338N, 156.008W, 15.vii.1949, Named for John Richard Vockeroth, in recogni- N.A. Weber, CNCD30669 (CNC). Canada: MB: tion of his life-long achievements in syrphidology Twin Lakes, 58.632088N, 96.788518W, 15.vii.2007, research. Dick was also in the process of revising P. Hebert, CNCD30696 (CNC); 23 km E Churchill, Sericomyia, so here we build on his knowledge of Ramsay Creek, 58.730108N, 93.771008W, the group. 3.vii.2007, P. Hebert, 07PROBE10093 (BIOG), CNCD30690 (2 ##, CNC); Churchill, 58.76888N, Distribution 94.17168W, 10, 12.vii.1948, 25, 28.vi.1952, Widespread but rarely collected across northern 24.vii.1952, J.G. Chillcott, L.A. Miller, Canada (AB, NT, QC, YT – Map 1). CNCD30104, 30688, 30692-4 (2 ##,3~~, CNC). NT: Coppermine, 62.562738N, 115.092768W, Remarks 3.viii.1951, S.D. Hicks, CNCD30689 (1 #,CNC); This is the ‘‘nov. spec.’’ from north central Norman Wells, 65.266678N, 126.816678W, North America given in the Arctophila key by 5.viii.1969, G.E. Shewell, CNCD2230 (1 ~,CNC); Thompson et al. (2000). Reindeer Depot, Mackenzie Delta, 68.668978N, 134.071578W, 2,4.vii.1948, J.R. Vockeroth, W.J. Revisionary notes Brown, CNCD30106-7, 30678-81, 30686-7, 30695 (8 ##,1~, CNC), Kidluit Bay, 1. Mallota powelli Nayar and Cole (1968: Richard Island, 69.326348N, 134.437248W, 28, 288) is merely a mis-identified species and is a 30.vii.1948, J.R. Vockeroth, CNCD30677. junior synonym of S. flagrans (Osten Sacken) 30691 (1 #,1~, CNC), 21 miles E Tuktoyaktuk, (syn. nov.). 69.426848N, 132.172478W, 8–12, 17–21.vii.1971, 2. The colour characters given by van Stack- D.M. Wood, CNCD2212, 30682, 30684-5 (4 ##, elberg (1927) and van Veen (2004) to separate CNC), Tuktoyaktuk, 69.433858N, 133.014738W, S. arctica from Sericomyia jakutica do not work. 15–18.vii.2010, H. Goulet, C. Boudreault, Nielsen (1997) also recognized this. JSS24604–6, 24959 (1 #,3~~,CNC),Sachs 3. Sericomyia jakutica is here recognized for Harbour, 71.986538N, 125.2438W, 19–22.vii.2010, the first time from the Nearctic Region, we have H. Goulet, C. Boudreault, JSS24953-4 (2 ~~, examined the following specimens from the CNC). YT: Summit Lake, N. Richardson Mountains,

᭧ 2012 Her Majesty the Queen in Right of Canada 232 Can. Entomol. Vol. 144, 2012

Fig. 8. Sericomyia nigra. A, Male habitus, dorsal (CNCD30637); B, female habitus, lateral (CNCD9764); C, male foreleg, lateral (CNCD30583); D, hypandrium (phallus removed; CNCD30591); E, surstyli and associated structures (CNCD30590); F, lateral of hypandrium and associated structures (CNCD30951). Scale bars for colour illustrations 5 2 mm; for genitalia 5 0.2 mm.

A B

E F

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Map 7. Range map for Sericomyia nigra.

Map 7

Map 8. Range map for Sericomyia chrysotoxoides (circles) and Sericomyia jakutica (triangles).

Map 8

68.164038N, 136.988258W, 3.vii.1987, S.A. and provide a database to assist with future identi- Marshall, debu1047493 (1 #, DEBU). ﬁcations (particularly of non-adults). Most (97 specimens of 14 species) sampled were Nearctic. All Species concepts morphological species except two were supported by the molecular data and all showed very little DNA barcode data (50 end of the COI) were intraspeciﬁc variation (Fig. 13). Uncorrected pair- collected for 101 Sericomyia specimens of 16 spe- wise divergences varied from 0% to 0.72% within cies to test current morphological species concepts species and 0% to 9.85% between species. Average

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Fig. 9. Sericomyia woodi. A, Male habitus, dorsal (CNCD30858); B, female habitus, lateral (CNCD30858); C, male foreleg, lateral (CNCD30856); D, hypandrium and associated structures (CNCD2232); E, surstyli and associated structures (CNCD2232); F, lateral of hypandrium and associated structures (CNCD2232). Scale bars for colour illustrations 5 2 mm; for genitalia 5 0.2 mm.

