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.

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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.

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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:

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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

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Great Lakes Entomologist 41: 129-168.

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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.

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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.

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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.

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FIGURES 1-2. Anaphoidea atomaria , holotype. 1, habitus; 2, type slide. Scale bar = 500

pm.

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FIGURES 3-4. Anaphoidea atomaria , holotype. 3, antennae; 4, fore wing. Scale bars =

100 pm.

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FIGURES 5-6. Anaphoidea atomaria , holotype. 5, mesosoma, lateral; 6, metasoma, lateral.

Scale bars = 100 pm.

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FIGURES 7-9. Anaphes 1 atomarius , reared female from Brazil. 7, head, anterior; 8,

antenna; 9, mesosoma, dorsal. Scale bars = 100 pm.

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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.

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FIGURES 12-15. Anaphes 1 atomarius, reared male from Brazil. 12, head, anterior; 13,

head, dorsolateral; 14, antenna; 15, wings. Scale bars = 100 pm.

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FIGURES 16-17. Anaphes latomarius , reared male from Brazil, lateral. 16, mesosoma; 17,

metasoma. Scale bars = 100 jam.

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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

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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

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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.

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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.

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Redescription of Anaphes atomarius (Brethes)

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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

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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

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De Santis, L. 1967. Catalogo de los himenopteros argentinos de la serie Parasitica

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Dymock, J. J. 1989. Listronotus bonariensis (Kuschel), Argentine stem weevil (Coleoptera:

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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

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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).

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Pheromone Races of Ostrinia nubilalis Infesting Grain Corn JESO Volume 146, 2015

Gas Chromatography

Pheromone extractions were analyzed with a Varian 3500 gas chromatograph

equipped with a Varian 8200 auto-sampler (Agilent Technologies, Santa Clara, California,

United States of America) using capillary techniques similar to those described by Field et al.

(1999) and DuRant et al. (1995). Three pi of solution from the sample were injected in the gas

chromatograph injector using a sandwich technique where a 0.5 pi upper air gap was placed

between the solvent plug and sample plug in a 10-pl syringe. The air gap resides between

the sample and the solvent that remains below the syringe plunger after rinsing the syringe.

The air gap prevents liquid-to-liquid contact and reduces the chance of sample contamination

from previous samples. A 0.8 pi lower air gap was used to reduce sample volatilization during

insertion of the needle into the hot injector.

During injection, the hot needle time was zero, the injection rate was 1.5 pi sec 1 ,

and the needle residence time was 0.02 min. The gas chromatograph was equipped with a

heated injector fitted with a 4 mm inside diameter open-top glass uniliner (Restek Corporation,

Bellefonte, Pennsylvania, United States of America) containing glass wool, a fused silica

capillary column (15 m x 0.25 mm) with 0.25 pm Stabilwax® film thickness (Restek

Corporation, Bellefonte, Pennsylvania, United States of America), a 5 m x 0.25 mm fused

silica guard column, and a flame ionization detector. The gas chromatograph was programmed

as follows: injector temp: 200 °C, splitless for 1.5 min, then set to split for the remainder of the

run (split ratio 50:1 set at 60 °C); detector temp: 250 °C, attenuation set at 32 xlO' 11 ; column

oven programmed at 80 °C, held 2.0 min, heat from 80 °C to 240 °C at 10 °C min' 1 , held at 240

°C for 5 min to end of the run; and total run time was 23 min. Hydrogen was used as carrier

gas at a flow rate of 20 cm sec -1 (6.5 psi head pressure) and nitrogen was used as makeup gas.

Under these conditions, the Z-7-14:OAc internal standard and the two pheromone isomers

eluted at « 13.1 - 13.5 min with each of the three peaks being separated by 0.2 to 0.4 min,

which allowed for distinct separation on the chromatogram and detection of these compounds

(Fig. 2).

FIGURE 1. Focations where Ostrinia nubilalis tested for pheromone race analysis were

collected in 1997, 2008, 2009, and 2010 in Canada.

45

Smith et al.

