ISSN 1713-7845 JOURNAL of the ENTOMOLOGICAL SOCIETY OF ONTARIO Volume One Hundred and Forty Six 2015 Published 2015 ISSN 1713-7845 JOURNAL of the ENTOMOLOGICAL SOCIETY of ONTARIO Volume One Hundred and Forty Six 2015 THE ENTOMOLOGICAL SOCIETY OF ONTARIO OFFICERS AND GOVERNORS 2014-2015 President: I. SCOTT Agriculture and Agri-Food Canada, 1391 Sandford St. London, ONN5V 4T3 ian.scott@agr.gc.ca President-Elect: J. GIBSON Royal BC Museum 675 Belleville Street, Victoria, BC V8W 9W2 JGibson@royalbcmuseum.bc.ca Past President: J. MCNEIL Department of Biology, University of Western Ontario Biological and Geological Sciences Building London, ON N6A 5B7 jmcneil2@uwo.ca Secretary: M. LOCKE Vista Centre, 1830 Bank St, PO Box 83025 Ottawa, ON K1V 1A3 entsocont.membership@gmail.com Treasurer: S. LI Natural Resources Canada, Canadian Forest Service 960 Carling Ave., Building 57 Ottawa, ON K1A 0C6 sli@nrcan.gc.ca Directors: D. BERESFORD (2015-2017) Biology Department, Trent University 1600 West Bank Drive, Peterborough, ON K9J 7B8 davi dberesford@ trentu. c a S. CARDINAL (2013-2015) Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Ave, Ottawa, ON K1A 0C6 sophie.cardinal@agr.gc.ca A. GUIDOTTI (2014-2016) Department of Natural History, Royal Ontario Museum 100 Queen's Park, Toronto, ON M5S 2C6 antoniag@rom. on.ca W. KNEE (2014-2016) Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Avenue, Ottawa, ON K1A 0C6 whknee@ gmail. c om B. SINCLAIR (2013-2015) Department of Biology, University of Western Ontario London, ONN6A 5B7 bsincla7@uwo. ca J. SMITH (2015-2017) University of Guelph Ridgetown Campus 120 Main St. E, Ridgetown, ON NOP 2C0 jocelyn. smith@uo guelph. ca ESO Regional Rep to ESC: P. BOUCHARD Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Avenue, Ottawa, ON K1A 0C6 Patrice.Bouchard@agr,gc>ca Librarian: J. BRETT Library, University of Guelph Guelph, ON NIG 2W1 j imbrett@uoguelph.ca Webmaster & Newsletter Editor: T. BURT Agriculture and Agri-Food Canada, K.W. Neatby Building, 960 Carling Avenue, Ottawa, ON K1A0C6 trevburt@ gmail. com Newsletter Editor: A. LINDEMAN Department of Biology, Carleton University 209 Nesbitt Building, 1125 Colonel By Drive Ottawa, ON K1S5B6 amanda .lindeman@ gmail .com Student Representative: L, DES MARTEAUX Department of Biology, University of Western Ontario Biological and Geological Sciences Building London, ON N6A5B7 ldesmart@uwo.ca Student Representative: C. PEET-PARE Department of Biology, Carleton University 1125 Colonel By Drive, Ottawa, ON K1S 5B6 cpeetpare@gmail.com JESO Editor: C. MACQUARRIE Natural Resources Canada, Canadian Forest Service Great Lakes Forestry Centre 1219 Queen Street East, Sault Ste. Marie, ON P6A 2E5 JE SOE di tor @ gmail. com Technical Editor: T. ONUFERKO Department of Biology, York University Lumbers Building (RM 345), 4700 Keele Street Toronto, ON M3J 1P3 onuferko@y orku. ca Associate Editors: A. BENNETT Agriculture and Agri-Food Canada 960 Carling Ave., Ottawa ON K1A 06C N. CARTER Engage Agro Corporation 1030 Gordon St., Guelph, ON, NIG 4X5 neilcarter@ engageagro .com J. SKEVINGTON Agriculture and Agri-Food Canada Eastern Cereal and Oilseed Research Centre 960 Carling Ave., Ottawa, ON K1A 0C6 JESO Volume 146, 2015 JOURNAL of the ENTOMOLOGICAL SOCIETY OF ONTARIO VOLUME 146 2015 This is my first volume as Editor of the Journal of the Entomological Society of Ontario. It is my great pleasure to take over the reins from John Huber. I would like to take this opportunity to thank John for his years of service to JESO, and for all his helpful advice over the past year as I have settled into the position. It is quite a daunting task to take on the editorship of JESO at this period in its history. As most readers of this journal are aware, the fate of this publication has been up in the air over the past few years. Changes in the publication landscape in the recent past have resulted in an explosion of journals, and there has never been more choice available to authors as to where they can submit their work. This has had the rather unpleasant side effect of contributing to the reduction in the number of papers that are submitted to JESO. So where in that vast landscape of publication choices does this leave a small outpost like JESO? That is a question I hope to be able to answer during my tenure. It is clear that if JESO is to survive it must adapt. We are fortunate that some of this work has begun and that we have a society and an executive that is supportive of further adaptations of JESO. We are also fortunate that there is a strong foundation of almost 150 years of publication upon which we can build. Under John’s leadership, JESO has also begun some of the renovations necessary to bring us into the modern era, notably the adoption of an online-only publication and the archiving of the back catalogue with the Biodiversity Heritage Library. My task is to continue this work. My intent for 2016 is to complete the second stage of renovations to JESO by bringing us to an integrated submission and publication system that will allow our authors to submit papers using a modern web interface and allow our editorial team to more quickly review, edit, and publish your work. With the move to the online publication system, I also plan to begin the task of amalgamating JESO’s back catalogue with the existing online offerings from the past few years. The long-term goal is that within the next five years the entire back catalogue of JESO will be online, searchable and indexed and available at one location. My hope is that through this process we can raise the profile of JESO to the point that it becomes a first-choice for many authors. However, at the same time I am a realist. JESO will never challenge the large journals of the two major North American entomological societies. But that is okay. We can, though, become a journal of choice for regional contributions, particularly for works of taxonomy and systematics, and applied entomology, which have been traditional strengths for JESO. I also see JESO as a place where natural history about insects and arthropods can find a welcoming home. Moreover, I think JESO can serve as a home for those authors that have a passion for entomology, 1 JESO Volume 146, 2015 even if entomology may not be their profession. These are potential authors who may never think of submitting to academic journals but still have important stories to tell. JESO may never win the war of the impact factors, but we can be a place where good work will find a welcome home. So I end with what may be called the editor’s plea: Send us your submissions! JESO has survived because of the support of its readers. I’m willing to bet that each of you has at least one paper that is looking for a welcome home but languishes at the back of your file cabinet (be it digital or physical). This is the kind of support that JESO needs as we begin to adapt to the new reality. During this period we also ask that you liken us to your favourite local establishment: We may be under new management and the sign out front says ‘Under Renovations’ but we still want your business and are no less dedicated to your satisfaction. Please stick with us while we go through this process. It will be worth it. Whereas in previous years publications were made annually and undesirably after the NSERC application cycle, articles can now be posted online instantly upon acceptance. This is a major benefit to graduate students in particular who wish to submit their work for publication prior to applying for scholarships. The other major benefit is that we are now an online open access publication, and do not penalize authors for submitting large numbers of pages or images. Consider us as you contemplate on where to submit those lengthy taxonomic revisions. Before I go, I would like to end with a thank you to the associate editors of JESO and the reviewers who gave their time to help us with this issue. I would also like to thank Neil Carter for his service to JESO over the years and welcome Jocelyn Smith as a new associate editor. Technical editing of JESO is done by Tom Onuferko. Together these folks have helped put together the following 54 pages. I hope you enjoy reading them. Submissions for volume 147 are now being accepted and can be sent to JESOEditor@gmail.com. Chris MacQuarrie Editor 2 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 A LIST OF BEE SPECIES (HYMENOPTERA: APOIDEA) RECORDED FROM THREE MUNICIPALITIES IN THE NIAGARA REGION OF ONTARIO, INCLUDING A NEW RECORD OF LASIOGLOSSUMFURUNCULUM GIBBS (HALICTIDAE) IN CANADA T. M. ONUFERKO 1 *, R. KUTBY 2 , M. H. RICHARDS 3 department of Biology, York University, 4700 Keele Street, Toronto, Ontario, Canada M3J 1P3 email, onuferko@yorku.ca Abstract J. ent. Soc. Ont. 146: 3-22 We carried out an extensive survey of bee species in the Niagara Region of Ontario, Canada, by sampling various sites within three municipalities from 2003 to 2013. The municipalities were St. Catharines, Port Colborne, and Wainfleet. Sampling mainly consisted of pan-trapping, but also included sweeping through vegetation and targeted collection from flowers. In the longest ongoing survey of a bee community to date in Canada, we collected 51,842 bee specimens comprising nearly 150 valid species, of which 30 were not previously recorded for the region. We also present the first record of the rare sweat bee species Lasioglossum furunculum Gibbs (Hymenoptera: Halictidae) in Canada, which was previously known only from Massachusetts, United States of America. Published November 2015 Introduction Our first survey of a bee community in the Niagara Region of southern Ontario, Canada, was carried out in 2003 at 8 sites on the Brock University campus and the adjacent Glenridge Quarry Naturalization Site in St. Catharines in the northeastern tier of the Niagara Peninsula (43.1 °N, 79.2 °W; Richards et al. 2011). The St. Catharines sites included relatively undisturbed meadows and fields on the Brock University campus, as well as regeneration sites at the Glenridge Quarry Naturalization Site, a former landfill. * Author to whom all correspondence should be addressed. 