ISSN 1713-7845
JOURNAL
of the Hie
ENTOMOLOGI@ a
SOCIETY —
OF
ONTARIO
Volume
One Hundred and Forty Four
2013
Published December 2013
ISSN 1713-7845
JOURNAL
of the
ENTOMOLOGICAL SOCIETY
of
ONTARIO
Volume One Hundred and Forty Four
2013
THE ENTOMOLOGICAL SOCIETY OF ONTARIO
OFFICERS AND GOVERNORS
2012-2013
President: J. SKEVINGTON
Agriculture and Agri-Food Canada,
K.W. Neatby Building
960 Carling Avenue, Ottawa, ON K1A 0C6
jskevington@gmail.com
President-Elect: J. MCNEIL
Department of Biology
Biological and Geological Sciences Building
University of Western Ontario, London, ON N6A 5B7
jmeneil2@uwo.ca
Past-President: B. GILL
Entomology Unit, Ontario Plant Laboratories,
Canadian Food Inspection Agency. Building 18 C.E.F.
960 Carling Ave., Ottawa, ON K1A 0C6
bruce. gill@inspection.ge.ca
Secretary: N. MCKENZIE
Vista Centre, 1830 Bank Street, P.O. Box 83025
Ottawa, ON K1V 1A3
nicole_mcekenzie@he-sc.ge.ca
Treasurer: S. LI
Pest Management Centre, Building 57
Agriculture and Agri-Food Canada
960 Carling Ave., Ottawa, ON K1A 0C6
sli@nrean.ge.ca
Directors:
C. BAHLAI (2012-2014)
Deparment of Entomology, Michigan State University
Center for Integrated Plant Systems
578 Wilson Rd., East Lansing MI USA 48824
cbahlai@msu.edu
R. BUITENHUIS (2011-2013)
Vineland Research and Innovation Centre
4890 Victoria Ave. North, P.O. Box 4000
Vineland, ON LOR 2E0
rose.buitenhuis@vinelandresearch.com
S. CARDINAL (2013-2015)
Agriculture and Agri-Food Canada
960 Carling Ave., Ottawa ON K1A 0C6
sophie.cardinal@agr.ge.ca
J. GIBSON
Department of Integrative Biology
University of Guelph
Guelph,ON N1G 2W1
jfgibson@uoguelph.ca
B. SINCLAIR
Department of Biology
University of Western Ontario
London, ON N6A 5B7
bsincla7@uwo.ca
(2012-2014)
(2013-2015)
ESO Regional Rep to ESC: H. DOUGLAS
Canadian Food Inspection Agency
960 Carling Ave., Ottawa ON K1A 06C
douglash@inspection.ge.ca
Librarian: J. BRETT
Library, University of Guelph
Guelph, ON NIG 2W1
jimbrett@uoguelph.ca
Newsletter Editor: A. GRADISH
School of Environmental Science
University of Guelph, Guelph ON NIG 4Y2
agradish@uoguelph.ca
Student Representative: A. FREWIN
Integrated Biology, University of Guelph
Guelph, ON NIG 2W1
afrewin@uoguelph.ca
Website: M. JACKSON
School of Environmental Science
University of Guelph, Guelph, ON NIG 2W1
jackson@uoguelph.ca
JESO Editor: J. HUBER
Canadian National Collection of Insects
Agriculture and Agri-Food Canada
960 Carling Ave. Ottawa, ON,K1A 0C6
John.Huber@agr.ge.ca
Technical Editor: J. VICKRUCK
Dept. of Biological Sciences, Brock University
500 Glenridge Ave., St. Catharines, ON L2S 3A1
jess.vickruck@pbrocku.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, N1G 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 144, 2013
JOURNAL
of the
ENTOMOLOGICAL SOCIETY OF ONTARIO
VOLUME 144 2013
Prior to the successful Joint Annual Meeting of the Entomological Societies of
Ontario and Canada in Guelph your Board met and discussed how to improve the profile
of JESO amidst all the competition from the numerous new electronic journals. In his 1999
editorial, past editor Dolf Harmsen had foreseen some of the problems the Proceedings
would face with the advent of electronic publishing. Since then, past editor Miriam
Richards overcame some of the hurdles and for almost a decade JESO has been publishing
electronically. It continues to be printed as hard copy as well. Having both formats has its
merits and for historical reasons—it is one of the longest running entomology journals in the
world, with no publication breaks since 1871—the hard copy will continue to be published.