A B

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Fig. 10. A, Sericomyia chalcopyga male habitus, dorsal (CNCD1843); B, S. chalcopyga male habitus, lateral (CNCD1843); C, S. chalcopyga male ﬁeld photo (John Davis); D, S. chalcopyga hypandrium (CNCD33896); E, Sericomyia chrysotoxoides male habitus, dorsal (CNCD29977); F, S. chrysotoxoides male head, lateral (CNCD505); G, S. chrysotoxoides female ﬁeld photo (Virginia, Dave Cheung); H, S. chrysotoxoides hypandrium (CNCD29795). Scale bars for colour illustrations 5 2 mm; for genitalia 5 0.2 mm.

A B

black supra-alar pile

D H

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Map 9. Range map for Sericomyia chalcopyga.

Map 9

Map 10. Range map for Sericomyia sexfasciata.

Map 10

pairwise divergence between species was 5.36% and the result of a sequencing error. These species are most species were more than 2% different. Excep- dramatically different morphologically and have tions were Sericomyia slossonae and Sericomyia been treated as belonging to separate subgenera woodi (0.73–1.21% different) and S. jakutica and historically. S. tolli has pronounced sexual Sericomyia tolli. The latter two species were found dimorphism (sexes are similar in S. jakutica), dif- to be genetically identical. Several specimens were ferent colour patterns from S. jakutica,andagreatly sampled for each species to ensure that this was not projecting face (face of S. jakutica is half the length).

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Fig. 11. Sericomyia sexfasciata. A, Female habitus, dorsal (CNCD507); B, female habitus, lateral (CNCD507); C, male ﬁeld photo (Churchill, Manitoba, J.H. Skevington); D, lateral of hypandrium and associated structures (CNCD30648); E, hypandrium, ventral (CNCD30648); F, surstyli and associated structures (CNCD30648); G, hypandrium, dorsal (CNCD30648). Scale bars for colour illustrations 5 2 mm; for genitalia 5 0.2 mm.

A B

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Fig. 12. A, Sericomyia arctica male habitus, dorsal (CNCD29517); B, S. arctica male habitus, lateral (CNCD29517); C, S. arctica surstyli and associated structures (CNCD29518); D, S. arctica lateral of hypandrium and associated structures (CNCD29518); E, S. arctica hypandrium and associated structures (CNCD29518); F, Sericomyia jakutica male habitus, dorsal (genitalia twisted out for examination – CNCD30683); G, S. jakutica male habitus, lateral (CNCD30683); H, S. jakutica surstyli and associated structures (CNCD30107); I, S. jakutica lateral of hypandrium and associated structures (CNCD30107); J, S. jakutica hypandrium and associated structures (CNCD30669). Scale bars for colour illustrations 5 2 mm; for genitalia 5 0.2 mm.

A B

saw-like teeth on hypandrium D

F G

hypandrium smooth with sharp, divergent tips

H I

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Fig. 13. Neighbour-joining phenogram of Sericomyia species produced from analysis of COI data.

Cheilosia sp. 09BBEDI-0756 CNCD-2214 S. bifasciata CNCD-2220 C. sp.| JSS21087 S. sachalinica CNCD14810 C. prima S. bifasciata CNCD-2215 C. sp. nov. 1 CNCD9545 S. militaris CNCD9376 C. sp. 09BBEDI-0751 S. bifasciata CNCD-2213 C. sp. nov. 2 CNCD9158 S. militaris CNCD-4997 S. bifasciata 09BBEDI-0250 C. nigrofasciata CNCD11268

C. sp. JSS21085 CNCD-2224 S. bifasciata 09BBEDI-0458 S. militaris C. tristis 07WNP-10967 08TTML-0547 To 1 S. chrysotoxoides S. militaris CNCD-2225