2 Department of Biological Sciences, University of Calgary, Calgary, Alberta, Canada T2N 1N4 3 Department of Biological Sciences, Brock University, St. Catharines, Ontario, Canada L2S 3A1 3 Onuferko et al. JESO Volume 146, 2015 Descriptions of the St. Catharines sites and the history of the area (natural and in terms of human activity) were provided in Richards et al. (2011). The St. Catharines sites were sampled each year from 2004 until 2013, except 2007, for a total of ten years of sampling. From 2011 to 2013, bees were also systematically sampled at two landfill regeneration sites in southern Niagara Peninsula, the Elm Street Naturalization Site in Port Colborne, Ontario (42.9 °N, 79.3 °W), and the Station Road Naturalization Site in Wainfleet, Ontario (42.9 °N, 79.4 °W). The sites in Port Colborne and Wainfleet are located on sites that from the 1950s until 2009 and 2008, respectively, functioned as municipal landfills. By 2011, these landfills had been capped with clay, covered with soil, and planted with an array of flowering plant species native to North America. Our objective in the present study is to provide the list of bee species collected from our sites in these three municipalities of the Niagara Region. All three sampling areas are within the Carolinian Zone, which includes tallgrass prairie and woodland communities. Considering the proximity (< 30 km) of the sites and the longer sampling effort at St. Catharines, we expected that the species lists compiled for the Port Colborne and Wainfleet municipalities would be subsets of the St. Catharines list. Methods Bees were collected using three methods: pan-traps (2003-2006, 2008-2013), sweep-netting vegetation (2003-2005), and aerial netting from flowers (2003-2005, 2011-2013). Details on sampling methodology are provided in Richards et al. (2011) and Rutgers-Kelly and Richards (2013). While combining specimens from all these collecting methods maximized the number of species likely to be collected (Wilson et al. 2008), non¬ standard sampling across years and sites means that it is difficult to quantify and compare the proportional representations of bee species in the community. All specimens were pinned and labelled, and are currently deposited in the research collection of M. H. Richards at Brock University. The majority of specimens collected in St. Catharines after 2003 were identified by T M. Onuferko, and those collected in Port Colborne and Wainfleet from 2011-2013 were identified by R. Kutby and T. M. Onuferko. The following taxonomic keys were used to identify specimens in conjunction with online keys available on Discover Life (Ascher and Pickering 2015): Colla et al. (2011) for Bombas Latreille (Hymenoptera: Apidae); Gibbs (2010,2011) for Lasioglossnm Curtis (Hymenoptera: Halictidae) subgenus Dialictus Robertson; Gibbs et al. (2013) for Lasioglossnm subgenera Evylaeus Robertson, Hemihalictus Cockerell, and Sphecodogastra Ashmead; Mitchell (1960, 1962) for Halictus Latreille (Hymenoptera: Halictidae) and Sphecodes Latreille (Hymenoptera: Halictidae); McGinley (1986) for Lasioglossnm subgenera Lasioglossnm and Leuchalictus Warncke; Rehan and Richards (2008) and Rehan and Sheffield (2011) for Ceratina Latreille (Hymenoptera: Apidae); Rightmyer (2008) for Triepeolus Robertson (Hymenoptera: Apidae); and Sheffield et al. (2011b) fox Megachile Latreille (Hymenoptera: Megachilidae). Discover Life keys (Ascher and Pickering 2015) were used for the following genera: Agapostemon Guerin-Meneville (Hymenoptera: Apidae), Anthidinm Fabricius (Hymenoptera: Megachilidae), Anthophora Latreille (Hymenoptera: Apidae), Calliopsis Smith (Hymenoptera: Andrenidae), Chelostoma Latreille (Hymenoptera: Megachilidae), 4 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 Coelioxys Latreille (Hymenoptera: Megachilidae), Heriades Spinola (Hymenoptera: Megachilidae), Hoplitis Klug (Hymenoptera: Megachilidae), Hylaeus Fabricius (Hymenoptera: Colletidae), Melissodes Latreille (Hymenoptera: Apidae), Osmia Panzer (Hymenoptera: Megachilidae), and Stelis Panzer (Hymenoptera: Megachilidae). Females of the following species pairs are very difficult to differentiate, and identifications were largely based on male characters: Ceratina dupla Say versus C. mikmaqi Rehan and Sheffield, and Hylaeus affinis (Smith) versus H. modestus Say. Specimens of Nomada Scopoli (Hymenoptera: Apidae), a genus in need of revision, were kindly identified by Sam Droege (US Geological Survey, Patuxent Wildlife Research Center, Beltsville, Maryland). All Andrena Fabricius (Hymenoptera: Andrenidae) designations were made by Cory Sheffield (Royal Saskatchewan Museum, Regina, Saskatchewan), for which we are most grateful, and Jason Gibbs (Michigan State University, East Lansing, Michigan) graciously helped to identify many of the Lasioglossum specimens, including one new record for Canada. Results A total of 51,842 bee specimens were collected, comprising 149 species and 1 morphospecies of Nomada. Richards et al. (2011) had previously identified 124 species and morphospecies from the 2003 samples, including four distinct morphospecies and one unknown species of Nomada. In the present study, these Nomada have been collapsed into a single bidentate morphospecies group, as suggested by taxonomic expert Sam Droege. In the present study, 30 valid species not recorded by Richards et al. (2011) were identified (see Table 1 for a list of these and all other species sampled). All species belonged to the five most common bee families occurring in North America (Andrenidae, Apidae, Colletidae, Halictidae, and Megachilidae); no bees of the small and uncommon family Melittidae were sampled or observed. Of the 30 bee genera represented, only the cleptoparasitic genus Triepeolus was not previously recorded by Richards et al. (2011). Almost a third (9/30) of the new species added are cleptoparasitic or socially parasitic, one of which is described for the first time in Canada in the section that follows. The most speciose family sampled was Halictidae (54 species), and the least speciose was Colletidae (11 species). The present ranking of families by morphospecies richness (Halictidae > Apidae > Megachilidae > Andrenidae > Colletidae) generally matches that of Richards et al. (2011) (Halictidae > Apidae = Megachilidae > Andrenidae > Colletidae). New record for Canada: Lasioglossum ( Dialictus ) furunculum Gibbs Lasioglossum furunculum is a species that was recently described from Massachusetts, United States of America, from a single specimen (Gibbs 2011). It is most similar to Lasioglossum izawsum Gibbs (Hymenoptera: Halictidae), but differs in having no preapical tooth on the mandible (Fig. 1A) and an inner metatibial spur with four rather than three branches (Gibbs 2011). Females of another similar species, Lasioglossum simplex (Robertson) (Hymenoptera: Halictidae), lack a carinate pronotal ridge present in the two abovementioned species (Gibbs 2011). In all three species, the gena is subequal in width to the compound eye when viewed from the side (Fig. IB); it is conspicuously wider in other parasitic species (Gibbs 2011). 5 TABLE 1: A complete checklist of bee species sampled from 2003-2013 in southern St. Catharines, in Port Colborne, and in Wainfleet, Ontario, Canada. Species collected since the initial 2003 survey (Richards et al. 2011) are indicated by an asterisk (*). Species for which a life history trait is suspected but not confirmed, as in Lasioglossum spp., are indicated by a question mark (?). The presence of a species within a particular municipality is denoted with an ‘X’. Foraging habit is listed as N/A for parasitic species, which do not forage. Family and species Life History trait Origin Municipality Onuferko et al. JESO Volume 146, 2015 C /5 Id S-H Co R % oc § ~ O cd *-H Id S-H Id S-H 2 CD G CD o £ 6 £ f ° o ^2u B o U CD G (D a CD G G G R O cd CD O U R 53 CD 'r' R ^3 R J3 5s R X5 R U f R R R ^3 R 3s C3 R ^3 R CO 53 R 5s R ^3 R JR J 5! R X5 R X) * CD R •HSi R •SP R CD 'o' R ^3 R 55 5s R Jg R3 R X) cd R3 2 R R ^3 R 55 -R & R R ^3 R X| O cd R O § R R ^3 R 53 5s R ^3 R X| 3 S-H CD G CD o R CD -R G 'r R ^3 R 53 5s R R3 R X) X X X X X CD CD CCS Uh X5 G G3 G -X T3 G -a G -X -a G -X T3 G -X T3 G -X -a G -X T3 G -X -a G -X X5 G -X -a G -X -a G -X T3 G -X -a G -X -a G -X — • g — • g D g -H g -H g -H g -H g — » g — • g -H g -H g g — • g -H g -H g -J g O a a a o o a a O a a a O a a a b b b b b b b b b b b b cd cd cd cd cd cd cd cd cd cd cd cd cd cd cd _+^ o ’o ’o o "o o o O o o ’o O o o ’o c/^ in m m CO c/^ m GO m m GO GO GO GO GO G G G G S-H OD cd O o ^3 O C/5 ti OD X) O O G X S-H CD -G r ■» CD 2 ■g -G s—* S -G s—* S ISO OD -Q O C/5 C/5 CD S-H u T3 S-H CD S-H CD M CD O -G C/5 ti OD X) o O C/5 ti OD _Q CD ’£ OD cd X _S-H o X. a S-H o £ O u ■c o Ph SB CD G Th g -G U t si g ’ 5 b ’C o X> S 3 X &D g ’ 5 b 3 S-H O X X 3 X bfl c X S 3 X o o t/3 CD s> s> S> Sx* \> r\ r\ r\ r\ rS X X X X X X X .> ’X ’X '■0 ’X ’X X 0 X ’X ’X ’X ’X ’X ’X ’X ’X c3 cd c3 c3 c3 >< c3 c3 c3 c3 c3 c3 c3 z Z z z z w z Z z z z z z z z 13 13 13 O c 3 13 13 13 13 13 13 *■■* ~G o C/3 Gs 'I*** o R jg IS R G & jg R G 1 GS a Cu 03 0. SB ‘3 a SB o -S Q. u © E •a s 3 u © a. SB <*5 © 3 2 ’a < ~a o o CD - 4 -* C/3 b 3 o C/3 G O CO SB a u G R *s R "i .8 **»* O I o ■R & R -a G 3 a o b 3 o C/3 £ Lh X G 3 o lR O ~R G cu 5 o -R §* -R R £ u T3 G 3 O Lh EP CD T 3 G X T 3 G "S © ”2 G cd G 3 ,3 3 a *5 a o So T 3 G 3 G G • 3 = Lh 3 PQ Lj SB Q o Oslh R R a a bb° -a s»ep 'GGS" R ^ 12 G r— 1 CL) rO 1 SO S gX •§ cqp - 4 -* CO ■ 4 —* CO -LH CB -L-* SB 13 13 "3 "3 CD CD <5 © G G G G (D CD CD © a a O O "O "O TS "O G G G C G G Lh 3 3 O J-H O i~C 0 0 2 EP -a G EP CD T 3 G G 3 G 3 a in O O 0 s §0 .2 .2 .2 .2 CD .2 .2 .2 .2 .2 ’3 3 '3 '3 1 G ^ 'CD S ’3 '3 ’3 ’3 '3 0 0 0 0 0 0 0 0 0 CO CO CO CO CO CO CO CO CO G G G G O c 3 G G G G G w w W uq Oh W W W W W 03 3 © 3 G g 23 G R •k^G X X) _ Vh 5 o o • ?««* § -R R O 2 - a r S -fi V a .> > .> .> > > .> .> .> .> 3 3 3 3 3 3 3 t3 t3 3 3 3 3 3 3 z z z z z z z X X Z Z Z Z Z Z CO _ 3 g S-H 2 3 3 3 ^0? G CD •4—* CO -4-* CO -4-* CO -4—» CO -4-* CO a ■ »■** .2 .2 3 J dh 3 3 13 S-H 13 i-H 13 i-H 13 J-H 13 "3. as Oo H, a 0 Oh ' '33 a T3 o a 3 3 a a O 03 S-H O 1> G a> O CD G a> a »* O (Aste N/A N/A N/A N/A N/A N/A N/A X X X X T3 3 3 CD —< •3 3 —5 —4 g —4 a —> a O 0 0 o O ^ DO §3 to P bo3 Z dP d 3 b b b b b b b G G G G G G G •GI •^H •Gi •1h •G! •G ’o O O 'o O O O m m m m m m C/^ C/5 03 s-h G a. o -<—> Gh ° S3 £ CO ^ *2 +-4 *£< G co cs — 1 O hJ o *1*^ Co P ,a> as o to =o "5 a o *. - d =0 w oj 2 3 -73 34 § y Co A as O DC Co I O U T3 3 3 co ^3 O DC DC S tq ^ ^ X. ^ C3"^ O a: O 50 &) fo «0 G *0 I II co m s si 2d 30 3 53 2 g| o a S,S J2 ■> as < 2 3 34 CD o U •>*** a a -a -a a a § 3 O DC d 3 -3 O d a o CO DO 8 ! 3 ai a a § * 3 o DC d 3 -3 d 3 a 3 a "a a 3 o DC 3 -3 Si 'I Jo DO a cu o a a a § 30 3 a 3 •<»»» 3 ~a *§ a a I * 3 o DC -d 3 -3 S DO as o a a § * 8 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 a. OD U G O C/5 C/5 OD S-H u s a CD Q 5 § * G O C/5 C/5 2 o Q CD Q I a; as a X 3 O £ C/3 C/3 &b § 5 cD d a a r*» a a 3 D ,a -a to <0 X ,a ^a 53 "0 "a d a a, 5T • !■*» Ei; a. a § * * 03 A as (W D X a o S, 3 | ~ D X 33 t» — g 1—1 d -S ^ 33 5, .2 b, 82 8 5d -2 BbS d' '?u 3 O C/5 C/5 2 u a a 8 *N>* CD s Q CD G O co CO CD S-H u co s a 33 Si, 5 o a ^3 a c o s X § 33 I 33 33 O C/3 c/3 CD S-H u to R a to a 000 ODD c/3 3 CD 33 33 2 to R 33 a S3 33 1? R CD a a ,a £ 2 OD CD O U CD J3 co CD CO S CD t a 3D J3 £ G O co CO 2 u a 5 33 a, Df <33 • HH CD co S CD £ * 33 o 03 d CD X O o-n Qh X w X 00 t—> '»■*« Co R CD •2 o R CO R a 2 & CO 2 d, CD £ & CO 2 co R CD OD ■4—» CO _ G G G G G G G G G G G CD ■G •G G G G G G G G G O O O ’o ’o O ’o ’o O O O GO GO C/) CO CO CO CO CO CO CO CO C/3 to A is O & CO 2 a. CD ^2 &X) 33 X CD 1 CD S-H X C/3 a Hh, a a **«» 3 . 