It is an important part of ESO’s heritage. Greater visibility for JESO on the internet is now
needed and some good ideas for obtaining this were presented. Ideas for improving JESO’s
impact were also discussed. One action approved by the Board and promptly implemented
by past president Jeff Skevington was to sign an agreement with the Biodiversity Heritage
Library to have all back issues scanned, starting with Volume | of the Annual Report and
make them available on the website, with a two year embargo on the most recent volume.
This year’s volume contains five scientific notes and one scientific paper. All but
one (on taxonomy) report new species records for Ontario, new distributions or new host
records. Two papers are overviews of the past 60 years of JESO papers on two topics.
One, on Taxonomy and Faunistics, is mostly a summary. The other, much more detailed
and comprehensive is on Biological Control. Both were written to commemorate the 150"
meeting of ESO. If there is any trend in the kinds of papers submitted in recent years it
is more towards more papers in these areas of entomology and fewer on economically
important pest species and their control. Papers on all aspects of entomology are, of course,
welcome and my hope is that you, the readers, will continue to find JESO a good place to
publish your research.
John T. Huber
Editor
JESO Volume 144, 2013
New range records of mosquitoes from Northern Ontario JESO Volume 144, 2013
NEW RANGE RECORDS OF MOSQUITOES (DIPTERA:
CULICIDAE) FROM NORTHERN ONTARIO
J. L. RINGROSE', K. F. ABRAHAM?, D. V. BERESFORD"
'Department of Biology, Trent University,
2140 East Bank Drive, Peterborough, ON K9J 7B8
email: davidberesford@trentu.ca
Abstract J ent. Soc. Ont. 144: 3-14
A survey for mosquitoes at 23 sites in the Ontario Shield and Hudson
Bay Lowlands of northern Ontario, Canada, in 2011 and 2012 yielded 19
species, including 16 of Aedes, and one each of Anopheles, Coquillettidia,
and Culesita. One species, Aedes pullatus (Coquillett) is newly recorded for
Ontario. Eleven northern range extensions and one southern range extension
are reported.
Published December 2013
Introduction
The distributions of many mosquito species (Diptera: Culicidae) in Canada are
incomplete. Jenkins and Knight (1952) conducted a survey of larval mosquitoes in southern
James Bay. Steward and Mc Wade (1960) published range summaries of species in Ontario.
Wood et al. (1979) compiled the most complete account of mosquito distribution in Canada
The Canadian Endangered Species Conservation Council (CESCC 2011) assessed the
status of many species, including mosquitoes. Yet, areas such as northern Ontario are still
relatively little sampled.
Northern Ontario has become the focus of increased mineral exploration and
development (FNSAP 2010). Additionally, the area is projected to undergo significant
ecological transformation over the next several decades due to climate change (FNASP
2010). Together, these two driving forces create a need for better knowledge of species’
distributions in northern Ontario before significant changes occur. A biological diversity
survey of different taxa in northern Ontario was initiated in 2009 to address this issue
(OMNR 2012). The species composition and diversity information obtained will help
determine land use, and management and conservation planning, as well as provide baseline
information to determine the impact of mining and climate change.
* Author to whom all correspondence should be addressed.
* Wildlife Research and Development Section, Ontario Ministry of Natural Resources, 2140
East Bank Drive Peterborough, ON K9J 7B8
Ringrose et al. JESO Volume 144, 2013
Mosquito species lists for particular geographic areas include species that have not
been collected there but are assumed to be present based on information from adjacent areas
(e.g., Wood et al. 1979; Darsie and Ward 2005). Thus, it is reasonable to expect species to be
found in northern Ontario if they have been found in similar habitats and at similar latitudes
elsewhere, i.e., in spite of regional climatic differences, we expected to find species that
have existing records from both adjacent western Quebec and northern Manitoba because
of the large scale continuity of the ecosystems in the boreal and subarctic forests that
span these three provinces. For mosquito species whose known distributional limits were
either south or north of our study areas, we expected to extend known ranges north or
south, respectively. Following this reasoning, and based on the range maps provided by
Wood et al. (1979) and Darsie and Ward (2005), we predicted a maximum of 31 species
in our surveys. In this paper we report new information on occurrences of known species
(range extensions), new collection locations and records of species new to the province
for Culicidae in Ontario from surveys of previously unexplored areas of the far north of
Ontario. We use both rarefaction and a lognormal analysis to explore the maximum number
of species predicted in these areas and to gauge their relative abundances.