S. chrysotoxoides CNCD-4996 S. militaris 08|08BBDIP-1367 S. chrysotoxoides CNCD-4995

S. militaris 09BBEDI-0098 CNCD-2217 S. chalcopyga CNCD-1902 S. chrysotoxoides S. chalcopyga CNCD-2241 S. chrysotoxoides 09BBEDI-0248 S. militaris 09BBEDI-0153 S. chalcopyga CNCD-929 S. flagrans 08|08BBDIP-1022 S. chrysotoxoides CNCD-2218 S. militaris CNCD-4998 S. flagrans CNCD-1567 S. flagrans USNMENT35369 S. chrysotoxoides 09BBEDI-0245 S. militaris 07PROBE-04537 S. flagrans CNCD1564 09BBEDI-0249 S. flagrans CNCD1976 S. chrysotoxoides S. flagrans USNMENT35368 S. militaris 07PROBE-04538 S. chrysotoxoides 09BBEDI-0255 S. lata 08BBDIP-0373 CNCD-2222 08TTML-0548 S. lata S. chrysotoxoides 09BBEDI-0260 S. militaris S. lata CNCD-18 09BBEDI-0196 09BBEDI-0247 S. lata S. chrysotoxoides S. militaris 08TTML-2434 S. lata CNCD-2223 S. chrysotoxoides 09BBEDI-0252 S. jakutica 07PROBE-10093 S. militaris 08TTML-2451 S. jakutica JSS24953 S. chrysotoxoides 09BBEDI-0258 S. jakutica JSS24954 S. militaris 08BBDIP-1656 S. tolli CNCD-2234 S. chrysotoxoides 09BBEDI-0040 S. jakutica CNCD-2230 08BBDIP-1365 09BBEDI-0259 S. militaris S. jakutica CNCD-2212 S. chrysotoxoides S. jakutica JSS24959 S. chrysotoxoides 09BBEDI-0254 S. militaris 09BBEDI-0149 S. jakutica debu1047493 S. tolli CNCD30742 1 S. chrysotoxoides 09BBEDI-0246 09BBEDI-0150 S. tolli CNCD30712 S. militaris S. nigra CNCD-2228 S. chrysotoxoides CNCD-65 2 S. nigra CNCD-2227 S. militaris 09BBEDI-0151 S. chrysotoxoides 08TTML-2408 S. transversa CNCD-2236

S. transversa CNCD-2237 S. militaris 09BBEDI-0152 S. chrysotoxoides 08TTML-2457 S. woodi CNCD-2240 S. woodi CNCD-2232 S. chrysotoxoides 08TTML-2449 S. militaris 09BBEDI-0154 S. woodi debu1047488 S. slossonae debu170285 S. chrysotoxoides 08TTML-2441 S. militaris 09BBEDI-0155 S. slossonae 30699 S. chrysotoxoides 08TTML-2439 S. sexfasciata CNCD9573 S. militaris 09BBEDI-0156 S. sexfasciata CHU06-SYR-123 S. chrysotoxoides 08TTML-0557 S. sexfasciata 07PROBE-10084 S. militaris 08BBDIP-1657 S. sexfasciata CNCD9475 S. chrysotoxoides 08TTML-0555 S. sexfasciata CNCD9518 09BBEDI-0256 S. sexfasciata CNCD9609 S. chrysotoxoides S. lappona CNCD-2111 S. sexfasciata CNCD35781 S. chrysotoxoides 09BBEDI-0257 S. sexfasciata CNCD-2235 S. lappona CNCD-2229 S. vockerothi JSS19210 JSS19703 S. vockerothi To 2 S. lappona CNCD-2221

Some species that show substantial variation less than 0.69% for COI and no differences were in colour patterns among individuals were found found within the genitalia. Similarly, six speci- to be genetically cohesive. For example, 22 mens of S. ﬂagrans were sampled and included specimens of Sericomyia militaris were sampled dark specimens, predominantly yellow speci- from across the species range, including a vari- mens and specimens with considerable orange ety of phenotypes. These specimens varied by pile. They varied by less than 0.61% for COI and

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Fig. 14. Strict consensus cladogram of Sericomyia species produced from parsimony analysis of COI data. Bremer supports followed by bootstraps indicate clade support.

Cheilosia sp. nov. CNCD9545 3, 73 C. nigrofasciata CNCD11268 1 6, 85 C. prima CNCD14810

C. tristis 07WNP-10967

Sericomyia bifasciata 09BBEDI-0250

S. chalcopyga CNCD929 2, 82 1

S. sachalinica CNCD2220

1 S. chrysotoxoides CNCD2217 1, 79 S. lappona CNCD2111 1

S. militaris 07PROBE-04537

S. flagrans CNCD1564

5, 85 S. lata CNCD2222 1

S. transversa CNCD2237

S. nigra CNCD2228

S. sexfasciata CNCD35781

4, 81 S. woodi CNCD2240 3, 96

S. slossonae debu170285

S. vockerothi JSS19210

S. tolli CNCD30742 5, 96

S. jakutica debu1047493 also showed no variation in genitalic characters. Phylogeny Specimens showing pronounced differences in colour pattern were also collected at the same Parsimony analysis of COI data using single location on the same dates, further supporting exemplars for each species available produced their treatment as a single species. 12 equally parsimonious cladograms (Fig. 14). The molecular data from this study are avail- The strict consensus tree produced is very able from GenBank and can be analysed and similar to the topology of the Bayesian con- explored in the public project on BOLD sensus tree (Fig. 15). The only difference is that ‘‘Sericomyia – Skevington (SERSK)’’ (http://www. the Bayesian tree shows some additional reso- boldsystems.org/views/projectmenu.php?&). lution. Two major lineages of Sericomyia are

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Fig. 15. 50% majority rule consensus cladogram of Sericomyia species produced from Bayesian analysis of COI data. Clade supports shown are posterior probabilities.