33 a a, d a £ * 9 Onuferko et al. JESO Volume 146, 2015 73 CD 3 a • i—H "S O o M P PQ £ b 13 2 - 'o '2 3 s O U o Oh tz> CD _g "C 3 X t3 u C/2 c CD > ’■& '■& ’■& ’-P "P 'P 'P 'P 'P 'P *c c3 c3 c3 c3 C3 c3 c3 C3 c3 c3 c3 c3 c3 c3 c3 c3 c3 o z z z z z z z z Z z z z z z z z z b o HH C/5 E ,3 X> 3 -3 OX) 3 ’5b 3 S-H O Oh -3 3 33 OD G "-3 05 3 X CD (D O. 3 33 3 3 2^ 3 Oh XXX X X X X X X XXX X X X X X X s> s> Sx< Sx< Sx< \> rS rS rS rS rS rS CD P C/5 P CD ^ P / ^ S Qu 3 42 o e a C3 -o a 3 O d 5 53 C3 •»*«* S^, a- a S»»» as a a 3 a -a 2 2 Oh 3 5 C/2 3 5 a 5 o 3 a •§ ja *»» 7s 3 3 a 3 a a a Q. Q Q S O O O O co O 5 O 2 O g Q r/) ^ Qx Q <| daj Sf^ co to O X X K> K> K> K> K> K> K> K> K> K> K> r\ r\ r\ r\ r\ r\ r\ r\ r\ r\ r\ 13 13 13 13 13 "2 P CD CD ^ cd cd 13 13 13 13 13 13 13 ¥-3 3 2 13 (D (D (D CD *•«» ^ Q q. q q Q o a -p a a "a? a Geo C co _ co co«o O^o o C<3 ^5 C<3 .2 S-2 Si 552 02 &1 S’! SI g>| S" Co o Co O Co 3 <5 S 1 ^ w-^ S-hJ ainp 10 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 s S-H o X ~o U t: o Oh OT s> s> s> s> rN rN rN rS rS X X XXX X X X X X X s> s> s> s> s> s> s> rN rN rS rN rN rS rS X XXX CD 3 X I 5 O 3 f x £5 ^ £ § 8 S -S 2 Z e QO CD -a o I -a 3 3 2 O T3 3 3 2 a T3 3 3 2 O (D X CD ■SP § a • »«. Q a 03 03 o SP ■§ XI 5 "a o 3 a 3 S a a 03 03 o £ 1 XI ■SC¬ OT X X O OT X X O p § 'Ki "a *)•>«» Q a 03 00 O & 'I XI CO 5 a S3 § a "a Q a 03 03 o 'I XI Co Co 3 cl <3 03 a a Q o a o CD a a a ,a "a ■*-* S3 bo x So X >0 > 0 g3 o CD ^ o S2 S-. ^o (Tj ^o G X E a to Cl S A ,0 03 <0 o 'I XI ^ a Lh 03 CD 03 X o ■ a-Bb 3 X *« "a P Co =0 -s O | §> k*- 2 8 as a a a as 2 a cl X -a 3 3 2 a a 5 o a a 6 «-2 3 03 X 53 ox X X ’-G ’G *G "G ’G *G "G "G "G *G *G "G ’G 'G 'G G G c3 z z z z z z z z z z z z z z z z 03 o3 G o3 03 03 o3 03 03 03 03 03 o3 <3 < a3 a3 a3 CD (D < < a3 CD (D D P P I CD 3 Sh O X O U o Oh c/5 CD _g "C 3 X b u C/2 3 ’5b ’C o X 3 -3 3D 3 '5b 3 S-H O Oh -3 3 33 3D G H c/5 3 X 33 3 -3 CD o 30 CD (D P. C/5 33 3 3 3 Oh <1> d) d) d> d) d> d> d> d> d) _o 0 a/ > > .> > '> > '> > > > > .> '■& ’-p 'P P P P p p p p p p p 0 0 P c3 c3 c3 c3 c3 c3 C3 c3 c3 c3 c3 c3 c3 X X c3 z z Z Z Z z z z z z z z z w w z X 73 3 3 2 o a Cj • >•*. Q -a 3 3 2 O \j>* S> \j>* s> XN XN XN XN xS XXX -4—* C/5 -4-* C/5 ■4—» C/5 -4—* C/5 -4-* C/5 -4—* C/5 O- -4—* C/5 ■4—> C/5 ■4—» C/5 -4—* C/5 -4—* C/5 -4—* C/5 -4-* C/5 -4-* C/5 -4-* C/5 -4-* C/5 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 13 a/ a3 CD CD CD CD d> CD D D D D D D D D c p P P P P p P p p P P P P p P a> a/ a/ 2 2 2 2 2 2 2 —4 2 —4 2 —4 2 —4 2 2 2 b O a O a O a O O 0 O O O O O- p D O C/5 o- .*2 CD O C/5 0 3 X 0 3 X ."2 0 0 C/5 ^2 D O C/5 O. ."2 O O C/5 .^2 CD O C/5 ."2 D O C/5 c- b b c- b b P P 3 3 p P P P P "o 0 H w b b H W W W W to CO b b b b b b b b b b "o 13 ”0 0 0 CO CO CO CO CO 3 3 o hJ Co S S3 £ 3 3 § S3 £ CD 3 O hJ "§ • ^ •I*** § CD CD § C/2 a !< £ s- M 3 53 53 •i*** •»««» Q Q ^5 CD A ■—. H 3 K a 3 ,7 b Q ^ 5S SS a jS O' Co _ C^ 3 co O jo 2-2 3-2 j >-2 x -2 ■§ § 8 8 co *o O P &0 O =0 C/5 O i'll 1 •i*«» » J —' Co O to S3 (V S3 <1 0<) co to O co Co O So - O 33 So ■g b-S S3 a =o <1^2 * =o co O 1? 'I I Co S S3 -a •»*«» 5 - 5 a 7s a o 3 a a S3 -a *2 S a -p a s 03 3 c^ 33 co O to D C\ r/"> C7 oSS v . a J 5 -S 5 S’ | g? C *1*^ *!-•» co O =o £ 53 c> C2 0<1 3 CD Cj 53 •t*»» a O 3 ?of •2 o 8 S3 tE"§ a S Jo 'a x> a o ’S? oi S 3 CD _ 5 a ? ~ a co '5,-2 a^ So o ~ o s 1^ -2 —I D -a Cl DT co ^0 O I 3 sg-~ B § X /NxNxNxNk^k^k^xnxnxnxNxNxnxnxnxnxn 12 quebecense (Crawford) *Sphecodes atlantis Mitchell Cleptoparasite Nests of N/A Native nest-building _ Lasioglossnm _ TABLE 1 continued... A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 .4? ”2 .& o '2 3 s b O -4-* C/5 X ,3 ■3 ^*5 >3 ■3 to ^0 rTS ^5 O C^H X =o O c^ 3 X i d .CJ s-h Ofl ° O O W 3 a S-H tLO 0 O O LO 3 X i .CJ s-h be o o 5 -g Sj CXI S .CD D d* c ax P G a^: G G l '-«} G ■"B G ‘^-1 Z< 'S 5 oo a “ m > 13 S3 d x c a G *~Ci o3 1 —-3 D -HH on cS S-H 3 o. o -on G. D U t/5 as S-H as Gh O h-» G. D u D _-hh oo as S-H ra Gh O -4-» CL D U C/5 § 2 -a g to -G IS o g G d: Cl X 1 G O oo d D X c§ g 2 G -a to -G d o g G •d Cl X 1 G o oo d D X c§ a g a a K 2 to A IS o g •d Cl X 1 - 4 —» C /5 - 4 —* C /5 s—* C /5 G G 13 (D (D (D G G G CD CD CD O O a d m nl v G o -ss g .-3 oo ;> G r g O U C/5 CD G £ u as ‘3 D Q. as fS w D 3 ■a D 3 6JD D s S a ,a o G H—hhC ^ U -5 -a o d JTx ^ £ R a S3 >2 s >S -a a a i? ~ o S d a d a p g a c g 3E a a a 4= a*^ X X X X X X X (D CD CD (D o o o CD ‘-G ‘-G ‘G ’G d o d o d o ‘G G G G G X X X G X X X X w w w z a a cs I CJ £ o b b b b "o O O "o t/5 t/5 t/5 t/5 CD G CD O iS 3 G G Oh ! C/5 CD z C/5 G !-h G Gh o - 4 —> Gh CD s =1 D § MS,, ‘ G G\ CD B§ D a S B CD Is O CXO g O o 2) a G a a Cl a U G GO a s a is S G O £ "g o G O 2 o I "g o CJ 13 Megachile brevis , M. centuncularis, and M. mendica _ TABLE 1 continued... Onuferko et al. JESO Volume 146, 2015 A 13 2 - 3 '2 =3 b o -4—> C/5 2 , .> .> > > > .> ■ 2 > ’-G "G '-G ’-G ’G G G o G o G G G C/5 C/5 o d o G G G G G G G X G X G G G o X z z z z z Z Z W Z W Z Z Z Ph Sz W cc3 35 ■M C/5 s—* C/5 -4—* C/5 -4-* C/5 -4-* C/5 -4-* C/5 -4—* C/5 -4—* C/5 ■4—* C/5 -4-* C/5 -4-* C/5 -4-* C/5 -4—* C/5 -4-* C/5 -4—* C/5 OX) _G 13 13 G 13 13 13 13 13 13 13 13 G G 13 G 5b < <5 (D D5 KS S-H o5 d O 53 CD 53 *n 35 O d, C5 O’ w 5 35 ■g 1 Oh D. % S "5 o a o o s b g • *»». "a o O -a S-H 1 G S-H u Cl I o I to ,A> 53 • HHs L. I 55 O do t/3 CD S-H u 53 O 5. 53 > a a l O to d d 53 • HHs L. I 55 O di di (D S-H u 5 53 S: '<■*>* fc. CJ 6 CL I a § to ,a d 53 • H>H L. I G O C/5 C/5 5 R £ >5 o 53 5 ~53 ‘ HH .*>5 I G O C/5 C/5 2 u 5J -a g d. 2? CD 5 -a I S-H CD -G o G G > 2 Oh, 5 53 3 a, CD 5 -a y* I 33 xj a -S & i § 2 -2 £ o ^ sS d C A{J CD «o '—- 33- a: cd 53 a -a . a is -a a £ a I * &5 ■<; ^ a o a to fc d S CD d X> -2 2 a 05i s — tq CD w G 5J a, ? in > g -a -S' ?| .2 5 S-H O X) dG a2i a -a -a 5 d a S g^ G O C/5 C/5 2 u a a a 2 S' • HHi -a a §> a o S 55 O DC DC i ^ cd ai - Dh - ^ IJ^) -a ® -a a -2 a S -a S bo a s£* ad I al ' HHi -a a a X SXSXSX G G G G G G G G G 55 d 2 § G G G ’> ’> ’> ’> ’> ’> ’> ’> ’> 2 > § o ’> ’> '> d c3 G G G G G G G G G 2 2 K 2)0 G G G Su u u u u U u u u o febu u o u bbbbbbb b b b b bb bb ^3 £ ^3 ^3 ^3 ■2 ^3 ^3 ^3 o o O "o O o o o o o "o "o "o 3 05 t/3 t/5 03 O) 05 03 03 05 05 05 03 03 05 05 55 O DC DC CD S-H u a 5- -2 C/5 S S' 2 ° _ a a S'-S 3 a cd 3a ;s 1= -a a -a a s: a B S S I 3 2I 3 I al -a a a 3 a .2 o d “a a 5S a o a 5 5a a -a a a S Sd g 3d -a I a 14 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 s> s> s> s> rS rS rS rN rS S> S> S> Sx* rN rN rS rS r% (D (D (D (D (D (D (D (D (D (D CD .> .> .> .> .> .> .> .> .> .> G G G G G G G G G G G cd G G G G G G G G G G z z z z z z z z z z z G G G G G CD CD CD CD CD G G G G G CD CD (D CD CD a a a a a S 'M G ^ co CD co £ o g G J-H G CD a G s-h CD G CD a z r ^ CO _CD G X 73 d 73 3 13 G co 73 co • 2 G G CD co X CO • 2 G Ca to 0 R CO ^ R t S 0 Xh i G 1) X' CD 0 0 1 3 2 a 3 2 a G G GO 0 1 '1 u S CD -*—» GO 0 1 i u d R, 3 Q co O • l-H R- ■R g R, R, 5 Co 4) O CD .2 2 2 2 2 2 2 2 2 2 0 "0 0 "o 0 0 0 0 0 0 C/3 C/3 C/3 C/3 C/3 C/3 C/3 C/3 C/3 C/3 (« S3 S-. 3 X, O -4-* X 3 u C/3 "§ •§> A 3 R Q 03 o ■s R ,3 ~R O |) 3 O c/3 c/3 3 R u gS R 5 3 a G 03 O ■s R § *R O I ■ Sb £ tZ3 3 R Q 03 a a S3 vt o sS s § -a o |) 3 O Vi G G O Ui 3 a O CO CO CO CO CD G O 2 u 2 S-H O co CO 2 5 u .co CD S-H C/3 a 53 c u 53 R a b 3 E? R CD X* R 0 a 0 g> bo T-H4 R R R a Rh R - CO s: CO R R R R O Co Vi v, Vi 'Ki 0 O O O C^ CJ • **H Cj • N4 s R 53 R 53 R 53 6 <3 * 3 3 3 C SI R ?3 R S3 3 3 G O 03 C/3 2 u .03 i-a "S 5 -. 333 2 Co 2 Co O 1 X 3 S3 a 3 «? o ~ R; > bp .S3 X O a •|*»H **« X &3 X 1 -3 ^ ^ Vs S s| a x > § x ^ X 03 R E? 3b a T3 (U -o ’> 2 x C/3 G O 3 d 35 C/3 CD 73 C/3 =3 _o '> (D a X X 3 O R 3) ?3 03 O '§ <1 X CD 3) d S3 X CD CD CD d "§ b <3 dr 5 "a/ d X ^ 2 d w S3 v> X 'C £ 2 d jp C/3 S3 X 2 o d 1 CD X J3 & "S G S a 03 03 x 1 o C3 Q a 3 to 'O O n C/3 X S-H S3 X CD S-H CD T3 d G T3 CD 2 % CO .2 0 CD Gh co txD G '% o 53 R R s cb U a 03 03 X § ■ ^ _r> 1 I - ^ o <3 a; II $U Cb a ?a to O l 3 '1 "O CD -+-> CO S-H CD ts CD N £ S s 5 3 Hh 53 R k. CJ &0 bS) .G co cd CD td G CD T3 cd 2 % 2 CD -G G X) o CD Gh CO T3 (D jg *C o CO CD -G G G d S-H CD > CD CO CD -G ^G 13 _G G O 'G CD ’o CD S-H G O « G O ^ Z 1 R I i o; | w o X "bp -2 o S R ® $b O ^ X 3 a 5 X ~ a X) -R a - I £ a, £ ng 5p a R S bp >} o Oh '£3 • ■ 3! Vi hJ CD ^ t/3 X CD PR $ t: G G 0/ X c3 s X Ld a co £3 Rs o 53 "5c •2 ^ § CD C/3 o X -s-» -a G S3 G O X R O a R 53 .2 a R ■>-H a 53 a =<3 X R > CD S-H X o X (D CD d a s a Yi S>3 X O Z*) > I "o 5^ C5 «o ^0 O co 2 G -S CD o 0 ) 00 CD 1 CQ S3 X ~a d S3 d S3 2 CD C/3 3 O CQ X so X -2 w < CD X 3 2 5 c2 c2 o R3 8 § oT 00 C35 S-H (D G CD cd Ph G cd Uh cd5 00 ^ 2 X w C\ x C- I) ® OX) c CO G o 2 ^ cd S ^ x o O'n o 4—1 -a „ C/2 O X Vd 2 C- o x rl I £ CD d ■x o S XI C/3 CN 3 X a ___ < O . . X C/3 ’-' CD c/3 2 -2 2 a 2 O ox) x d o •R 00 > ON O -H X 3 2 o ^ 8 8 ° S, x r- b 3 a 3 "R 53 R I X o C/3 CD '5 3 Q. D3 c/s 3 3 ’0 a on £ H X X 0X5 0° d C3N °5 15 Onuferko et al. JESO Volume 146, 2015 We collected a single female specimen of L. furunculum on 9 September 2009 from St. Catharines on the periphery of the Glenridge Quarry Naturalization Site, just south of the Niagara Escarpment. Males of the species are unknown (Gibbs 2011). Given the similarity of this species to L. izowsum, the lack of DNA barcodes, and the limited number of individuals available for both species, it is possible that L. furunculum and L. izawsum are synonymous (Gibbs 2011). However, the two species are currently recognized as valid, and the St. Catharines specimen best matches the description of L. furunculum . The species is presumably a cleptoparasite or a social parasite of one or more of the nest-building Lasioglossum (Dialictus) species present at our St. Catharines sites. Discussion Observed morphospecies richness of the entire 10-year sample from the St. Catharines sites (147 species) effectively matched that predicted by the abundance-based coverage (ACE, 147 species) and Chaol estimators (mean ± SD = 145 ± 9.6) based on combined pan-trap, sweep-net, and flower-net collection data from 2003 (Richards et al. 2011). Only three species present in Port Colbome and Wainfleet were entirely absent in samples from St. Catharines; these were Lasioglossum obfongum (Lovell) (Hymenoptera: Halictidae), Lasioglossumpilosum (Smith) (Hymenoptera: Halictidae), and Osmiageorgica Cresson (Hymenoptera: Megachilidae). We collected far more species and morphospecies in St. Catharines (147) than in Port Colborne (64) and Wainfleet (61) (Table 1), which was expected since St. Catharines samples were collected for 10 years and in relatively undisturbed as well as regenerating sites. The current list undoubtedly reflects some biases in the sampling protocols used, and there are certain taxa that appear to be underrepresented in terms of diversity, or should be present given records in areas neighbouring the Niagara Peninsula but are entirely absent FIGURE 1: Face (A) and lateral view (B) of a female specimen of Lasioglossum furunculum Gibbs collected in