Materials and Methods
Sampling took place within two different northern Ontario ecozones: the Ontario
Shield and Hudson Bay Lowlands ecozones (Crins et al. 2009) in 2011 and 2012, hereafter
referred to the western and eastern study areas, respectively, as part of a larger biological
survey of animal and plant taxa undertaken by the OMNR (2012). The 2011 sampling
occurred within 150 km of the First Nations communities near Big Trout Lake and Sandy
Lake in the western study area. The 2012 sampling occurred within 150 km of the First
Nations community of Fort Albany in the eastern study area. In each year 12 sample sites
were randomly selected from the computer generated grid of National Forest Inventory
(NFI) points (Gillis et al. 2005). Actual sample locations sometimes differed by as much
as 15 km from the NFI coordinates depending on feasibility of landing a helicopter. Our
plot locations are the sites at which field camps were established (OMNR 2012). Sample
locations were within | km of the field camp, which was verified using a handheld GPS
(Garmin Rino 530HCx, NAD83, +3m accuracy). Sampling occurred from 29 May to 17
July in 2011 and 4 June to 5 July in 2012.
Habitats at these sites were dominated by coniferous and shrub wetlands comprised
largely of black spruce (Picea mariana Britton, Sterns & Poggenb.) and tamarack (Larix
laricina (Du Roi) K. Koch) as well as shrub and sedge fens, and sphagnum bog. The sites
sampled in 2012 in the eastern study area generally had more standing water than those
sampled in 2011 in the western study area.
In both years the mosquito component of the sample regimen included daily sampling
both by individual collection (ad hoc, when mosquitoes were present, approximately 30
minutes total), and a dusk and dawn sweeping with an insect net for 6 minutes at each
sampling location. Individual collection consisted of catching mosquitoes that landed on the
face, arms, and legs of field crew members using snap cap vials (2.0 ml) before they had a
chance to bite. These collections occurred throughout the day and late evening. Individual
New range records of mosquitoes from Northern Ontario JESO Volume 144, 2013
specimens in snap vials were preserved dry in the capture vials. Adult mosquitoes collected
by sweeping were placed in labeled sample jars with a silica desiccant to prevent deterioration
from moisture. A large proportion of them had scales on their thoraces abraded and so could
not be identified to species. Therefore, more effort was placed on individual collection in
2012. All specimens were pinned and identified by JLUR and DVB using the keys of Wood et
al. (1979), and Thielman and Hunter (2007). Nomenclature was based on the WRBU Online
Catalog (2013). Voucher specimens were assigned individual specimen numbers (Table 2)
and are stored at the Trent University Biology Department in Peterborough, Ontario. Some
vouchers are deposited in the Canadian National Collection of Insects, Ottawa.
Analysis
Rarefaction analysis for the 2011 and 2012 catch data was performed using software
on the University of Alberta website (http://www.biology.ualberta.ca/jbrzusto/rarefact.
php). This method relates sampling effort to number of species caught. The total number of
species caught each year is used to calculate the expected number of species (with standard
deviation) that would have been caught if fewer mosquitoes were sampled overall. Different
species numbers for the same total catch sizes indicate community differences such as those
due to site, e.g., habitat or phenological, or procedural differences.
We also fit the catch data (Table 1) to a lognormal distribution using the sum of
squares method, i.e., Preston’s method as described in Ludwig and Reynolds (1988). This
allowed us to calculate the expected number of species by estimating the number of rare
species not found in the samples. Essentially, it assumes that species of low abundance, e.g.,
about | per 1000 individuals, will only be found if at least 1000 individuals are collected.
The lognormal distribution uses the abundance of different species and groups them into
octaves or doubled catch classes, e.g., 0-1 individuals, 1—2 individuals, 24 individuals,
4-8 individuals and so on, and fits these frequencies to a lognormal curve by aligning the
mode. Species that had only one individual caught could go into either the first or second
class, so the number was divided between these classes, e.g., if one catches 5 species with
only one individual each, then half of these (2.5) are assigned to the 0-1 class, and 2.5 to the
1-2 class (Ludwig and Reynolds, 1988). One of the assumptions of this method is that very
rare species will not be sampled, but can be calculated from the area of the normal curve
to the left of the 0—1 class or veil line. The biological interpretation is that this class (0-1)
would become the 1—2 class if our total catch size was increased. This analysis requires an
iterative method to find values for two parameters that provide the best fit: a (width), and
So (height). We used the SOLVER optimization add-in function in Microsoft Excel 2007
version for this task.