JESO Volume 146, 2015

Chromatogram results for female moths for which pheromone glands were excised and

analyzed by gas chromatography were categorized by pheromone race based on the percentage

ratio of the two pheromone isomers. The percentages were detennined by comparison of peak

heights of the isomers at the appropriate retention times on the chromatogram. Samples with

the peak height consisting of 95 % or more of the E isomer compared to the Z isomer were

classified as E-race, those with 5 % or less of the E isomer were classified as Z-race, and those

with intermediate percentages of E isomer were classified as hybrids, whereby the percentages

with very few exceptions fell within the 20 % to 80 % range. Although these criteria have a

broad range, analyses by C.E.M. of approximately 1000 O. nubilalis showed that E and Z

phenotypes do not fall outside the 5 % range for the minor isomer of each phenotype (Coates et

al. 2013). Allelic variation in a fatty-acyl reductase gene essential for pheromone biosynthesis

accounts for the phenotypic variation in female pheromone production (Lassance et al. 2010).

Mean percentages and standard deviations for the E isomer are typically about 98.5 ± 0.5 %

for the E race, 67 ± 10 % for the hybrid, and 3 ± 1.0 % for the Z race.

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FIGURE 2. Representative chromatograms from pheromone analyses of Ostrinia nubilalis

in Canada showing peaks for internal standard (IS), Z-ll-14:Oac (Z), and E-ll-14:Oac

(E).

Results and Discussion

In all years of sampling, only Z-race O. nubilalis were found in collections from

grain corn in Essex, Chatham-Kent, Middlesex, Huron, Niagara, Leeds and Grenville

Counties in Ontario, and from Grey County in Manitoba (Table 1, Fig. 1). Both E- and Z-

race phenotypes were identified in samples collected in grain corn from Ottawa, Ontario

and Saint-Anicet, Quebec (Table 1). The Ottawa population collected in 2008 contained

21.4 % hybrid females where significant quantities of both E and Z isomers were measured

by gas chromatography; however, the majority of individuals were Z-race (71.4 %) and 7.1

% were E-race (Table 1). Of the individuals tested from Quebec, the majority tested were E

46

Pheromone Races of Ostrinia nubilalis Infesting Grain Corn JESO Volume 146, 2015

and Z hybrids (66.7 %), and there was a greater proportion of E-race insects (20.0 %) than

Z-race (13.3 %) (Table 1).

Our results show that the Z-race of O. nubilalis is the dominant pheromone

race found infesting grain corn in the major corn producing regions of Canada, present

throughout Ontario from Essex to Ottawa Counties and in southern Manitoba. E-race O.

nubilalis were only present in colonies derived from collections from grain corn in eastern

Ontario near Ottawa and Quebec, which indicates the presence of E-race within these

regions. The proportion of Z or E races among founder males and females from original

field collections and the resulting frequency of hybridization among offspring prior to GC

analysis are unknown. Detection of E-ll-14:OAc isomers in our analysis is evidence that

E-race O. nubilalis were originally present in the area sampled. However, where no E-11-

14:OAc isomers were detected there is a degree of uncertainty as to whether the E-race was

lost through generations of rearing in the laboratory or because the small number of founder

individuals in some collections may not have been sufficient to detect E-race individuals.

A relatively new method of distinguishing pheromone races of O. nubilalis using SNP

markers enables high throughput processing of larger sample sizes and has greater than

98 % correlation with results from GC analysis (Coates et al. 2013; Lassance et al. 2010).

Therefore we are confident in our results indicating E- and Z-race phenotypes.

Klun et al. (1975) reported some males collected in E-race pheromone traps at

Chatham and Simcoe, Ontario and St. Jean, Quebec. It is possible that some E-race O.

nubilalis were present in these areas in 1973 and 1974 when Klun etal. (1975) conducted their

study. Our results indicate it is unlikely that the E-race is currently present in southwestern

Ontario. Since we found the E-race phenotype present in the eastern range of populations

we studied, it is likely that the E-race is present further east from St. Anicet, Quebec and

Ottawa, Ontario. Although we did not show the presence of the E-race phenotype at the

Kemptville collection site, the E-race may be present there now since the samples we

analyzed were from a decade earlier in the 1997 collection; however, testing of current

populations must be completed for confirmation. A more in-depth study of populations of O.