southern St. Catharines in 2009. Note the absence of a preapical tooth on the mandible and narrow gena in lateral view. Scale bars = 1 mm. 16 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 from our samples. We never sampled from trees or tall shrubs, so the species presented here likely reflect a large subsample of the resident bee communities that forages at or near ground level. This might explain the comparatively low diversity of the genus Andrena , which frequently visit spring blooming trees and include multiple specialists of willows and other spring blooming plants. Given that pan-trapping was the main sampling method employed among years, it is not surprising that in our samples Colletes Latreille (Hymenoptera: Colletidae), a genus usually sampled very well in nets, not pans (Wilson et al. 2008), were low in both number and diversity. Forty bee genera occur in southern Ontario (Packer et al. 2007; Gibbs et al. 2014), and several of these ( Dieunomia Cockerell (Hymenoptera: Halictidae), Dianthidium Cockerell (Hymenoptera: Megachilidae), Paranthidium Cockerell and Cockerell (Hymenoptera: Megachilidae), Epeoloides Giraud (Hymenoptera: Apidae), Macropis Panzer (Hymenoptera: Melittidae), and Svastra Holmberg (Hymenoptera: Apidae)) are too rare or transient to be expected in our sites. The melittid genus Macropis ranges from Nova Scotia to Washington and south to Georgia (Hurd 1979) and collects floral oils from loosestrife (Lysimachia) Linnaeus (Primulaceae) (Cane et al. 1983), which was rare in our study sites. Some bees like Peponapis pruinosa (Say) (Hymenoptera: Apidae) are expected to be relatively common in the Niagara Region, but this species is restricted to areas where cultivated cucurbits, Cucurbita Linnaeus (Cucurbitaceae), are present. Perdita Smith (Hymenoptera: Andrenidae), another genus that should be present in our region, is largely composed of small, floral specialists. Holcopasites Ashmead (Hymenoptera: Apidae), absent from our collections, are small cleptoparasites of Calliopsis ; the latter was uncommon in our sites. Epeolus Latreille (Hymenoptera: Apidae), a genus of cleptoparasites of Colletes , was absent from our samples, although over half a dozen species are known from southern Ontario, including two species recorded from Port Colborne (Romankova 2004). Cleptoparasite and social parasite diversity overall may have been underrepresented in our collections. Our main method of sampling, pan trapping, likely biases collection toward small sweat bees (Halictidae), and underrepresents parasitic species, which spend more time searching for host nests than foraging, and larger bee species that can crawl out of pan traps should they fall in inadvertently (Cane et al. 2000; Wilson et al. 2008). A study by Cane et al. 2000 demonstrated that pan traps failed to catch most species of floral specialists associated with the creosote-bush, Larrea tridentata (DC.) Coville (Zygophyllaceae), below which the traps were set. The few specialist species present in our collections were mainly sampled from flowers or sweeps through vegetation. The number of species found in the present study is lower than that known from the Caledon Hills, located north and east of the Niagara Escarpment and close to the eastern limit of the Carolinian Zone in Ontario. Between two surveys there, one by MacKay and Knerer (1979) in 1968-1969 and another by Grixti and Packer (2006) in 2002-2003, 165 species were recorded, excluding honey bee, Apis mellifera Linnaeus (Apidae), and bumble bees, Bombus Latreille (Apidae), which were not sampled. Bee surveys taken between 1957 and 1984 at an abandoned field bordering forests comprised of oak, Quercus Linnaeus (Fagaceae) and hickory, Cary a Nuttall (Juglandaceae), (also within the eastern deciduous- Carolinian forest region) at the Edwin S. George Reserve in Livingston County, Michigan, United States of America, yielded a similar number of species (172) (Evans 1986). Given the longer species lists from these similar studies and factors related to sampling, it is likely that at least some additional species occur within or near our study areas, and still more 17 Onuferko et al. JESO Volume 146, 2015 within the greater Niagara Region. Exotic species ranged from well-established introductions such as A. mellifera and Megachile rotimdata (Fabricius) (Hymenoptera: Megachilidae) to more recent colonists (Table 1). The Palaearctic leafcutter bee Megachile ericetorum Lepeletier (Hymenoptera: Megachilidae), first discovered in the New World in St. Catharines in 2003 (Sheffield et al. 2010), is now well-established in the Niagara Region, based on subsequent captures of more than a dozen individuals in St. Catharines in 2006, 2010, 2012, and 2013 and in Port Colborne in 2012 and 2013; and recently in Rochester, New York, United States of America (Jacobi and Stafford 2012). We also collected two introduced Hylaeus species of the subgenus Spatulariella. Hylaeus hyalinatus Smith (Hymenoptera: Colletidae) was first reported in North America in 2001 (Ascher 2001) and then in St. Catharines almost every year from 2003 (Richards et al. 2011) to 2013, and was also found in Wainfleet in 2012. Hylaeuspunctatus (Brulle) (Hymenoptera: Colletidae) was first recorded in Canada in 2011 by Sheffield et al. (2011a), and was subsequently discovered in our St. Catharines samples from the same year. Anthidium manicatum (Linnaeus) (Hymenoptera: Megachilidae), which was found in almost every sampling year in St. Catharines, was also found in Pt. Colborne and Wainfleet. We also sampled a related introduced species, Anthidium oblongatum (Illiger) (Hymenoptera: Megachilidae), which is Palaearctic in origin and has been in Ontario since at least 2002 (Sheffield et al. 2011a). Exotic species established in eastern North America for some time include Lasioglossum leucozonium (Schrank) (Hymenoptera: Halictidae), L. zonulum (Smith) (Hymenoptera: Halictidae) (our only members of the subgenus Leuchalictus ) (Giles and Ascher 2006), Chelostoma rapunculi (Lepeletier) (Hymenoptera: Megachilidae) (Buck et al. 2005), and Megachile sculpturalis Smith (Hymenoptera: Megachilidae) (Paiero and Buck 2003). The leafcutter bee Megachile centuncularis (Linnaeus) (Hymenoptera: Megachilidae) has traditionally been considered to be a Holarctic species, though now there may be reason to suspect that it is exotic in North America as well (Giles and Ascher 2006; Sheffield et al. 2011b). Additional collections after 2003 of some of the abovementioned exotic species in the Niagara Peninsula may be indicative of their establishment within Ontario. Continued surveying within the present study region may be important in detecting future introductions as southern Ontario seems to have the highest number of introduced bee species in Canada (16 out of 17 exotic species in Canada (Sheffield et al. 2011b)), with one first record for North America of an Old World species discovered in St. Catharines. Our 10 years of collections represent the most extensive survey of the bee fauna in the Niagara Peninsula to date, and to our knowledge this is the longest continuous survey of any regional bee fauna in Canada. Although rare, transient, or extremely localized species may be discovered in the future, the current list likely encompasses the majority of common species present within the three sampled municipalities. To better detect the regional distribution patterns of bees, comprehensive sampling at additional localities is needed. Acknowledgements In addition to the taxonomic experts mentioned in the Methods who identified a large number of bees and verified many of our designations, we thank Rodrigo Leon Cordero, Jessi de Haan, and Konrad Karolak for help in preliminary taxonomic sorting of 18 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 specimens. This study was made possible with funding and other means of support from the Canadian Pollinator Initiative (CANPOLIN) strategic network, funded by the Natural Sciences and Engineering Research Council (NSERC). Lastly, we thank two anonymous reviewers and an associate editor for their suggestions to improve the manuscript. References Arduser, M. S. 2002. The bees of Missouri: A Preliminary report. Missouri Department of Conservation, Saint Louis, Missouri. Ascher, J. S. 2001. Hylaeus hyalinatus Smith, a European bee new to North America, with notes on other adventive. Proceedings of the Entomological Society of Washington 103: 184-190. Ascher, J. S., Kornbluth, S., and Goelet, R. G. 2014. Bees (Hymenoptera: Apoidea: Anthophila) of Gardiners Island, Suffolk County, New York. Northeastern Naturalist 21: 47-71. doi: 10.1656/045.021.0105 Ascher, J. S. and Pickering, J. 2015. Discover Life bee species guide and world checklist (Hymenoptera: Apoidea: Anthophila). [online] http://www.discoverlife.org/mp/ 20q?guide=Apoidea_species [accessed 9 October 2015] Bohart, G. E. and Nye, W. P. 1956. Bees. Foraging for Nectar and Pollen. Gleanings Bee Cult 84: 602-606. Bouseman, J. K. and LaBerge, W. E. 1978. A revision of the genus Andrena of the western hemisphere. Part IX. Subgenus Melandrena. Transactions of the American Entomological Society 104: 275-389. Buck, M., Paiero, S., and Marshall, S. A. 2005. New records of native and introduced aculeate Hymenoptera from Ontario, with keys to eastern Canadian species of Cerceris (Crabronidae) and eastern Nearctic species of Chelostoma (Megachilidae). Journal of the Entomological Society of Ontario 136: 37-52. Cane, J. H., Eickwort, G. C., Wesley, F. R., and Spielholz, J. 1983. Foraging, grooming and mate-seeking behaviors of Macropis nuda (Hymenoptera, Melittidae) and use of Lysimachia ciliata (Primulaceae) oils in larval provisions and cell linings. American Midland Naturalist 110: 257-264. doi: 10.2307/2425267 Cane, J. H., Minckley, R. L., and Kervin, L. J. 2000. Sampling bees (Hymenoptera: Apiformes) for pollinator community studies: pitfalls of pan-trapping. Journal of the Kansas Entomological Society 73: 225-231. Colla, S. R., Richardson, L., and Williams, P. 2011. Guide to the Bumble Bees of the Eastern United States. United States Department of Agriculture Forest Service and the Pollinator Partnership, [online] http://www.xerces.org/wp-content/ uploads/2008/09/Eastem_Bumble_Bee.pdf [accessed 9 October 2015] Eickwort, G. C. and Abrams, J. 1980. Parasitism of sweat bees in the genus Agapostemon by cuckoo bees in the genus Nomada (Hymenoptera: Halictidae, Anthophoridae). Pan-Pacific Entomologist 56: 144-152. Evans, F. C. 1986. Bee-flower interactions on an old field in southeastern Michigan. Pp. 103-109, In G. K. Clambey and R. H. Pemble, (eds). North American Prairie Conference, 1984. Center for Environmental Studies, Fargo, North Dakota/ 19 Onuferko et al. JESO Volume 146, 2015 Moorhead, Minnesota, United States of America. Gibbs, J. 2010. Revision of the metallic species of Lasioglossum (Dialictus) in Canada (Hymenoptera, Halictidae, Halictini). Zootaxa 2591: 