Results
We caught 896 mosquitoes in 2011 and 826 in 2012. Mosquitoes caught directly
from the face and arms and housed in vials could all be identified to species, whereas
only 117 (13%) of individuals from 2011 and 192 (21%) from 2012 sweeping could be
identified to species. Species collected and collection locations are summarized in Tables
1 and 2. Twelve species were collected in the western study area in 2011 and 16 species
JESO Volume 144, 2013
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New range records of mosquitoes from Northern Ontario JESO Volume 144, 2013
in the eastern study area in 2012 (Fig. 1, Table 1). The most abundant species identified in
both years was Coquillettidia perturbans (Walker). Rare species, i.e., those represented
by a single individual collected in either year were Aedes cinereus Meigen, Ae. nigripes
(Zetterstedt), Ae. provocans (Walker), Ae. pullatus (Coquillett), Ae. rempeli Vockeroth and
Culiseta impatiens (Walker).
Fitting to the lognormal distribution (Fig. 2), the expected number of species was
14.75 from the 2011 catches (fitted parameters a = 0.24, So = 2.0, Chi sq = 1.23, p = 0.94,
d.f. = 5) and 23.4 species in the 2012 catches (fitted parameters a = 0.225, So = 2.97, Chi
sq = 5.46, p = 0.36, d.f. = 5). By combining the two year’s totals, our expected number of
species for northern Ontario was 28.2 species (fitted parameters a = 0.21, So = 3.35, Chi sq
= 2.94, p = 0.82, d.f. = 6).
Interpretations of new records and range extensions are based on comparison with
range maps in Wood et al. (1979).
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Big Trout Lake and Sandy Lake in 2011 (closed circles) and within 150 km of Ft. Albany
in 2012 (open circles). The inset map of Ontario shows the sampling locations in 2011 and
2012.
Ringrose et al. JESO Volume 144, 2013
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2011 because collections from one of the July 12-21 sites were damaged by a bear. TUIC numbers are voucher specimens in the Trent
0003 | 4edes cinereus
TABLE 2. Mosquito species found at each sampling site. Dates indicate when sampling was conducted. Only 11 sample sites listed in
University Insect Collection.
JESO Volume 144, 2013
New range records of mosquitoes from Northern Ontario
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New Ontario record
Aedes pullatus has two distinct distributions, an eastern population in northern
Quebec and Labrador and the western population in Alberta, British Columbia, and the
Yukon (Wood et al. 1979). The single specimen we collected in the eastern study area is the
first record in Ontario and extends the range of the eastern population westward.
Northward range extensions
Aedes canadensis (Theobald) is a widely distributed species found in forested
regions of all Canadian provinces and the Yukon (Steward and McWade 1960). It is known
to be found in Moosonee and Moose Factory in Ontario. Our collection was in the eastern
study area.
Aedes cinereus is a common species in Ontario and has been found in Moosonee,
Moose Factory and the town of Kenora (Steward and McWade 1960). Jenkins and Knight
(1952) noted that Ae. cinereus was the most common larval species that they collected in
the southern James Bay area but, oddly, they collected no adults. Our single specimen was
collected in the eastern study area.
Aedes dorsalis (Meigen) is a rare northern species and in Ontario has only been
collected in Moosonee and Moose Factory (Steward and McWade 1960). It was only
collected in the eastern study area, which is not surprising because of its relative proximity
to these communities.
3
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64-128
FIGURE 2. Fitted lognormal distributions of mosquito catches within 150 km of Big Trout
Lake and Sandy Lake in 2011 and within 150 km of Ft. Albany in 2012. The area of the
region left of the veil line represents species that were too rare to be sampled with our
methodology.
10
New range records of mosquitoes from Northern Ontario JESO Volume 144, 2013
Aedes implicatus (Vockeroth) is common in the northern and central parts of
Ontario and has been collected in Moose Factory (Steward and McWade 1960). It was
collected in both study areas.
Aedes excrucians (Walker) is found throughout North America (Wood et al. 1979).
It was collected by Jenkins and Knight (1952) in Moose Factory and Moosonee and by
Steward and Mc Wade (1960). Our collection from the western study area provides a record
for the gap between the eastern James Bay coast and Manitoba.
Aedes intrudens Dyar is found south of the tree line in late spring (Wood et al.
1979). It has been recorded from all provinces (Steward and McWade 1960). The species
was common in the eastern study area, but was not found in the western study area.