nubilalis collected from a wider host range within the regions studied would provide more

conclusive information on the pheromone race composition of O. nubilalis in Canada. This

is the first documented evidence of the E-race in Canada corresponding with the eastern corn

growing region, which is a similar distribution pattern as in the U.S. These results provide

useful information for pheromone trap monitoring of O. nubilalis in Eastern Ontario and

Quebec, and support observations of infestations in non-corn crops. O. nubilalis is also a

significant pest of potato in Quebec, New Brunswick, and Prince Edward Island (Noronha

et al. 2008). Consequently, the E-race of O. nubilalis very well could be prominent in these

areas east of where we conducted our study. Further analysis of populations collected from

these regions is needed to determine this.

Acknowledgements

The authors wish to thank John Gavloski from Manitoba Agriculture, Food and

Rural Initiatives, Carman, Manitoba and Francois Meloche (retired) from Agriculture

and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Ottawa, Ontario for

47

Smith et al.

JESO Volume 146, 2015

sending collections of O. nubilalis from Manitoba, and Ottawa and Quebec, respectively. We

also wish to acknowledge Jennifer Bruggeman and Emily Burggraaf for their insect rearing

efforts. We are grateful for the excellent anonymous review comments and suggestions.

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Klun, J. A. and Brindley, T. A. 1970. cis-ll-Tetradecenyl Acetate, a Sex Stimulant of the

European Corn Borer. Journal of Economic Entomology 63: 779-780. doi: 10.1093/

jee/63.3.779

Klun, J. A. 1975. Insect Sex Pheromones: Intraspecific Pheromonal Variability of Ostrinia

nubilalis in North America and Europe. Environmental Entomology 4: 891-894.

doi: 10.1093/ee/4.6.891

48

Pheromone Races of Ostrinia nubilalis Infesting Grain Corn JESO Volume 146, 2015

Lassance, J.-M., Groot, A. T., Lienard, M. A., Antony, B., Borgwardt, C., Andersson, F.,

Hedenstrom, E., Heckel, D. G., and Lofstedt, C. 2010. Allelic variation in a fatty-

acyl reductase gene causes divergence in moth sex pheromones. Nature 466 : 486-

489. doi:10.1038/nature09058

Mason, C. E., Eheresman, N. R, He, K., Ilalia, A. D., and Pesek, J. D. 1997. Performance

of three commercial pheromone sources for trapping European corn borer. In

Ingerson-Mahar, J., (ed). 31st Northeast Regional Field Crops Insect Conference

at Burlington, VT, Rutgers University, New Brunswick, NJ. Pp. 27-34.

Mason, C. E., Rice, M. E., Calvin, D. D., Van Duyn, J. W., Showers, W. B., Hutchison,

W. D., Witkowski, J. F., Higgins, R. A., Onstad, D. W., and Dively, G. P. 1996.

European Com Borer Ecology and Management. North Central Regional Extension

Publication, Ames, Iowa.

McLeod, D. G. R., Ritchot, C., and Nagai, T. 1979. Occurrence of a two generation strain of

the European corn borer, Ostrinia nubilalis (Lepidoptera: Pyralidae), in Quebec.

The Canadian Entomologist 111 : 233-236. doi: 10.4039/Entl 11233-3

Noronha, C., Vernon, R. S.,and Vincent, C. 2008. Key pests of potatoes in Canada. Cahiers

Agricultures 17 : 375-381.

O’Rourke, M. E., Sappington, T. W., and Fleischer, S. J. 2010. Managing resistance to

Bt crops in a genetically variable insect herbivore, Ostrinia nubilalis. Ecological

Applications 20 : 1228-1236. doi: 10.1890/09-0067.1

Palmer, D. F., Schenk, T. C., and Chiang, H. C. 1985. Dispersal and Voltinism Adaptation of

the European Corn Borer in North America, 1917-1977. University of Minnesota,

Minneapolis, MN, USA. Pp. 31.