Cheilosia tristis 07WNP-10967 0.98 C. nigrofasciata CNCD11268 0.66 C.sp. nov. CNCD9545

C. prima CNCD14810

Sericomyia bifasciata 09BBEDI-0250 0.80

S. chalcopyga CNCD929

CNCD2217 1.00 S. chrysotoxoides

S. sachalinica CNCD2220 0.54

0.97 0.82 S. militaris 07PROBE04537

S. lappona CNCD2111

S. flagrans CNCD1564

S. lata CNCD2222 0.95 0.68 S. transversa CNCD2237

S. nigra CNCD2228 0.73 S. woodi CNCD2240 1.00 0.89 S. slossonae debu170285

S. sexfasciata CNCD35781

S. vockerothi JSS19210

S. tolli CNCD30742 1.00 S. jakutica debu1047493 0.20 0.10 0.0

supported but they do not reflect the current are in this same lineage and are closely related to subgeneric concepts. S. militaris. Sequence data for more Old World The close relationship between S. (Conosyrphus) Sericomyia are needed to find out if they are all tolli and S. (Sericomyia) jakutica supports our belief in this one clade. that Conosyrphus is a synonym of Sericomyia.The Within the other large lineage Sericomyia lata two Arctophila species included in the analysis and Sericomyia transversa are weakly supported (S. flagrans and S. vockerothi) are hypothesized to as sister taxa. These are both rather autapo- be in entirely different lineages, supporting our morphic Sericomyia that would be difficult to synonymy above as well as the contention of place morphologically so it will be interesting if Thompson et al. (2000) that Arctophila is not future research using more genes supports this monophyletic. In the future it would be inter- relationship. Within this lineage Sericomyia esting to add some Old World Arctophila species nigra, S. slossonae, and S. woodi are supported to the analysis to see how many times bumble as closely related. Their relationship was bee mimicry arose within the group. anticipated based on morphology as they all A few other patterns appear from this analysis. have continuous bands on abdominal terga 2–4 The two Old World species that we have data for and have very similar genitalia. S. slossonae and