Aedes provocans is a forest species and is a southern species in Ontario (Wood
et al. 1979), except for a single record from Great Slave Lake, Northwest Territories. We
collected a single specimen in the eastern study area.
Aedes rempeli is one of the rarest Canadian species (Vockeroth 1954). However,
Wood et al. (1979) suggested that this species may be widely but sparsely distributed in
northern Ontario. We caught a single specimen along the Albany River about 150 km
upstream from the James Bay coast.
Anopheles earlei Vargas is the most common species of this genus in Ontario. Our
collections of this species in both study areas extend the known range.
Coquillettidia perturbans is common in southern Ontario (Wood et al. 1979).
Jenkins et al. (1952) found that this species was very abundant in a spruce forest west of
Cochrane, Ontario. In both study areas it was our most abundant species.
Culiseta impatiens is a northern species usually found in forested regions and has
been recorded from Moose Factory (Steward and McWade 1960). Our single specimen
came from the western study area, providing a westward extension of the known range.
Southward range extensions
Aedes nigripes is an arctic species whose range, according to Wood et al. (1979),
did not extend southward into Ontario. However, one recent record exists from Polar Bear
Provincial Park (Beresford 2011). One specimens was collected in the western study area in
2011, even farther south than Polar Bear Provincial Park.
Range gap infills
Aedes abserratus (Felt and Young) is an uncommon species in Ontario (Wood et
al, 1979). Steward and McWade (1960) reported the species from Moose Factory. Beresford
(2011) collected it in Polar Bear Provincial Park. Our collection of this species in both study
areas fills the gap. Aedes communis (De Geer) is one of the most widely distributed species
in the northern hemisphere. Beckel (1954) stated that this species was rarely collected in the
Churchill area of Manitoba because it is non-biting in that area. In Ontario, records show it
to be generally present and often abundant throughout the province. This species was well
represented (9.4%) in our collections from the western study area, but less so (1%) in the
eastern study area.
Ringrose et al. JESO Volume 144, 2013
Aedes hexodontus Dyar has been collected in Churchill, Manitoba both as larvae
(Vockeroth 1954) and as adults (Beckel 1954), and also from western Quebec and western
Ontario (Wood et al, 1979). Our collection fills the gap.
Aedes impiger (Walker) is generally found in Nunavut and the Northwest Territories
(Steward and McWade 1960). It has been caught in Ontario at Moose Factory and along the
Albany River (Steward and McWade 1960) and in Manitoba at Churchill (Downes 1965).
Our collections from our western study area fill a gap between Churchill and the James Bay
coast in Quebec. Surprisingly, we did not find any in our eastern collections, which are close
to James Bay.
Aedes pionips Dyar is found in the forests of central and northern Canada, and has
been collected from Moose Factory, Ontario (Steward and McWade 1960) and Churchill,
Manitoba (Beckel 1954). Not unexpectedly, our collections fill the gap.
Aedes punctor (Kirby) is a common species in Ontario and throughout Canada
(Steward and McWade 1960). Records are from Moosonee (Jenkins and Knight 1952) and
Churchill, Manitoba (Beckel 1954). Our collections are within the expected range but fill
distributional gaps in northwestern Ontario.
Discussion
As expected we produced new distributional records, including both northward and
southward range extensions, and filled gaps in known ranges. All of the species we collected
are considered by CESCC (2011) to be secure (relatively widespread or abundant), except
for five with undetermined status: Aedes impiger, Ae. implicatus, Ae. pionips, Ae. rempeli
and An. earlei.
The rarefaction analysis, which standardizes across different sample sizes, indicates
that the eastern region (2012) had slightly more species than the western region (2011). For
example, in collections of 100 individuals we would only have been able to catch about 13
species in the east compared to 11 in west (Fig. 1). The lognormal analysis shows the same
pattern, with 23.4 species predicted to be in the eastern region compared to 14.75 in the
western region (Fig. 2). These analyses reveal that this difference in species richness may be
a function of the different regions (e.g., habitats) rather than catch effort. The 2012 eastern
study area collections were from sites with lower elevations (1—88 m) than the western sites
(148-379 m). However, because these two regions were sampled in different years, we
cannot attribute this difference to region alone.
From our survey of the range maps we expected to find up to 31 species. Fitting the
lognormal distribution to our overall catch numbers, our expected number of species was
28, a good estimate of species richness of this region.