Laurent, P. and Frerot, B. 2007. Monitoring of European Com Borer with Pheromone-Baited

Traps: Review of Trapping System Basics and Remaining Problems. Journal of

Economic Entomology 100 : 1797-1807. doi:10.1603/0022-0493(2007)100[1797:

MOECBW]2.0.CO;2

Roelofs, W. L., Carde, R. T., Bartell, R. J., and Tierney, P. G. 1972. Sex Attractant Trapping

of the European Corn Borer in New York. Environmental Entomology 1 : 606-608.

doi: 10.1093/ee/l .5.606

Roelofs, W. L., Du, J. W., Tang, X. H., Robbins, PS., and Eckenrode, C. J. 1985. Three

European corn borer populations in New York based on sex pheromones and

voltinism. Journal of Chemical Ecology 11 : 829-836. doi: 10.1007/bf01012071

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of Males to Synthetic Lure and to Females of Different Strains. Environmental

Entomology 3 : 51-58. doi: 10.1093/ee/3.1.51

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50

The historical spread of Hermetia illucens (L.) in Canada

JESO Volume 146, 2015

THE HISTORICAL SPREAD OF THE BLACK SOLDIER FLY,

HERMETIA ILLUCENS (L.) (DIPTERA, STRATIOMYIDAE,

HERMETIINAE), AND ITS ESTABLISHMENT IN CANADA

S. A. MARSHALL 1 *, N. E. WOODLEY 2 , M. HAUSER 3

1 School of Environmental Sciences, University of Guelph,

University of Guelph, Guelph, Ontario, Canada NIG 2W1

email, samarsha@uoguelph.ca

Scientific Note J. ent. Soc. Ont. 146: 51-54

The Black Soldier Fly, Hermetia illucens (Linnaeus), is a synanthropic,

polysaprophagous fly native to the Neotropics, but now found in every zoogeographic

region following decades of spread throughout the warmer parts of the world. In the last

few decades there has been considerable interest in using larvae of H. illucens for organic

waste control, composting, and as animal food supplements. It has been studied as an

agent for manure control (Sheppard et al. 1994), for controlling house fly infestations in

chicken production (Furman et al. 1959, Sheppard 1983), and as a food supplement for fish

(Bondari and Sheppard 1981) and swine (Newton et al. 1977). More recently, there has

been considerable interest in using H. illucens as an agent for composting (Lalander et al.

2015). An internet search on “Black Soldier Fly” demonstrates that this use is reaching a

household level.

Although it has been suggested that H. illucens might have been first brought to

Europe around 500 years ago (Benelli et al. 2014), the first verifiable Palaearctic record of

the species is from southern Europe (Malta) in 1926 (Lindner 1936). The subsequent spread

of this large, easily recognized species in Europe has been mainly along the Mediterranean

coast of Spain, France, and Italy in the 1950s and 1960s (Leclercq 1969, 1997). In more

recent years, the species has been documented spreading northwards in Central Europe:

Ssymank and Doczkal (2010) recorded it from Gennany and Rohacek and Hora (2013)

recorded it from the Czech Republic. We have seen specimens from South Africa collected

as early as 1915 and from Malaysia, Hawaii, Solomon Islands, New Caledonia, Mariana

Islands, Palau, and Guam in the 1940s. Probably by the 1960s, H. illucens had spread over

most of the range it occupies today. The apparent spread of this species along coastlines

and islands suggests that maritime transport may have played a role in repeated accidental

introductions.

Published December 2015

* Author to whom all correspondence should be addressed.

2 Systematic Entomology Laboratory -Agricultural Research Service, United States

Department of Agriculture, Smithsonian Institution NHB-168

P. O. Box 37012, Washington, DC, United States of America 20013-7012

3 California Department of Food and Agriculture, Plant Pest Diagnostics Branch, 3294

Meadowview Road, Sacramento, California, United States of America 95832-1448

51

Marshall et al.