᭧ 2012 Her Majesty the Queen in Right of Canada 242 Can. Entomol. Vol. 144, 2012

S. woodi have virtually identical genitalia, are Coquillett, D.W. 1907. New genera and species of very similar genetically and are best separated Diptera. The Canadian Entomologist, 39: 75–76. based only on leg colour. Their disjunct dis- Coquillett, D.W. 1910. The type-species of the North American genera of Diptera. Proceedings of United tributions support their continued treatment as States National Museum, 37: 499–647. separate species but more study is needed. Curran, C.H. 1934. Notes on the Syrphidae in the Adding additional genes and a morphological Slosson collection of Diptera. American Museum dataset is clearly the next step to more fully Novitates, 724: 1–7. understand the relationships and evolution of Felsenstein, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39: Sericomyia species. 783–791. Fluke, C.L., Jr. 1939. New Syrphidae (Diptera) from Acknowledgements Central and North America. Annals of the Entomological Society of America, 32: 365–375. Thanks to Barry Flahey for producing the Frey, R. 1915. Diptera Brachycera aus den arktischen excellent genitalia drawings and to Scott Kelso Kunstengegenden Sibiriens. Zapiski Imperatorskoi for developing protocols for sequencing historical Akademii Nauk, ser. 8, 19(10): 1–35. pinned specimens and for collecting the COI Gaunitz, S. 1937. Eine neue Syrphidengattung. Entomologisk Tidskrift, 58: 91. barcoding data for many of the species of Gaunitz, S. 1963. Notes on Syrphidae. Entomologisk Sericomyia. This study was supported by fund- Tidskrift, 84: 15–17. ing to Agriculture and Agri-Food Canada, the Hajibabaei, M., deWaard, J.R., Ivanova, N.V., International Barcode of Life Project (iBOL) Ratnasingham, S., Dooh, R.T., Kirk, S.L., et al. through the Canadian Centre for DNA Barcoding, 2005. Critical factors for assembling a high volume of DNA barcodes. Philosophical Transactions of from the Ontario Genomics Institute, Genome the Royal Society B: Biological Sciences, 360: Canada, the Ontario Ministry of Research and 1959–1967. Innovation, and the Natural Sciences and Engi- Hebert, P.D.N., Cywinska, A., Ball, S.L., and neering Research Council of Canada. NSERC deWaard, J.R. 2003. Biological identifications Canpolin provided the framework for this and other through DNA barcodes. Proceedings of the Royal Society of London (B), 270: 313–322. current research on Syrphidae (this is Canpolin Hull, F.M. 1949. The morphology and inter- publication no. 22). The following curators and relationship of the genera of syrphid flies, recent curatorial assistants provided specimens and data and fossil. Transactions of the Zoological Society, used in this study: V. Levesque-Beaudin (BIOUG), 26: 257–408. S.A. Marshall (DEBU), B. Hubley (ROME), International Commission on Zoological Nomenclature. 1963. Opinion 678. The suppression under the and F. Sperling and D. Shpeley (UASM). Steve plenary powers of the pamphlet published by Marshall, Jason Dombroski, Dave Cheung, and Meigen, 1800. Bulletin of Zoological Nomenclature, John Davis provided some of the field photographs. 20: 339–342. We first discovered John’s excellent photo of Latreille, P.A. 1810. Conside´rations geńe´rales sur Sericomyia chalcopyga on BugGuide (http:// l’ordre naturel des animaux composant les classes des Crustace´s, des Arachnides, et des Insectes; avec bugguide.net). Martin Hauser, Ximo Menguel, un tableau me´thodique de leurs genres, dispose´s en Bradley Sinclair, and Robb Bennett reviewed the familles. Schoell, Paris. manuscript and helped to improve the final pro- Loew, H. 1863. Diptera Americae septentrionalis duct. Skevington was responsible for the DNA indigena. Centuria tertia. Berliner Entomologischer work and took the lead in the preparation of the Zeitschrift, 7: 1–55. Macquart, P.J.M. 1842. Dipteres exotiques nouveaux manuscript. Thompson contributed to the mor- ou peu connus. Tome deuxieme. – 2e partie. phological work and the identification key. Me´moires de la Socie´te´ Royale Sciences, de l’Agriculture et des Arts, Lille, 1841(1): 65–200. References Meigen, J.W. 1800. Nouvelle classification des mouches a` deux ailes (Diptera L.) d’apre`s un plan Brunetti, E. 1907. Notes on Oriental Syrphidae. Records tout nouveau. Perronneau, Paris. of the Indian Museum, 1: 379–380, pls. 11–13. Meigen, J.W. 1803. Versuch einer neuen Gattungs- Brunetti, E. 1908. Notes on Oriental Syrphidae with Eintheilung der europaïschen zweiflu¨gligen Insekten. descriptions of new species. Part I. Records of the Magazin fu¨r Insektenkunde, 2: 259–281. Indian Museum, 8: 49–96. Metcalf, C.L. 1917. Two new Syrphidae (Diptera) Coe, R.L. 1966. A new Sericomyia from Turkey (Diptera: from eastern North America. Entomological News, Syrphidae). Entomologisk Tidskrift, 87: 118–120. 28: 209–212, 1 pl.