In fact, we found only 19 species and four of the species we did catch were not
expected from the range map analysis: Aedes nigripes, Ae. provocans, Ae. pullatus, Ae.
rempeli. This means that 16 species from the range map analysis were expected but not
found, either due to our sampling methods, phenology, or habitat preferences. Of these,
Wyeomyia smithii (Coquillett) is fully autogenous and has not been reported bloodfeeding;
Ae. diantaeus Howard, Dyar and Knab is not found in coniferous forests; Ae. spencerii
(Theobald) is not found in forest regions; Ae. sticticus (Meigen) is generally restricted to
12
New range records of mosquitoes from Northern Ontario JESO Volume 144, 2013
floodwaters of rivers; Culesita morsitans (Theobald) and Culex restuans Theobald prefer
to bloodfeed from birds; Culex territans Walker prefers reptiles and amphibians; Culesita
alaskaensis (Ludlow) and Ae. mercurator Dyar are early spring species; An. walkeri
(Theobald), Ae. vexans (Meigen) and Ae. campestris Dyar & Knab are primarily nocturnal
biters. The remaining four of the expected species are rare, Ae. riparius Dyar & Knab, Ae.
flavescens (Miiller), Ae. fitchii (Felt & Young) and Ae. decticus (Howard, Dyar & Knab)
(Wood et al. 1979).
All collection methods have inherent biases associated with them (Muirhead-
Thomson 1991). Some important limitations to this survey are that collections occurred at
randomly chosen sites (i.e., not selected for high probability of detecting mosquitoes) and
using simple methods that were part of a larger diversity survey. The mosquito portion of
that survey was limited by the logistics of available time and equipment at these remote
sites. A collection effort that focused on targeting mosquitoes alone, within specific habitats,
would likely have produced more of the expected species, and the use of CO, traps of CDC
light traps would have produced far larger collections. Nevertheless, this study, despite
its limitations, indicates that surveys undertaken in under-sampled regions can produce
important baseline information that extends the previously known ranges.
Acknowledgements
The authors would like to thank the Ontario Ministry of Natural Resources
Northeast Science and Information Section and Wildlife Research and Development Section
for project coordination and logistics, and Far North Branch for funding. Additional travel
support for JLR was provided by a Northern Scientific Training Program (NSTP) grant
through Trent University. We give special thanks to Dean Phoenix and the field crews of
the Far North Biodiversity Project in 2011 and 2012. We would also like to thank the First
Nations communities of Kitchenuhmaykoosib Inninuwug, Keewaywin and Fort Albany for
their hospitality and generosity.
References
Beckel, W. E. 1954. The identification of adult female Aedes mosquitoes (Diptera, Culicidae)
of the black-legged group taken in the field at Churchill, Manitoba. Canadian
Journal of Zoology 32: 324-330.
Beresford, D. 2011. Insect collections from Polar Bear Provincial Park, Ontario, with new
records. Journal of the Entomological Society of Ontario 142: 19-27.
CESCC 2011. Wild species 2010: the general status of species in Canada. National General
Status Working Group. Canadian Endangered Species Conservation Council.
Available online at: http://www.wildspecies.ca/wildspecies20 1 0/downloads/wild-
species-2010.pdf
Crins, W. J., Gray, P. A.,Uhlig, W. C. and Wester, M. C. 2009. The ecosystems of Ontario,
Part 1. Ecozones and ecoregions. Report SIB TER IMA TR-01. Ontario Ministry of
Natural Resources Inventory, Monitoring and Assessment Section. Peterborough,
ON. 71 pp.
Ringrose et al. JESO Volume 144, 2013
Darsie, R. F. and Ward, R. A. 2005. Identification and geographical distribution of the
mosquitoes of North America, North of Mexico. University of Florida Press,
Gainesville, FL. 400 pp.
Downes, J. A. 1965. Adaptations of insects in the arctic. Annual Review of Entomology 10:
257-274.
FNSAP. 2010. Science for a changing far north. The Report of the Far North Science
Advisory Panel. A report submitted to the Ontario Ministry of Natural Resources.
Queen’s Printer for Ontario, Toronto, ON. 141 pp.
Gillis, M. D., Omule, A. Y. and Brierley, T. 2005. Monitoring Canada’s forests: the national
forest inventory. Forestry Chronicle 81: 214-221.
Jenkins, D. W. and Knight, K. L. 1952. Ecological survey of the mosquitoes of southern
James Bay. American Midland Naturalist 47: 456-468.