JESO Volume 146, 2015

Although we do not know the exact original distribution of the Black Soldier Fly

and it cannot be excluded that it originally occurred in the southeastern United States, its

current North American range seems to reflect a northward spread from a native range in

Central America and the northern parts of South America in historical times. It was present

in the southern United States by the late 1800s. The earliest specimen we are aware of is

from 1881 from Fernandina, Florida (United States National Museum). Riley and Howard

(1889) recorded it (misspelled as Hermetia mucens ) on the basis of a specimen collected

from beehives in Alabama in 1887. We know of further records from 1897 (Fouisiana), 1899

(Texas), 1911 (South Carolina), 1923 (southern California), 1926 (Virginia), 1931 (Iowa),

1938 (Ohio), 1940 (northern California), 1943 (Maryland), and 1945 (New York City). The

northernmost record we are aware of is from Warner, Merrimack County, New Hampshire,

16 September 1972 (University of New Hampshire Collection). The species was not listed

in works on the Diptera of Kansas (Adams, 1903) or Oregon (Cole and Fovett 1921). James

(1960) shows a distribution map for California, in which the most northern specimen is at

the northern end of the Central Valley. However, he also gives a North American map and

mentions records from Oregon, Washington state, and North Dakota, which he discusses as

“temporary local introductions”. Woodley (2001) did not include these records in his World

catalog. By now, there are many records available from Oregon and Washington State, and

we consider the species established in these states. One of us (NEW) did a considerable

amount of collecting in southeastern Washington from about 1972 to 1980, and H. illucens

was never seen. It now occurs there, indicating its spread in that area has been relatively

recent. We know of no verified records from western Canada, the Rockies, and most of the

Great Plains states.

Hermetia illucens is not currently recorded as occurring naturally in Canada,

which is a significant issue because there is a great deal of interest in utilizing this species in

Canada for waste processing, compost production, and protein production. Because Black

Soldier Flies are easily reared on a wide range of decomposing materials, including animal

and human waste, the large, slow-moving larvae of these flies have become popular both

for manure management and pet food, and are now widely mass-reared for a variety of

commercial uses in the United States, Costa Rica, Europe, and South Africa. At the moment,

this species can only be imported into Canada as sterilized larvae shipped without any manure

or sewage sludge (B. Gill, Canadian Food Inspection Agency, personal communication).

Therefore, the question of whether or not Hermetia illucens occurs naturally in Canada is

an important one.

There are no verifiable published Canadian records of the Black Soldier Fly,

although the species is erroneously recorded from Edmonton, Alberta in the CANADENSYS

database (Biodiversity Institute of Ontario, 2015). This entry is apparently an error due

to contamination; there are no Hermetia specimens from Alberta in the Biodiversity

Institute of Ontario collections (J. deWaard, Biodiversity Institute of Ontario, personal

communication).

Although the Edmonton records are in error, we here record H. illucens from Canada

for the first time on the basis of specimens collected in southern Ontario and deposited in

the University of Guelph insect collection as early as 2007 (Windsor, Black Oak Savannah

Park, 20 June 2007, S.M. Paiero, DEBU 002268353). This record is not surprising given

the many other new Canadian records from the same area (Paiero et al. 2010) and given the

52

The historical spread of Hermetia illucens (L.) in Canada

JESO Volume 146, 2015

known occurrence of H. illucens in nearby Michigan. Further Black Soldier Fly specimens

were documented in the course of a forensic investigation near Aurora, Ontario in August

2011 (S. VanFaerhoven, personal communication). So we can here confidently record H.

illucens as collected repeatedly in Ontario, and it is probably established here.

Several questions remain unanswered about the history and current status of the

Black Soldier Fly in Canada. The records reported here might reflect natural northward

movement of this synanthropic fly, but it is more likely that the captured flies were escaped

adults that emerged from imported larvae. These adults would be sterile if they hatched from

legally imported sterile larvae, but is also possible that this species has been established in

the province as a result of illegal importation and rearing of viable populations. Trinh et

al. (2013), for example, indicate that their viable cultures of Black Soldier Flies originated

with a supplier in Georgetown, Ontario, and it is likely that other viable cultures have been

brought across the border by well-intentioned organic gardeners, fishermen, or pet owners.

We suspect that the Black Soldier Fly is here to stay, but the data available to us do not allow

us to say how and when it arrived.

Acknowledgements

We thank Christopher C. Grinter, Richard S. Zack, and Robert Zuparko for checking

specimens in their collections, Sherah VanFaerhoven for sharing her unpublished records,

Bruce Gill for comments on an early draft of this note, Robert Walberg for encouraging us to

clarify the status of Black Soldier Fly in Ontario, and Steven Paiero for depositing his first

Ontario collections of Black Soldier Fly in the University of Guelph Insect Collection.