1-382. Gibbs, J. 2011. Revision of the metallic Lasioglossum (Dialictus) of eastern North America (Hymenoptera: Halictidae: Halictini). Zootaxa 3073: 1-216. Gibbs, J., Dumesh, S., and Griswold, T. L. 2014. Bees of the genera Dufourea and Dieunomia of Michigan (Hymenoptera: Apoidea: Halictidae), with a key to the Dufourea of the eastern United States. Journal of Melittology 29: 1-15. doi: 17161/jom. v0i29.4652 Gibbs, J., Packer, L., Dumesh, S., and Danforth, B. N. 2013. Revision and reclassification of Lasioglossum ( Evylaeus ), L. (Hemihalictus ) and L. (Sphecodogastra ) in eastern North America (Hymenoptera: Apoidea: Halictidae). Zootaxa 3672(1): 1-117. doi: 10.11646/zootaxa.3672.1.1 Giles, V. and Ascher, J. S. 2006. Bees of the Black Rock Forest Preserve, New York (Hymenoptera: Apoidea). Journal of Hymenoptera Research 15: 208-231. Grixti, J.C. and Packer, L. 2006. Changes in the bee fauna (Hymenoptera: Apoidea) of an old field site in southern Ontario, revisited after 34 years. The Canadian Entomologist 138: 147-164. doi: 10.4039/n05-034 Hartman, C. G., Hartman, P, Hartman, P, and Rettenmeyer, C. 1944. Note on the habits of Osmia georgica Cresson as ascertained by the glass-tube method. Psyche: A Journal of Entomology 51, 162-165. doi: 10.1155/1944/53098 Hurd, P. D., Jr. 1979. Superfamily Apoidea. Pp. 1741-2209, In Krombein, K. V., Hurd, P. D., Jr., Smith, D. R., and Burks, B. D. (eds). Catalog of Hymenoptera of America North of Mexico, Vol. 2. Smithsonian Institution Press, Washington, District of Columbia, United States of America. Jacobi, B. and Stafford, G. 2012. Discovery of the western palearctic bee Megachile ( Pseudomegachile ) ericetorum (Hymenoptera: Megachilidae) in Rochester, N.Y., USA. bembiX 35: 5-8. LaBerge, W. E. 1973. A revision of the bees of the genus Andrena of the western hemisphere. Part VI. Subgenus Trachandrena. Transactions of the American Entomological Society 99: 235-371. LaBerge, W. E. 1980. A revision of the bees of the genus Andrena of the western hemisphere. Part X. Subgenus Andrena. Transactions of the American Entomological Society 106: 395-525. LaBerge, W. E. 1985. A revision of the bees of the genus Andrena of the western Hemisphere. Part XI. Minor subgenera and subgeneric key. Transactions of the American Entomological Society 111: 441-567. LaBerge, W. E. 1987. A revision of the bees of the genus Andrena of the western hemisphere. Part XII. Subgenera Leucandrena , P til andrena, Scoliandrena, and Mel andrena. Transactions of the American Entomological Society 112: 191-248. LaBerge, W. E. 1989. A revision of the bees of the genus Andrena of the western hemisphere. Part XIII. Subgenera Simandrena and Taeniandrena. Transactions of the American Entomological Society 115: 1-56. LaBerge, W. E. and Ribble, D. W. 1972. A revision of the bees of the genus Andrena of the western hemisphere. Part V. Gonandrena , Geissandrena , Parandrena , 20 A list of bee species recorded in the Niagara Region JESO Volume 146, 2015 Pelicandrena. Transactions of the American Entomological Society 98: 271-358. LaBerge, W. E. and Ribble, D. W. 1975. A revision of the bees of the genus Andrena of the western hemisphere. Part VII. Subgenus Euandrena. Transactions of the American Entomological Society 101: 371-446. MacKay, P. A. and Knerer, G. 1979. Seasonal occurrence and abundance in a community of wild bees from an old field habitat in southern Ontario. The Canadian Entomologist 111: 367-376. doi:10.4039/Entl 11367-3 McGinley, R. J. 1986. Studies of Halictinae (Apoidea: Halictidae), I: Revision of New World Lasioglossum Curtis. Smithsonian Contributions to Zoology 429: 1-294. doi: 10.5479/si.00810282.429 Michener, C. D. 2007. The Bees of the World, 2nd edition. The Johns Hopkins University Press, Baltimore. Maryland, United States of America. Mitchell, T. B. 1960. Bees of the eastern United States, Volume I. North Carolina Agricultural Experiment Station Technical Bulletin 141: 1-538. Mitchell, T. B. 1962. Bees of the eastern United States , Volume II. North Carolina Agricultural Experiment Station Technical Bulletin 152: 1-557. Osgood, E. A. 1989. Biology of Andrena crataegi Robertson (Hymenoptera: Andrenidae), a Communally Nesting Bee. Journal of the New York Entomological Society 97: 56-64. Packer, L., Genaro, J. A., and Sheffield, C. S. 2007. The bee genera of eastern Canada. Canadian Journal of Arthropod Identification 3: 1-32. doi: http://dx.doi. org/10.3752/cjai.2007.03 Paiero, S. M. and Buck, M. 2003. The giant resin bee, Megachile sculpturalis Smith, and other newly introduced and newly recorded native Megachilidae and Andrenidae (Apoidea) from Ontario. Journal of the Entomological Society of Ontario 134: 141-143. Rehan, S. M. and Richards, M. H. 2008. Morphological and DNA sequence delineation of two problematic species of Ceratina (Hymenoptera: Apidae) from Eastern Canada. Journal of the Entomological Society of Ontario 139: 59-67. Rehan, S.M. and Sheffield, C.S. 2011. Morphological and molecular delineation of a new species in the Ceratina dupla species-group (Hymenoptera: Apidae: Xylocopinae) of eastern North America. Zootaxa 2873: 35-50. Ribble, D. W. 1967. The monotypic North American subgenus Larandrena of Andrena (Hymenoptera: Apoidea). Bulletin of the University of Nebraska State Museum 6: 27-42. Ribble, D. W. 1974. A revision of the bees of the genus Andrena of the Western Hemisphere, subgenus Scaphandrena. Transactions of the American Entomological Society 100: 101-189. Richards, M. H., Rutgers-Kelly, A., Gibbs, J., Vickruck, J. L., Rehan, S. M., and Sheffield, C. S. 2011. Bee diversity in naturalizing patches of Carolinian grasslands in southern Ontario, Canada. The Canadian Entomologist 143: 279-299. doi: 10.4039/nll- 010 Rightmyer, M. G. 2008. A review of the cleptoparasitic bee genus Triepeolus (Hymenoptera: Apidae). Part I. Zootaxa 1710: 1-170. Roberts, R. B. 1973. Bees of the northwestern American: Agapostemon (Hymenoptera: 21 Onuferko et al. JESO Volume 146, 2015 Halictidae). Agricultural Experiment Station, Oregon State University Technical Bulletin 125: 1-23. Romankova, T. 2004. Ontario bees of tribe Epeolini: Epeolus Latreille and Triepeolus Robertson (Hymenoptera: Apidae, Nomadinae). Journal of the Entomological Society of Ontario 135: 87-99. Rutgers-Kelly, A. C. and Richards, M. H. 2013. Effect of meadow regeneration on bee (Hymenoptera: Apoidea) abundance and diversity in southern Ontario, Canada. The Canadian Entomologist 145: 655-667. doi: 10.4039/tce.2013.42 Sheffield, C. S., Dumesh, S., and Cheryomina, M. 2011a. Hylaenspunctatus (Hymenoptera: Colletidae), a bee species new to Canada, with notes on other non-native species. Journal of the Entomological Society of Ontario 142: 29—43. Sheffield, C. S., Frier, S. D., and Dumesh, S. 2014. The bees (Hymenoptera: Apoidea, Apiformes) of the Prairies Ecozone, with comparisons to other grasslands of Canada. Arthropods of Canadian Grasslands 4: 427-467. Sheffield C. S., Ratti, C., Packer, L., and Griswold, T. 2011b. Leafcutter and Mason Bees of the Genus Megachile Latreille (Hymenoptera: Megachilidae) in Canada and Alaska. Canadian Journal of Arthropod Identification 18: 1-107. Sheffield, C. S., Richards, M. and Griswold, T. 2010. Discovery of the Old World bee, Megachile ( Pseudomegachile ) ericetorum (Hymenoptera: Megachilidae), in Ontario, Canada. Journal of the Entomological Society of Ontario 141: 85-92. Stockhammer, K. A. 1967. Some Notes on the Biology of the Blue Sweat Bee, Lasioglossum coeruleum (Apoidea: Halictidae). Journal of the Kansas Entomological Society 40: 177-189. Taraday, B. 1982. The effect of the cleptoparasitic bee Nomada maculata (Hymenoptera: Anthophoridae) on the biology of its host Andrena vicina (Hymenoptera: Andrenidae). M.Sc. thesis, Cornell University, Ithaca, New York, United States of America. Wilson, J. S., Griswold, T., and Messinger, O. J. 2008. Sampling bee communities (Hymenoptera: Apiformes) in a desert landscape: are pan traps sufficient? Journal of the Kansas Entomological Society 81: 288-300. doi: 10.2317/JKES-802.06.1 Wolf, A. T. and Ascher, J. S. 2009. Bees of Wisconsin (Hymenoptera: Apoidea: Anthophila). Great Lakes Entomologist 41: 129-168. 22 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 REDESCRIPTION OF ANAPHES ATOMARIUS (BRETHES) (HYMENOPTERA: MYMARIDAE) AND COMPARISON WITH SIMILAR SPECIES IN EUROPE AND NORTH AMERICA J. T. HUBER Natural Resources Canada, c/o Canadian National Collection of Insects, Agriculture and Agri-food Canada, 960 Carling Avenue, Ottawa, Ontario, Canada, K1A0C6 email, John.Huber@agr.gc.ca Abstract J. ent. Soc. Ont. 146: 23-39 Anaphes atomarius (Brethes) (Hymenoptera: Mymaridae) is redescribed based on the holotype and specimens reared from Listronotns bonariensis (Kuschel) (Coleoptera: Curculionidae) in Brazil that are assumed tentatively to be conspecific with the type. Anaphes archettii Ghidini from Italy is also redescribed, a lectotype designated, and both species are compared to A. listronoti Huber and A. victus Huber from North America. Published November 2015 Introduction The Argentine stem weevil, Listronotns bonariensis (Kuschel) (Coleoptera: Curculionidae) is native to South America. It was accidentally introduced into New Zealand where it was discovered in 1927 (Dymock 1989) and has become a major economic pest (Timlin 1964). A search for potential biological control agents was begun by staff at the Commonwealth Institute of Biological Control, South American Station, San Carlos de Bariloche, Argentina, and an egg parasitoid was found and identified as Anaphes atomarius (Brethes) (Hymenoptera: Mymaridae). In 1966 and 1967, consignments of parasitized eggs were sent to New Zealand (Clausen 1977) and specimens were released at Nelson, Lincoln (Canterbury) and Waikato but the species failed to become established as a result of not being able to overwinter (Ferguson et al. 2007). Ahmad (1977, 1978) detailed the rearing technique for L. bonariensis and the egg parasitoid. Because L. bonariensis may occur as a contaminant in grain shipments from New Zealand or elsewhere it is listed as a quarantine pest of pasture grasses and cereals in the European Union (Ostoja-Starzewski 2011). The original 5-line Latin description and sketchy line drawings of wings and antenna are inadequate to define Anaphes atomarius and because of its potential for biological control the species is redescribed here, based on the holotype and several other specimens reared from L. bonariensis in Brazil. It is compared with similar species reared from known hosts in Europe and North America. 