᭧ 2012 Her Majesty the Queen in Right of Canada Skevington and Thompson 243

Nayar, J.L., and Cole, F.R. 1968. A new species of Schirmer, K. 1913. Zwei neue Dipteren aus dem Mallota from North America (Diptera: Syrphidae). Norden und Suden Europas. Wiener Entomologische Pan-Pacific Entomologist, 44: 287–289. Zeitung, 32: 221–222. Nielsen, T.R. 1997. The hoverfly genera Anasimyia Shorthouse, D.P. 2010. SimpleMappr, a web-enabled Schiner, Helophilus Meigen, Parhelophilus Girschner tool to produce publication-quality point maps and Sericomyia Meigen in Norway (Diptera, [online]. Available from http://www.simplemappr. Syrphidae). Fauna Norvegica, Serie B, 44(2): 107–122. net [accessed 2 March 2011]. Nielsen, T.R., and Vockeroth, J.R. 2000. Description Skevington, J.H. 2003. Revision of Australian Eudorylas of a new Nearctic Sericomyia Meigen, 1803 Acze´l (Diptera, Pipunculidae). Studia Dipterologica, 9: (Diptera, Syrphidae). Dipteron, 3: 137–140. 621–672. Nixon, K.C., and Carpenter, J.M. 1994. On outgroups. Stackelberg, A.A. 1927. Ubersicht der palaarktischen Cladistics, 9: 413–426. Arten der Unterfamilie Cinxiinae (Diptera, Syrphidae), Osburn, R.C. 1908. British Columbia Syrphidae, new im Zusammenhang mit der Auffindung einer neuen species and additions to the list. The Canadian Art der Gattung Cinxia in Jakutien. Materialy Komissii Entomologist, 40: 1–14. po izucheniyu Yakutskoi ASSR [5Materiaux de la Osburn, R.C. 1926. A new species of the genus Condidea Comission pour l’etude de la Republique Autonome (Diptera, Syrphidae). Entomological News, 37: 51–53. Sovietique Socialiste Iakoute], 20: 1–27. Osten Sacken, C.R. 1875. A list of the North Swofford, D.L. 2001. PAUP*. Phylogenetic analysis American Syrphidae. Bulletin of the Buffalo using Parsimony (*and other methods). Sinauer Society of Natural Sciences, 3: 38–71. Associates, Inc., Sunderland, Massachusetts. Pape, T., and Thompson, F.C. (Editors). 2010. Thompson, F.C., Thompson, B.J., and Fairman, J.E. Systema Dipterorum, version 1.0 [online]. 2000. Only in Costa Rica: new Neotropical flower Available from http://www.diptera.org/ [accessed flies (Diptera: Syrphidae). Studia Dipterologica, 7: 2 March 2011]. 33–43. Portschinsky, J.A. 1873. Descriptions de quelques Thompson, F.C., Rotheray, G.E., and Zumbado, M.A. Dipteres nouveaux de la Siberie orientale. Horae 2010. Syrphidae (Flower Flies). In Manual of Societatis Entomologicae Rossicae, 9: 287–291. Central American Diptera. Edited by B.V. Brown, Portschinsky, J.A. 1881. Diptera europaea et asiatica A. Borkent, J.M. Cumming, D.M. Wood, N.E. nova aut minus cognita. Pars IIa. Horae Societatis Woodley, and M.A. Zumbado. NRC Research Entomologicae Rossicae, 16: 257–284. Press, Ottawa, Canada, pp. 763–792. Posada, D., and Crandall, K.A. 1998. MODELTEST: Thompson, F.C., Vockeroth, J.R., and Sedman, Y.S. testing the model of DNA substitution. Bioinformatics 1976. Family Syrphidae. A catalogue of the Applications Note, 14: 817–818. Diptera of the Americas south of the United Reemer, M., and Hippa, H. 2008. Review of the States 46, 195 pp. species of Pseudovolucella Shiraki, 1930 (Diptera: van Veen, M.P. 2004. Hoverflies of northwest Europe. Syrphidae). Tijdschrift voor Entomologie, 151: Identification keys to the Syrphidae. KNNV 77–93. Publishing, Utrecht, the Netherlands. Ronquist, F., and Huelsenbeck, J.P. 2003. MrBayes 3: Walker, F. 1849. List of the specimens of dipterous Bayesian phylogenetic inference under mixed insects in the collection of the British Museum. models. Bioinformatics Applications Note, 19: Parts II (231–484), III (485–687), and IV 1572–1574. (689–1172). British Museum (Natural History), Rotheray, G.E. 1993. Colour guide to hoverfly larvae London. (Diptera, Syrphidae). Dipterists Digest, 9: 1–156. Williston, S.W. 1887. Synopsis of the North American Schiner, J.R. 1860. Vorlaüfiger Commentar zum Syrphidae. Bulletin of U.S. National Museum, 31: dipterologischen Theile der ‘‘Fauna Austriaca,’’ 1–335. mit einer na¨heren Begru¨ndung der in derselben Wirth, W.W., Sedman, Y.S., and Weems, H.V., Jr. aufgenommenen neuen Dipteren-Gattungen. II. 1965. Family Syrphidae. Agriculture Handbook, Wiener Entomologische Monatschrift, 4: 208–216. 276: 557–625.

᭧ 2012 Her Majesty the Queen in Right of Canada 4 a.Etml o.14 2012 144, Vol. Entomol. Can. 244 Appendix 1. Sericomyia specimens sequenced

Database Accession number Country State Latitude decimal Longitude decimal Deposition Speciﬁc epithet GenBank number

09BEDI 458 Canada NL 48.49209 254.02230 BIOUG S. bifasciata HM435789 09BEDI 250 Canada NS 46.71310 260.38349 BIOUG S. bifasciata GU689904 CNCD 2215 Canada NS 46.73333 260.63333 CNC S. bifasciata JF442704 CNCD 2213 Canada QC 45.68333 276.08333 CNC S. bifasciata JF442702 CNCD 2214 USA NH 45.05 271.38333 CNC S. bifasciata JF442703 CNCD 1902 USA OR 44.23 2122.202 CNC S. chalcopyga JF442707 CNCD 2241 USA WA 48.7667 2121.8 CNC S. chalcopyga JF442706 CNCD 929 USA AK 59.889 2151.783 CNC S. chalcopyga JF442705 09BEDI 254 Canada NB 45.58219 264.98590 BIOUG S. chrysotoxoides GU689896 09BEDI 245 Canada NB 45.60319 264.98770 BIOUG S. chrysotoxoides GU689898 09BEDI 247 Canada NB 45.60319 264.98770 BIOUG S. chrysotoxoides GU689893 09BEDI 248 Canada NB 46.77149 264.96219 BIOUG S. chrysotoxoides GU689901 09BEDI 246 Canada NL 48.6236 253.96709 BIOUG S. chrysotoxoides GU689899 09BEDI 252 Canada NL 48.6236 253.96709 BIOUG S. chrysotoxoides GU689891 09BEDI 259 Canada NS 46.65530 260.3722 BIOUG S. chrysotoxoides GU689892 09BEDI 249 Canada NS 46.71310 260.38349 BIOUG S. chrysotoxoides GU689900 09BEDI 255 Canada NS 46.71310 260.38349 BIOUG S. chrysotoxoides GU689894