Ludwig J. A. and Reynolds, J. F. 1988. Statistical Ecology: a primer on methods and
computing. John Wiley & Sons., New York, NY. 337 pp.
Muirhead-Thomson, R. C. 1991. Trap responses of flying insects. The influence of trap
design on capture efficiency. Academic Press, San Diego, CA. 304 pp.
OMNR. 2012. Wildlife research: far North biodiversity. Ontario Ministry of Natural
Resources. Available online at: — http://www.mnr.gov.on.ca/en/Business/
Wildlife/2ColumnSubPage/STDPROD _099955.html.
Steward, C. C. and McWade, J. W. 1960. The mosquitoes of Ontario (Diptera: Culicidae)
with keys to the species and notes on distribution. Proceedings of the Entomological
Society of Ontario 91: 121-188.
Thielman, A. C., and Hunter, F. F. 2007. Photographic key to the adult female mosquitoes
(Diptera: Culicidae) of Canada. Canadian Journal of Arthropod Identification 4,
Available online at: http://www.ualberta.ca/bsc/ejournal/th 04/th_04.html, doi:
10.3752/cjai.2007.04
Wood, D. M., Dang, P. T. and Ellis, R. A. 1979. The mosquitoes of Canada (Diptera:
Culicidae). The Insects and Arachnids of Canada, Part 6: Agriculture Canada.
Publication 1686. 390 pp.
Vockeroth, J. R. 1954. Notes on the identities and distributions of Aedes species of northern
Canada, with a key to the females (Diptera: Culicidae). The Canadian Entomologist
6: 241-255.
WRBU. 2013. Traditional Mosquito Classification, July 2013. Walter Reed
Biosystematics Unit. Available online at: http://www.wrbu.org/docs/mq_
ClassificationTraditional201307.pdf.
Taxonomy and faunistics in Ontario, 1952-2012 JESO Volume 144, 2013
TAXONOMY AND FAUNISTICS IN ONTARIO, 1952-2012:
PUBLICATIONS IN THE “JOURNAL OF THE ENTOMOLOGICAL
SOCIETY OF ONTARIO”
J.T. HUBER
Natural Resources Canada c/o Agriculture and Agri-Food Canada, Research Centre,
960 Carling Avenue, Ottawa, ON K1A 0C6
email: john.huber@agr.gc.ca
Abstract J. ent. Soc. Ont. 144: 15-26
Publications on taxonomy and faunistics that appeared in the Journal of the
Entomological Society of Ontario over a 60-year period beginning in 1952
are tabulated. These consist of 60 papers on taxonomy with a total of 700
species, including 125 new ones, described and/or keyed. Almost 100 papers
on faunistics (lists, new distributions for North America or parts of North
America) were published, with a total of 4700 species mentioned. A brief
overview of taxonomy and faunistics as given in JESO volumes is provided.
Published December 2013
Introduction
Although the Entomological Society of Ontario (ESO) began in 1863, the first
report was published in 1871 (covering the year 1870) and publication continued as Annual
Reports up to 1958, then from 1959-2001 as the Proceedings of the Entomological Society
of Ontario (PESO), and finally from 2002—present as the Journal of the Entomological
Society of Ontario (JESO). Because the main goal of the Society at its inception was to
publish research on pest insects the first volume was titled “First Annual Report of the
Noxious Insects of the Province of Ontario” and subtitled “Prepared for the Agricultural and
Arts, and Fruit Growers’ Associations of Ontario, on Behalf of the Entomological Society
of Canada.” The first two articles in it were by the Rev. C. J. S. Bethune, entitled “Insects
affecting the apple” and “Insects injurious to grape.” The 61 magnificent black-and-white
illustrations throughout Vol.1 and the 694 others in the next twelve volumes (illustrations
summarized in detail in Vol. 13) stand as a testament to the careful attention to detail in the
published papers.