23 Huber JESO Volume 146, 2015 Methods Non-type specimens were slide mounted in Canada balsam using the method described by Noyes (1990). Photographs of slide preparations were taken with a ProgRes C14 plus digital camera attached to a Nikon Eclipse E800 compound microscope, and the resulting layers combined electronically using Auto-Montage® (Synoptics Group, Cambridge) or Zerene Stacker™ (http://zerenestacker.com) and, except for primary types, retouched as needed with Adobe® Photoshop (Adobe Systems for Windows). Measurements of morphological structures are given in micrometres (pm), following Huber (1992, 2006). Abbreviations used are: fl x = funicle or flagellar segment, mps = multiporous plate sensillum. Specimens are deposited in the following institutions. CNC - Canada, Ontario, Ottawa, Canadian National Collection of Insects. DEZA - Italy, Naples, Portici, Dipartimento di Entomologia e Zoologia Agraria delTUniversita degli Studi di Napoli «Frederico II». MACN - Argentina, Buenos Aires, Division Entomologia, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” Anaphes atomarius (Brethes) Anaphoidea atomaria Brethes, 1913: 100 (original description). Patasson atomarius: Ogloblin, 1964: 39 (generic transfer). Patasson atomarius : De Santis, 1967: 109 (catalogue). Patasson atomarius: Clausen, 1977: 272 (host, biological control). Patasson atomaria: De Santis, 1979: 371 (catalogue). Patasson atomarius: Ahmad, 1977: 151 (host, percent parasitism). Patasson atomarius: Ahmad, 1978: 161 (laboratory rearing, longevity). Patasson atomarium: Dymock, 1989: 23 (biological control). Anaphes atomarius: Huber, 1992: 72 (list, implied generic transfer). Type material. Holotype $ (MACN), on slide (Fig. 2) labelled: 1. “Patasson atomarius $ Brethes], Det. A. Ogloblin”. 2. “A 14”. 3. “Anaphoidea atomariaBr. 10545”. Some illegible letters in faded ink and the number 53 in pencil are also on the labels. Type locality: the original description gives the type locality and collecting date as General Urquiza and 1 .xi. 1912. The locality is now in Villa Urquiza, an area in greater Buenos Aires. Other Material Examined. BRAZIL. Rio Grande do Sul: Passo Fundo, 14.viii. 1985, D.N Gassen, ex. L. bonariensis (1$ and 4$, CNC). Diagnosis. Anaphes atomarius belongs to a complex of species with 2 mps on fl, of each antenna in females. The holotype differs from A. archettii (described below) and Anaphes listronoti Huber by the fore wing with double line of setae separating the medial space from the posterior margin of the wing (a single line in A. listronoti ), and narrower fore wing. 24 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 Incidentally, the line drawing of the fore wing of the holotype of A. listronoti (Huber et al. 1997, fig. 11) differs from its photograph (Huber 2006, fig. 23) in that the setal line between the medial space and posterior margin appears partly double in the former but single in the latter. I rechecked the holotype and the photograph showing a single setal line is correct. Anaphes atomarius differs from A. victus Huber by the narrower fore wing, with length to width ratio at least 8.0 (at most 6.7 in A. victus). The reared female from Brazil that I tentatively identify as A. atomarius has the fore wing with a single setal line separating medial space from hind margin and a slightly wider wing (length to width ratio of 7.37). Description. Female. Holotype (Fig. 1) body length 445 (mesosoma + metasoma only) (total length including head = 500 in original description). Head. Head width 189. Antenna. Length to width ratio of segments: scape + radicle 79/25 (3.16), pedicel 49/29 (1.69), fl T 21/14 (1.5), 11,57/19 (3.00), ff 57/21 (2.71), fl 4 57/22 (2.59), ff 56/21 (2.67), fl 6 52/22 (2.36), club 100/38 (2.63); ff-fl 6 each with 2 mps (Fig. 3). Wings. Fore wing (Fig. 4) length to width ratio 620/77 (8.05); longest marginal setae about 122, marginal space length 62, with double line of setae separating marginal space from hind margin (Fig. 4). Hind wing length 394, width 23, longest marginal setae about 109. Legs. Metatibia length 214, metatarsomere 1-4 lengths 32, 38, 34, 31; metatasomere 1 0.84 x length of metatarsomere 2. Metasoma. Ovipositor sheath length 277, extending under mesosoma to about level of anterior margin of mesocoxa (Fig. 5) and slightly exserted posteriorly (Fig. 6); ovipositor length to metatibia length ratio 1.29. Reared female specimen from Brazil. Body length 490 (mesosoma + metasoma only). Head. Head (Fig. 7) width 193. Antenna. Scape with faint oblique striations on inner surface (Figs 7, 8). Length to width ratio of antennal articles: scape + radicle 107/24 (4.46), pedicel 49/28 (1.75), fl, 26/16 (1.63), fl, 64/17 (3.76), ff 64/17 (3.76), fl 4 62/16 (3.88), ff 62/18 (3.44), fl 6 58/20 (2.90), club"l04/37 (2.81); fl,-fl 6 each with 2 mps (Fig. 8). Mesosoma. Scutellum (Fig. 9) with campaniform sensilla separated by 3.2 x their diameter. Wings. Fore wing length to width ratio 656/89 (7.37); longest marginal setae about 127, marginal space length 101, with single line of setae separating marginal space from hind margin. Hind wing length to width ratio (for a male specimen) 642/29. Legs. Metatibia length 208, metatarsomere 1-4 lengths 33, 39, 40, 35; metatasomere 1 0.85 x length of metatarsomere 2. Metasoma. Gaster (Fig. 10) about 0.9 x as long as mesosoma. Ovipositor length 294, extending under mesosoma to about level of anterior margin of mesocoxa (Fig. 11); ovipositor length to metatibia length ratio 1.41. Reared male specimens from Brazil. Body length (n=l, on slide) 645. Head as in Figs 12 and 13. Antenna. Length of segments (n=3) (Fig. 14): scape + radicle 91-97, pedicel 48, fl, 4-5, fl, 76-81, fl 3 85-86, fl 4 81-83, ff 80-82, fl 6 78-79, fl 7 76-82, fl g 76-78, fl 9 80- 84, fl, 0 76—80, fl,, 77-82. Length/width of ff. 3.75^4.04. Total flagellum length 797-815. Mesosoma. As in Fig 16. Wings. As in Fig 15. Metasoma. Gaster (Fig. 17) sligthly longer than high. Genitalia as in Fig. 18 (and see comments in Discussion). The four males are assumed to be conspecific with the reared female based on being obtained from the same rearing event. 25 Huber JESO Volume 146, 2015 FIGURES 1-2. Anaphoidea atomaria , holotype. 1, habitus; 2, type slide. Scale bar = 500 pm. 26 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 FIGURES 3-4. Anaphoidea atomaria , holotype. 3, antennae; 4, fore wing. Scale bars = 100 pm. 27 Huber JESO Volume 146, 2015 FIGURES 5-6. Anaphoidea atomaria , holotype. 5, mesosoma, lateral; 6, metasoma, lateral. Scale bars = 100 pm. 28 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 FIGURES 7-9. Anaphes 1 atomarius , reared female from Brazil. 7, head, anterior; 8, antenna; 9, mesosoma, dorsal. Scale bars = 100 pm. 29 Huber JESO Volume 146, 2015 FIGURES 10-11. Anaphes latomarius , reared female from Brazil, apex of mesosoma + metasoma; 10, dorsal surface; 11, ovipositor, dorsal view (as seen through metasoma). Scale bars = 100 pm. 30 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 FIGURES 12-15. Anaphes 1 atomarius, reared male from Brazil. 12, head, anterior; 13, head, dorsolateral; 14, antenna; 15, wings. Scale bars = 100 pm. 31 Huber JESO Volume 146, 2015 FIGURES 16-17. Anaphes latomarius , reared male from Brazil, lateral. 16, mesosoma; 17, metasoma. Scale bars = 100 jam. 32 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 FIGURE 18. Anaphes ? atomarius, reared male from Brazil, dorsal, genitalia. Scale bar = 100 pm. Anaphes archettii Ghidini Anaphes archettii Ghidini, 1945: 39 (original description). Anaphes archettii: Viggiani and Jesu, 1988: 1020 (host cited). Anaphes archettii'. Huber, 1992: 72 (list). Anaphes archettii: Viggiani, 1994: 472 (male genitalia). Anaphes archettii : Pagliano and Navone, 1995: 36 (list). Anaphes archettii: Jesu, 2002: 111 (host cited). Anaphes archettii: Pintureau, 2012: 33 (list). Type material. Lectotype $, here designated (DEZA), on slide (Fig. 20) labelled 1. Littoria, 13.v. 1943 ex Lixus junci coll. F.M Ghidini”. 2. “Lectotype $ des. Huber 2014”. 3. “ Anaphes archettii Ghidini 2$”. 4. “Paralectotype $ Anaphes archettii’. Type locality: Italy, Lazio, Agro Pontino [a plain in Latina Province south and southeast of the provincial capital, Latina). The former name of Latina was Littoria (used in the original description). Paralectotypes. 1$ and 1 $ (DEZA), with same data as lectotype. The female paralectotype is on the same slide as the lectotype, the male on another slide; both were examined. Three other specimens (DEZA) remain from the original series 33 Huber JESO Volume 146, 2015 but are in poor condition; they were not examined. All other original specimens are lost (G. Viggiani, personal communication). Diagnosis. Anaphes archettii belongs to the same species complex as A. atomarius. It differs from A. atomarius by four features: 1) longer body length (at least 770 long vs 500 in A. atomarius holotype), 2) fore wing with a single line of setae separating the medial space from the posterior margin (double line in atomarius holotype), 3) fore wing length to width ratio 5.39 (8.05 in atomarius holotype) and 4) ovipositor to metatibia length ratio 1.77 (1.29 in A. atomarius). The body length of A. archettii is at least 770, based on Ghidini (1945) compared to at most 693 in A. victus and 723 in A. listronoti. Description. Female. Lectotype (Fig. 19) body length (mesosoma + metasoma only) 792 (total length including head = 770-850 in original description). Head. Head width 314. Antenna. Length to width ratio of segments (scape-fh from paralectotype): scape + radicle 155/47, pedicel 65/38, f^ 36/19, fl 2 101/26, fl, 101/27, fl 4 99/30, fl. 91/29, fl 6 90/30, clava 145/45; fl 2 -fl 6 each with 2 mps (Figs 21, 22). Wings. Fore wing (Fig. 26 [male]) length to width ratio 1013/188 (5.39); longest marginal setae about 150, marginal space length 146, with single line of setae separating marginal space from hind margin. Hind wing length 904, width 58, longest marginal setae about 130. Legs. Metatibia length (paralectotype) 334, metatarsomere 1-4 lengths 61, 66, 58, 35; metatarsomere 1 0.92 x length of metatarsomere 2. Metasoma. Ovipositor sheath length 592, extending under mesosoma to about level of anterior margin of mesocoxa (Fig. 23); ovipositor length to metatibia length ratio 1.77. Male. Body length (from original description) 0.65-0.70 mm. Antenna. Length of segments (Fig. 24) (paralectotype): scape + radicle 128/40, pedicel 51/39, fl ] 9, fl, 125, fl 3 119, fl 4 118, fL 119/23, fl 6 116, fl 7 116, fl g 115, fl 9 111, fl ]() 109, fl n 114. Length to width ratio of fl. 5.04. Total flagellum length 1171. Fore wing as in Fig. 26. Genitalia as in Fig. 25 (and see comments in Discussion). Discussion Only four species of Anaphes have been described from South America: three in A. (Yungaburra ) and one, A. atomarius , in A. {Anaphes) (Huber 1992). Anaphes atomarius belongs to the crassicornis species group, in which the clava is 2-segmented. Among species described from the Western Hemisphere A. atomarius would key to couplet 12 in Huber (2006), which leads to A. listronoti, A. sordidatus (Girault) and A. victus. Anaphes victus and some specimens of A. listronotus Huber were reared from Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) among other species, and A. sordidatus was reared from Tyloderma foveolatum (LeConte) (Coleoptera: Curculionidae). Specimens of all three species sometimes or always have 2 mps on fl, of the female antenna, in contrast to other Anaphes species that have at most 1 or, usually, 0 mps on fl,. Several Old World (European) species also have 2 mps on fl 2 , but only one of them, A. archettii , is treated here for comparison with A. atomarius because the types were reared from a known host. The specimens from Passo Fundo, about 900 km from the type locality of A. atomarius, match the holotype fairly well but not perfectly. I tentatively treat the differences as intraspecific variation until shown otherwise by further rearing and morphological study 34 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 of additional specimens reared from L. bonariensis, preferably from nearer the type locality. Because of the slight morphological differences, the species name atomarius may not be correctly applied to the reared specimens I examined. Regarding specimens introduced into New Zealand, it is not known who made the species identification, whether voucher specimens from the releases or studies were kept or, if so, where they are deposited. Therefore their identity cannot be checked. Because no voucher specimens were located from previous FIGURES 19-22. Anaphes archettii, lectotype. 