᭧ CNCD 2218 Canada NS 46.73333 260.63333 CNC S. chrysotoxoides JF442712

02HrMjsyteQeni ih fCanada of Right in Queen the Majesty Her 2012 09BEDI 256 Canada NS 46.81230 260.76800 BIOUG S. chrysotoxoides GU689902 09BEDI 40 Canada NS 46.81940 260.62260 BIOUG S. chrysotoxoides GU689824 09BEDI 257 Canada NS 46.89670 260.78329 BIOUG S. chrysotoxoides GU689903 09BEDI 258 Canada NS 46.89670 260.78329 BIOUG S. chrysotoxoides GU689895 09BEDI 260 Canada NS 46.89670 260.78329 BIOUG S. chrysotoxoides GU689897 CNCD 2217 Canada ON 42.75 281.11666 CNC S. chrysotoxoides JF442710 08TML 547 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides GU806501 08TML 555 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides GU806732 08TML 557 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides GU806728 08TML 2408 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides HQ982383 08TML 2439 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides HQ982413 08TML 2441 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides HQ982415 08TML 2449 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides HQ982423 08TML 2457 Canada ON 43.44639 280.25119 BIOUG S. chrysotoxoides HQ982429 CNCD 4996 Canada QC 45.46638 274.32638 CNC S. chrysotoxoides JF442708 kvntnadThompson and Skevington Appendix 1. Continued

Database Accession number Country State Latitude decimal Longitude decimal Deposition Speciﬁc epithet GenBank number

CNCD 65 Canada QC 45.46638 274.32638 CNC S. chrysotoxoides JF442711 CNCD 4995 Canada QC 47.09558 269.986288 CNC S. chrysotoxoides JF442709 08BDIP 1022 Canada AB 49.00799 2114.0250 BIOUG S. flagrans JF442717 USNM ENT 35368 USA CA 37.63437 2118.25567 USNM S. flagrans JF442714 USNM ENT 35369 USA CA 37.63437 2118.25567 USNM S. flagrans JF442713 CNCD 1564 USA OR 42.66638 2118.5652 CNC S. flagrans JF442716 CNCD 1567 USA OR 42.66638 2118.5652 CNC S. flagrans JF442718 CNCD 1976 USA OR 42.63638 2118.5766 CNC S. flagrans JF442715 07ROBE 10093 Canada MB 58.73099 293.77999 BIOUG S. jakutica JF442723 CNCD 2230 Canada NT 65.26666 2126.8166 CNC S. jakutica JF442724 JSS 24959 Canada NT 69.43385 2133.01473 CNC S. jakutica JF442720 CNCD 2212 Canada NT 69.45 2133.0333 CNC S. jakutica JF442725 JSS 24953 Canada NT 71.98653 2125.243 CNC S. jakutica JF442722 JSS 24954 Canada NT 71.98653 2125.243 CNC S. jakutica JF442721 debu 1047493 Canada YT 68.16402 2136.9882 DEBU S. jakutica JF442719 CNCD 2111 Sweden 63.31666 212.1 CNC S. lappona JF442729 CNCD 2221 Sweden 63.31666 212.1 CNC S. lappona JF442727

᭧ CNCD 2229 Russia CNC S. lappona JF442728

02HrMjsyteQeni ih fCanada of Right in Queen the Majesty Her 2012 09BEDI 196 Canada NS 46.77939 260.33309 BIOUG S. lata GU689981 CNCD 18 Canada ON 45.96055 278.06111 CNC S. lata JF442732 08BDIP 373 Canada ON 48.59000 286.29000 BIOUG S. lata JF442730 CNCD 2222 Canada QC 45.48333 276.25 CNC S. lata JF442731 CNCD 2223 Canada QC 45.68138 276.05027 CNC S. lata JF442733 08BDIP 1365 Canada MB 50.87300 2100.0500 BIOUG S. militaris JF442735 08BDIP 1367 Canada MB 50.87300 2100.0500 BIOUG S. militaris JF442734 CNCD 9376 Canada MB 58.61861 293.82888 CNC S. militaris JF442738 07ROBE 4537 Canada MB 58.63000 293.81900 BIOUG S. militaris JF442739 07ROBE 4538 Canada MB 58.63000 293.81900 BIOUG S. militaris JF442740 09BEDI 98 Canada NB 45.54959 265.018898 BIOUG S. militaris GU689825 09BEDI 151 Canada NB 45.60319 264.98770 BIOUG S. militaris GU689978 09BEDI 152 Canada NB 45.60319 264.98770 BIOUG S. militaris GU689975