References

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JESO Volume 146, 2015

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southern Ontario tallgrass prairie. Pp. 199-225. In Shorthouse, J., and Lloate, K. D.

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54

JESO Volume 146, 2015

THE ENTOMOLOGICAL SOCIETY OF ONTARIO

OFFICERS AND GOVERNORS

2015-2016

President: J. GIBSON

Royal BC Museum

675 Belleville Street, Victoria, BC V8W 9W2

J G ibson@roy albcmuseum. be. ca

President-Elect: G. OTIS

School of Environmental Sciences

University of Guelph, Guelph, ON NIG 2W1

goti s@uoguelph. ca

Past President: L SCOTT

Agriculture and Agri-Food Canada,

1391 Sandford St. London, ON N5V 4T3

ian.scott@agr.gc.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

Shiyou.li@canada.ca

Directors*

D. BERESFORD (2015-2017)

Biology Department, Trent University

1600 West Bank Drive, Peterborough, ON K9J 7B8

davidberesford@trentu.ca

A. GUIDOTTI (2014-2016)

Department of Natural flistory. 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.com

A. SMITH (2016-2018)

Department of Integrative Biology

University of Guelph, 50 Stone Road East

Guelph, ONN1G2W1

salex@uoguelph.ca

J. SMITH (2015-2017)

University of Guelph Ridgetown Campus

120 Main St. E, Ridgetown, ON NOP 2C0

jocelyn.smith@uoguelph.ca

L. TIMMS (2016-2018)

laura.le.timms@gmail.com

ESO Regional Rep to ESC: S. CARDINAL

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K. SHUKLA

Student Representative: C. PEET-PARE

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JESO Editor: C. MACQUARRIE

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Technical Editor: T. ONUFERKO

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onuferko@yorku. ca

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ENTOMOLOGICAL SOCIETY OF ONTARIO

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and among the nine oldest, existing entomological societies in the world. It serves as an

association of persons interested in entomology and is dedicated to the furtherance of

the science by holding meetings and publication of the Journal of the Entomological

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An annual fee of $30 provides membership in the Society and a subscription to the

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Please send your name, address (including postal code) and email address to:

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or email: entsocont.membership@gmail.com

NOTICE TO CONTRIBUTORS

Please see the Society web site (http://www.entsocont.ca) for current instructions to

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FELLOWS OF THE ENTOMOLOGICAL SOCIETY OF ONTARIO

W. W. BILL JUDD

2002

FREEMAN MCEWEN

2010

C. RON HARRIS

2003

THELMA FINLAYSON

2013

GLENN WIGGINS

2006

JOHN STEELE

2014

BERNARD PHILOGENE

2010

CONTENTS

I. FROM THE EDITOR.1-2

II. ARTICLES

T. M. ONUFERKO, R. KUTBY, and M. H. RICHARDS — A list of bee species (Hymenoptera:

Apoidea) recorded from three municipalities in the Niagara region of Ontario, including a new

record of Lasioglosum furunculum Gibbs (Halictidae) in Canada.3-22

J. T. HUBER — Redescription of Anaphes atomarius (Brethes) (Hymenoptera: Mymaridae)

and comparison with similar species in Europe and North America.23-39

J. L. SMITH, T. S. BAUTE, and C. E. MASON — Pheromone races of Ostrinia nubilalis

Hiibner (Lepidoptera: Crambidae) infesting grain corn in Manitoba, Ontario, and Quebec

provinces of Canada.41-49

S. A. MARSHALL, N. E. WOODLEY, and M. HAUSER — . 2015. The historical spread of

the Black Soldier Fly, Hermetia illucens (L.) (Diptera, Stratiomyidae, Hermetiinae), and its

establishment in Canada.51-54

III. ESO OFFICERS AND GOVERNORS 2015-2016.55

IV. ESO OFFICERS AND GOVERNORS 2014-2015.inside front cover

V. FELLOWS OF THE ESO.inside back cover

VI. APPLICATION FOR MEMBERSHIP.inside back cover

VII. NOTICE TO CONTRIBUTORS

.inside back cover

JOURNAL OF THE ENTOMOLOGICAL SOCIETY OF ONTARIO - VOLUME 146, 2015