19, habitus; 20, type slide, 21, paralectotype antenna, from scape (radicle missing) to ff; 22, lectotype antenna, from ff (part) to clava. Scale bars: 19 = 1000 pm, 21 and 22 = 200~ pm. 35 Huber JESO Volume 146, 2015 FIGURES 23-26. Anaphes archettii, types. 23, female paralectotype, body dorsal; 24, male paralectotype, head + antenna; 25, male genitalia, lateral; 26, fore wing. Scale bars = 200 pm. 36 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 publications that use the name A. atomarius I cannot be sure whether the species name was correctly applied in those publications either. Like most species of Anaphes , the holotype of A. atomarius was not reared so its host is unknown. It would be expedient to assume that the name A. atomarius was correctly applied to all specimens reared from L. bonariensis because then the name would be associated with specimens reared from a known host that happens also to be a pest of economic importance. But this cannot be done until more evidence of conspecificity is obtained. That may be impossible because the holotype is slide mounted so other lines of evidence such as DNA barcoding or biological information cannot be obtained from it for comparison with freshly reared specimens from known hosts. The possibility exists that a complex of similar Anaphes species in South America uses L. bonariensis as a host, just as a complex of species exists on L. oregonensis in North America. Species in the latter complex differ in biology, e.g., in the number of individuals reared from a single host egg of L. oregonensis — A. listronotus is gregarious and A. victns is solitary (Huber et al. 1997). Unfortunately, publications on the biology of A. atomarius do not state how many adults emerge from a single host egg and this information was not recorded in the five reared specimens in this study. Another possibility is that A. atomarius is the same as one of the North American species. The fact that one species was described from Brazil and the others from Canada or the United States of America is not a problem because various species of Mymaridae in the Western Hemisphere are known to have wide distributions that extend from Canada, or at least somewhere north of Mexico, to Argentina. Additional rearing is needed of A. ‘atomarius ’ from Listronotus spp. in South America for detailed morphological study and, if colonies can be established, laboratory crossing experiments with the North American species, preferably with the addition of molecular evidence to see if species are the same or different. Ghidini (1945) reared numerous specimens of A. archettii from Lixiisjunci Boheman (Coleoptera: Curculionidae) on sugar beet (Beta vulgaris Linnaeus) (Chenopodiaceae) in Italy but did not state how many emerged from a single weevil egg. Apart from the specimens discussed above, the original material is lost (Viggiani, personal communication). Viggiani (1994) illustrated the male genitalia (photographed in Tig. 25) and showed that various Anaphes species could be distinguished by measurements of various genitalic parts. The problem is that association of males with females is only certain for the few Anaphes species reared from economically important hosts, whereas descriptions of most Anaphes species are based on females only, the corresponding males being unknown or not certainly associated. Because the genitalia of only three males of A. atomarius from Brazil and one of A. archettii are available for study little can be said about variation. In any case, there appears to be no difference in measurements. Body length in the four Anaphes species discussed above may be correlated with host egg size and number of individuals developing in a single egg. The gregarious or solitary nature of A. atomarius and A. archettii must first be determined, however. A host for each of the four species is known if one expediently, but perhaps incorrectly, assumes that specimens reared from L. bonariensis are indeed A. atomarius. If eggs of L. junci are larger than those of any of the Listronotus species that may account for the larger body size of A. archettii compared to the other Anaphes species. It would be interesting to obtain living A. archettii from L. junci and try to rear them on L. oregonensis in order to detennine whether the host range can be extended and, if so, see if specimens reared from L. oregonensis are 37 Huber JESO Volume 146, 2015 smaller than when reared on L.junci. If they are, then the body length difference proposed above to separated, archettii from 4. listronoti or A. victus does not distinguish these species and other differences need to be found. Ultimately, molecular evidence and cross breeding may be needed to distinguish correctly these (and other) Anaphes species. It may show that at least two of them are conspecific. Acknowledgements I thank J. Martinez (MACN) for the loan of the holotype of A. atomarius and G. Viggiani (DEZA) for the loan of three syntypes of A. archettii and information on the type locality. D. Ward, New Zealand Arthropod Collection and D. Gunawardana, Plant Health and Environment Laboratory, Ministry of Agriculture and Forestry, Auckland, searched for voucher specimens of A. atomarius but could not locate any. J. Read (CNC) is gratefully acknowledged for preparing the plate of illustrations. References Ahmad, R. 1977. Zur Kenntnis von Hyperodes bonariensis Kuschel (Col., Curculionidae) und seiner Feinde in Argentinien. Anzeiger fur Schadlingskunde, Pflanzenschutz, Umweltschutz 50: 150-151. Ahmad, R. 1978. Note on breeding the Argentine stem weevil Hyperodes bonariensis [Col. : Curculionidae] and its egg parasite Patasson atomarius [Hym. : Mymaridae], Entomophaga 23: 161-162. doi: 10.1007/BF02371722 Brethes, J. 1913. Himenopteros de la America meridional. Anales del Museo Nacional de Historia Natural de Buenos Aires 24: 1-165. Clausen, C. P. 1977. Curculionidae. In Clausen, C.P (ed.) Introduced parasites and predators of arthropod pests and weeds: a world review. United States Department of Agriculture, Agriculture Handbook 480. Agricultural Research Service, United States Department of Agriculture, Washington, DC, USA. Pp. 259-276. De Santis, L. 1967. Catalogo de los himenopteros argentinos de la serie Parasitica incluyendo Bethyloidea. Publicacion de la Comision de Investigacion Cientifica de Buenos Aires. Comision de Investigacion Cientifica, Provincia de Buenos Aires Gobernacion, La Plata, Argentina. De Santis, L. 1979. Catalogo de los himenopteros chalcidoideos de America al sur de los Estados Unidos. Memorias de la Comision de Investigaciones Cientificas de la Provincia de Buenos Aires. Publicacion especial. Comision de Investigacion Cientifica, Provincia de Buenos Aires Gobernacion, La Plata, Argentina. Dymock, J. J. 1989. Listronotus bonariensis (Kuschel), Argentine stem weevil (Coleoptera: Curculionidae). In Cameron, P. J., Hill, R. L., Bain, J., and Thomas, W. P. (eds) A review of biological control of invertebrate pests and weeds in New Zealand 1874 to 1987. CAB International Institute of Biological Control, Technical Communication 10. Wallingford, UK. Pp. 23-26. Ferguson, C. M., Moeed, A., Barratt, B. I. P, Hill, R. L., and Kean J. M. 2007. BCANZ - 38 Redescription of Anaphes atomarius (Brethes) JESO Volume 146, 2015 Biological Control Agents introduced to New Zealand [online], http://www.b3nz. org/bcanz [accessed December 2014], Ghidini, G. M. 1945. Osservazioni biologiche sul Lixus junci Boh. con descrizione di un suo nuovo parassita : Anaphes Archettii n. sp. Atti dell’Istituto Botanico della ‘Giovanni BriosV e Laboratorio Crittogrmico di Universita di Pavia, Serie V 6: 29—42 + Plate 3. Huber, J. T. 1992. The subgenera, species groups, and synonyms of Anaphes (Hymenoptera: Mymaridae) with a review of the described Nearctic species of the fuscipennis group of Anaphes s.s. and the described species of Anaphes (Yungaburra ). Proceedings of the Entomological Society of Ontario 123: 23-110. Huber, J. T. 2006 [2004], Review of the described Nearctic species of the crassicornis group of Anaphes s.s. (Hymenoptera: Mymaridae). Journal of the Entomological Society of Ontario 135: 3-86. Huber, J. T., Cote, S., and Boivin, G. 1997. Description of three new Anaphes species (Hymenoptera: Mymaridae), egg parasitoids of the carrot weevil, Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae), and redescription of Anaphes sordidatus Girault. The Canadian Entomologist 129: 959-977. doi: 10.4039/ Ent 129959-5 Jesu, R. 2002. Description of Anaphes maradonae n. sp. from Italy (Hymenoptera: Chalcidoidea: Mymaridae). Bollettino del Laboratorio di Entomologia Agraria «Filippo Silvestri» 58: 107-115. Noyes, J. S. 1990. Section 2.7.2.5. Chalcid parasitoids. Pp. 247-262 In Rosen, D. (ed.) The armoured scale insects, their biology, natural enemies and control, Vol. B. Elsevier Science Publishers, Amsterdam, The Netherlands. Ogloblin, A. 1964. Notas sobre algunas especies descritas por el Dr. Juan Brethes (Hym. Mymaridae). Neotropica 10: 39—40. Ostoja-Starzewski, J. 2011. Argentine stem weevil Listronotus bonariensis (Kuschel). The Food and Environmental Research Agency, Sand Hutton, York, United Kingdom. Pagliano, G. and Navone, P. 1995. Fascicolo 97. Hymenoptera Chalcidoidea. In Minelli, A. Ruffo, S., and La Posta, S. (eds), Checklist delle specie della fauna Italiana, 97. Calderini, Bologna, Italy. Pp. 1—40. Pintureau, B. 2012. Les Hymenopteresparasitoides oophages d Europe. Editions Quae c/o INRA, Versailles, France. Timlin, S. J. 1964. Diagnosis and extent of damage by stem weevil. Proceedings of the New Zealand Weed and Pest Control Conference 17: 149-151. Viggiani, G. 1994. L’armatura genitale esterna maschile di alcune species di Anaphes Haliday. Memorie della Societci Entomologica Italiana 72: 469-483. Viggiani, G. and Jesu, R. 1988. Considerazioni sui mimaridi italiani ed i loro ospiti. In Atti XV Congresso Nazionale Italiano di Entomologia, Accademia Nazionale Italiana di Entomologia, L’Aquila , Italy. Pp. 1019-1029. 