09BEDI 153 Canada NB 45.60319 264.98770 BIOUG S. militaris GU689982 245 4 a.Etml o.14 2012 144, Vol. Entomol. Can. 246 Appendix 1. Continued

Database Accession number Country State Latitude decimal Longitude decimal Deposition Speciﬁc epithet GenBank number

09BEDI 155 Canada NB 46.77149 264.96219 BIOUG S. militaris GU689980 09BEDI 150 Canada NB 46.81999 264.95369 BIOUG S. militaris GU689977 09BEDI 156 Canada NB 46.81999 264.95369 BIOUG S. militaris GU689973 09BEDI 154 Canada NL 49.62450 257.92269 BIOUG S. militaris GU689976 CNCD 2225 Canada NS 46.73333 260.63333 CNC S. militaris JF442744 09BEDI 149 Canada NS 46.89670 260.78329 BIOUG S. militaris GU689979 CNCD 2224 Canada ON 43.23722 281.87083 CNC S. militaris JF442743 08TML 548 Canada ON 43.44639 280.25119 BIOUG S. militaris GU806532 08TML 2434 Canada ON 43.44639 280.25119 BIOUG S. militaris HQ982408 08TML 2451 Canada ON 43.44639 280.25119 BIOUG S. militaris HQ982425 08BDIP 1656 Canada ON 48.58900 286.29199 BIOUG S. militaris JF442737 08BDIP 1657 Canada ON 48.58900 286.29199 BIOUG S. militaris JF442736 CNCD 4998 Canada QC 45.46638 274.32638 CNC S. militaris JF442741 CNCD 4997 USA NH 44.45 271.55 CNC S. militaris JF442742 CNCD 2228 Canada QC 53.71666 276.06666 CNC S. nigra JF442746 CNCD 2227 Canada YT CNC S. nigra JF442745 CNCD 2220 Japan 36.2 2137.75 CNC S. sachalinica JF442726

᭧ CNCD 9475 Canada MB 58.61861 293.82888 CNC S. sexfasciata JF442752

02HrMjsyteQeni ih fCanada of Right in Queen the Majesty Her 2012 CNCD 9518 Canada MB 58.61861 293.82888 CNC S. sexfasciata JF442751 CNCD 9573 Canada MB 58.61861 293.82888 CNC S. sexfasciata JF442750 CNCD 9609 Canada MB 58.61861 293.82888 CNC S. sexfasciata JF442749 CHU06-SYR 123 Canada MB 58.62900 293.79799 CNC S. sexfasciata JF442753 07ROBE 10084 Canada MB 58.63000 293.81900 BIOUG S. sexfasciata JF442748 07ROBE-JW 952 Canada MB 58.73059 293.78040 BIOUG S. sexfasciata HM860971 07ROBE-JW 953 Canada MB 58.73059 293.78040 BIOUG S. sexfasciata HM860972 CNCD 35781 USA AK 65.58333 2156.50056 CNC S. sexfasciata JF442747 debu 170285 Canada ON 47.5635 284.7186 DEBU S. slossonae JF442755 CNCD 30699 USA ME 44.81666 270.98333 CNC S. slossonae JF442756 CNCD 30712 Canada NT 71.58863 2123.4441 CNC S. tolli JF442760 CNCD 2234 Canada YT 69.58333 2139.0833 CNC S. tolli JF442761 CNCD 30742 Canada YT 69.58843 2139.0838 CNC S. tolli JF442759 CNCD 2237 Canada NS 46.73333 260.63333 CNC S. transversa JF442762 kvntnadThompson and Skevington Appendix 1. Continued

Database Accession number Country State Latitude decimal Longitude decimal Deposition Speciﬁc epithet GenBank number

CNCD 2236 Canada ON 45.33333 275.58333 CNC S. transversa JF442763 CNCD 2235 Canada QC 54.51666 269.86666 CNC S. transversa JF442754 JSS 19703 Canada QC 48.82944 272.35115 CNC S. vockerothi JF442758 JSS 19210 Canada QC 49.00028 272.34763 USNM S. vockerothi JF442757 CNCD 2240 Canada YT CNC S. woodi JF442767 debu 1047488 Canada YT 65.84188 2137.7567 DEBU S. woodi JF442764 CNCD 2232 Canada YT 66.125 2136.1166 CNC S. woodi JF442765 ᭧ 02HrMjsyteQeni ih fCanada of Right in Queen the Majesty Her 2012 247