39 Huber JESO Volume 146, 2015 40 Pheromone Races of Ostrinia nubilalis Infesting Grain Corn JESO Volume 146, 2015 PHEROMONE RACES OF OSTRINIA NUBILALIS HUBNER (LEPIDOPTERA: CRAMBIDAE) INFESTING GRAIN CORN IN MANITOBA, ONTARIO, ANDQUEBECPROVINCESOFCANADA J. L. SMITH 1 *, T. S. BAUTE 2 , C. E. MASON 3 department of Plant Agriculture, University of Guelph Ridgetown Campus Ridgetown, Ontario, Canada NOP 2C0 email, jocelyn.smith@uoguelph.ca Abstract J. ent. Soc. Ont. 146: 41-49 Ostrinia nubilalis (Hubner) (Lepidoptera: Crambidae), European corn borer, is an economic pest of Zea mays (Linnaeus) (Poaceae) and other vegetable crops that is distributed throughout the agricultural production regions of Ontario, Quebec, and Manitoba in Canada. Two phenotypic races of O. nubilalis have been identified that differ in the proportion of isomers of 11 - tetradecenyl acetate (ll-14:OAc) in their sex pheromone. The Z-race (Z-ll- 14:OAc) is the predominant race in the United States of America, known to inhabit Zea mays as its primary host, whereas the E-race (E-ll-14:OAc) infests a wider host range, including many vegetable crops, and is only found within the Eastern coastal states of the United States of America. Collections of O. nubilalis were made from grain corn in agricultural regions of Ontario, Quebec, and Manitoba in 1997, 2008, 2009, and 2010, and females were analyzed for pheromone race using gas chromatography (GC). Only Z-race O. nubilalis were found in Ontario (from Essex to Leeds and Grenville Counties) and in Southern Manitoba. E-race individuals were detected in collections from Ottawa, Ontario and St. Anicet, Quebec, with an increasing proportion of E-race phenotypes in samples from west to east. This is the first report of pheromone race determination using GC among Canadian O. nubilalis populations and the first documentation of E-race O. nubilalis in Canada using GC. Published December 2015 * Author to whom all correspondence should be addressed. 2 Ontario Ministry of Agriculture, Food and Rural Affairs Ridgetown, Ontario, Canada NOP 2C0 3 Department of Entomology and Wildlife Ecology University of Delaware, 531 S. College Avenue, Newark, Delaware, United States of America 19716-2160 41 Smith et al. JESO Volume 146, 2015 Introduction Ostrinia nubilalis (Hiibner) (Lepidoptera: Crambidae), European corn borer, has been an economic pest of corn, Zea mays (Linnaeus) Poaceae, throughout North America since introduction early in the 20 th century (Caffrey and Worthley 1927; Mason et al. 1996). Two phenotypic races of this species have been identified that differ in the proportion of 11- tetradecenyl acetate (ll-14:OAc) geometrical isomers in their sex pheromone composition (Klun and Brindley 1970). Although O. nubilalis is reported to utilize over 200 host plants, it predominantly infests corn, as its common name implies; however, the E-race (E-11-14: OAc) inhabits a wider host range, including peppers Capsicum spp. (Linnaeus) Solanaceae, potato Solanum tuberosum (Linnaeus) Solanaceae, tomato Solanum lycopersicum (Linnaeus) Solanaceae, and wheat Triticum aestivum (Linnaeus) Poaceae, as well as corn, whereas the Z-race (Z-ll-14:OAc) has a strong fidelity to corn (Bontemps et al. 2004; Mason et al. 1996). The Z-race is present throughout the North American range of O. nubilalis (Palmer et al. 1985); populations within the United States of America Corn belt are dominated by the Z-race (Mason et al. 1996; Showers et al. 1974), whereas the Northeastern coastal states contain greater proportions of the E-race (Klun and Brindley 1970; Mason et al. 1996; O’Rourke et al. 2010; Roelofs et al. 1972; Roelofs et al. 1985). Regional pheromone race identification of O. nubilalis is important for effective integrated pest management in agricultural crops including population monitoring using pheromone traps (DuRant et al. 1995) and for resistance management implications (Bontemps et al. 2004; O’Rourke et al. 2010). The major com producing areas in Canada are in southern portions of Manitoba, Ontario, and Quebec (Hamel and Dorff 2013); however, the pheromone race composition of O. nubilalis from these regions has not been reported. Klun et al. (1975) reported results for captures of O. nubilalis males in pheromone traps from several locations in Canada. They tested blends of the two pheromone isomers ranging from dominance of Z at one extreme to dominance of E in the lures at the other end of the spectrum. Their trapping data showed that more than 15% of males trapped at Simcoe, Ontario and St. Jean, Quebec were attracted to E pheromone blends out of a total catch of at least 45 moths in each case. At three other locations, moths predominantly were caught in traps baited with Z blends. Although pheromone trapping of males can provide an indication of presence of E and Z races in a population, this method is not definitive because males exhibit different levels of response to E and Z independently baited lures (Glover et al. 1991; Mason etal. 1997; Pelozuelo and Frerot 2007). McLeod etal. (1979) reported that male O. nubilalis collected from two Ontario locations and one population from St. Remi, Quebec responded most strongly to Z-ll-14:OAc using electroantennograms. However, another population that infested corn later in the same growing season from the Quebec location responded with greater affinity to E-l l-14:OAc. The most reliable method of race determination is analysis through gas chromatography of excised female pheromone glands or by analysis of race-specific single nucleotide polymorphism (SNP) genetic markers (Coates et al. 2013). Although grain corn is the second largest crop produced in Ontario by acreage, vegetable crops such as field tomatoes, sweet corn, and peppers also provide substantial farm value to the agricultural economy within the province (Hagerman 1997). The presence of significant acreages of fruit and vegetable crops in Essex, Chatham-Kent, and Niagara 42 Pheromone Races of Ostrinia nubilalis Infesting Grain Corn JESO Volume 146, 2015 Counties in Ontario, which have the potential to support E-race O. nubilalis , and reports of infestation of winter wheat T. aestivum (Linnaeus) Poaceae in Quebec and eastern Ontario (F. Meloche, personal communication) prompted the investigation of the composition of pheromone races in Canadian populations of O. nubilalis. Although Klun et at. (1975) and McLeod (1979) provided results for males collected in E- and Z-baited traps, the pheromone composition has not been documented with race-specific analysis through gas chromatography or SNP analysis. Although there is some hybridization in the field, E and Z populations are usually isolated due to multiple reproductive barriers (Dopman et al. 2010). The present study represents the first report of race-specific testing of Canadian O. nubilalis populations using gas chromatography; these results were generated prior to the development and publication of methods for SNP analysis (Coates et al. 2013). Populations of O. nubilalis were collected from commercial grain corn fields in Ontario, Quebec, and Manitoba in 1997, 2008, 2009, and 2010, and sent to C.E.M. at the University of Delaware for pheromone gland analysis of females using gas chromatography. Materials and Methods Insect Specimens O. nubilalis larvae were collected in September or October of each sampling year from commercial grain corn fields that had not been planted with transgenic hybrids that express Bacillus thuringiensis (Berliner) (Bt) Bacillales insecticidal proteins (fir-corn) or from non-fir refuge plants within fir-corn fields (Table 1, Fig. 1). In 1997, 50 field-collected larvae from each location were cooled and directly shipped, in cardboard larval rearing rings with artificial diet, to C.E.M. for pheromone analysis. In 2008, 2009, and 2010, com stalks containing diapausing larvae were removed from growers’ fields and kept over winter in a non-heated bam at the University of Guelph Ridgetown Campus (Ridgetown, Ontario). Following termination of diapause, larvae were extracted from the corn stalks and transferred into rearing dishes with cardboard pupation rings, which were placed in growth chambers maintained at 16:8 L:D, 27 °C photoperiod, 18 °C scotoperiod, and 75 % relative humidity (RH) to establish laboratory colonies; original colony sizes ranged from 20-70 individuals. After multiple generations of laboratory rearing (Table 1), pupae were removed from the colony, sexed, and female pupae were shipped to C.E.M. for pheromone analysis. Upon receipt by C.E.M., individual larvae and/or pupae were housed in 28 ml plastic food service cups containing cotton rolls saturated with water, and these were placed in a growth chamber set on a reversed photoperiod to facilitate gland removal at regular working hours. Through pupation and eclosion, conditions were set at 25 °C, 16:8 (L:D) photoperiod, and 50-80 % RH. Drinking water was provided for newly emerged moths, and females were set aside for pheromone analysis. Pheromone ring glands of females were excised with micro-scissors at the non- sclerotized terminal segment, just anterior of the single ring gland, during the 7 th h of scotophase the second day after eclosion (2U48 h old). Each gland was placed into a 50-pl point-tipped auto-sampler vial containing 5 pi of heptane and an internal standard of 4.5 ng cis-7-tetradecenyl acetate (Z-7- 14:OAc). Samples were held for > 30 min at room temperature or stored in a - 20 °C freezer before analysis. 43 TABLE 1. Location of Canadian field collections of Ostrinia mibilalis in 1997, 2008, 2009, and 2010, and the percentage of E and Z pheromone races or hybrids determined using gas chromatography (GC). Smith et cd. JESO Volume 146, 2015 CD OD cd S3 CD o S-H CD Ph PP X N W Id .22 c/s c/s =3 >? o 13 o a cd S u go o- s . a pp o p—> co o O CD c/s CD ad o o o o o • *-H PP Oh cO ^H OQ O CD O CD tp =3 h-> ■ ^H OD O o HP CD tp o cd v CD S-H cd CD z cO 4* cd ;_0 a £>o o -o o oooooooo o' o' o' o' o' o' o' o' r i oooooooo o' o' o' o' o' o' o' o' oooooooo !"■ OOOOOOOO-H o' o' o' o' o' o' o' o' A oo c- m —< . (N co cs c i co c- c- — Ot^t^t^^-Ti-Td-Td-co O Cl Cl O 0- O O oo Cl ci ci ci ci co’ ci ci in ^|- ^oi- ^|- -^i- -^i- ^t- S3 S3 CD v ^ 5 2 Sh "Td S-h S-h O rrt aBV, § doo § 2 hChP^C^hPPQ^IoI^O CD S3 SP CD CD XX * BBv ad cd c3 S3 o cd 2 o-S "go cd cd S i S_h S-h cd <33 cd v > 3 W8g XggPo O O o o' o o' ICS C- Ph O O O Cl C- co c- o 'Tl- pp CD CD S-H u w S3 o cd PP O H-* • i—H S3 cd r- o' o co co’ o o' Cl ICS >cs Cl Ph OO o o Cl n CD co lo § 3 a ^ CD ^ OO QJ O a cf O S-H v CD 3 N § % 22 -2 cr ’B S3 CD W 2 'g UU $ ^ c3 w cp P S3 33 c/s O O S-H