Forward to the 1950s. Glen (1956) compiled a historical overview of entomology
in Canada with contributions by different authors in 16 categories, one being Systematic
Entomology by G. Holland. One category not treated by Glenn as a separate subdiscipline
was faunistics, probably because most taxonomy and biology papers included information
on insect distributions, even if it was for a single (usually pest) species, so “faunistics” was
too vague to treat as a subdiscipline. To mark the 150" Anniversary of the Entomological
Society of Ontario, papers in these two subdisciplines are compiled and briefly discussed
15
Huber JESO Volume 144, 2013
here. Only the past 60 years are treated, beginning with publication date 1953 (vol. 84) to
provide a slight overlap with Holland (1956). The subdiscipline of taxonomy, “systematic
entomology” of Holland, is complemented with a summary of papers on faunistics. The
latter were written by both taxonomists and non-taxonomists, but the non-taxonomists
relied heavily on taxonomists for specimen identifications. Over the past six decades both
groups of entomologists added a lot of new information on insect distributions in Ontario
or Canada. Because they do not include identification keys or taxon descriptions the papers
on faunistics are summarized separately from those on taxonomy. Except for 19 papers
on particular insect species, and five on insect associations with certain plant species, the
faunistics papers exclude studies that detail the biology of single species, most of which are
economically important as pests or biological control agents. Such papers are treated by P.
Mason (this volume).
Taxonomy
Only about | taxonomy paper per year (60 in total) was published over the past
60 years (Table 1). These covered almost 700 species of which 125 were described as
new. Twenty-five of the papers treated Ontario insects only. Most of them (47) included
identification keys, usually to adults but sometimes to larvae or pupa. Somewhat surprisingly,
25 of the papers treated Diptera, 25 treated Hymenoptera but only 7 treated Coleoptera and
1 treated Lepidoptera. About 36 family group taxa and 50 genera were covered. A few
papers were more general, treating Lepidoptera, Aculeata, and Symphyta. Two were on
nomenclature and type specimens, respectively.
Although most publications in the Annual Reports over its first 80 years treated
pest biology and control, the occasional paper foreshadowed the trend over the next 60 years
towards more papers on taxonomy and faunistics of insects in general. Fletcher (1902), the
founder of the Canadian National Collection of Insects, therefore began an Entomological
Record. His aim was not to record facts connected to economic entomology—he called it
“practical” entomology—but instead to publish information about other insects, including
1) arecord of special rarities taken by collectors, with the various locations and dates, 2) the
names of specialists who have devoted particular attention to some order, genus, species,
or phases of taxonomic study, 3) the names of any books of note affecting entomology,
or connected with any branch of it, which may have been published during the year. For
the year, Fletcher summarized collecting thus “The season of 1901 in almost all parts
of Canada has been characterized as ‘poor’ by nearly all collectors heard from.” Most of
Fletcher’s publication gives a literature summary, lists of names and locations of collectors
(36 for Lepidoptera, 10 for Coleoptera, and three each for Hymenoptera and Orthoptera),
and 8 pages of “notes on captures” compiled by himself and, for Orthoptera, by E. M.
Walker. The next year, Fletcher (1903) stated that he hoped at least some of the general
collectors in all parts of the country might become specialists on particular taxa because
they were urgently needed. He noted that the Lepidoptera and Coleoptera were always
fairly well worked but specialists in the other orders were few. Fernald’s (1916) paper on
life zones in entomology and Felt’s (1926) paper on insect distributions presage Walker’s
(1955) discussion on climate change, mentioned below. So right from the beginning of
the 1900s there was interest and concern about taxonomy and distributions, particularly
changing ones, of the insect fauna of Canada in general and Ontario in particular.
16
Taxonomy and faunistics in Ontario, 1952-2012 JESO Volume 144, 2013
Faunistics
In 1961, C.G. MacNay’s yearly article “A summary of the more important insect
infestations and occurrences in Canada in 19xx” ceased to be published. Although this series
of articles focused almost exclusively on pest insects from across Canada, other noteworthy
species were occasionally mentioned. Thus, for 1950 (eighty-first Annual Report) one and
a half lines were written on one species not considered a pest: “The noticeable scarcity of
reports of this insect [the Painted Lady, Vanessa cardui L. (Lepidoptera: Nymphalidae)]
contrasts with its widespread abundance in 1949”. The rest of the 19-page article summarized
the abundance of pest species under several subheadings: general feeders, field crop
insects, vegetable insects, fruit insects, insects affecting greenhouse and ornamental plants,
insects affecting man and domestic animals, household insects, stored product insects. All
the other articles in the volume related to pesticides—it was, after all, shortly after start
of the pesticide heyday/revolution. Similar examples occur in the 1951 Annual Report
[one mention of a Mourning Cloak butterfly larva, Nymphalis antiopa (L.) (Lepidoptera:
Nymphalidae)]. Almost invariably the record was in the context of damage to something
of economic interest. Preventing damage to crops/animals/humans was seen as perhaps
the most important task of entomology. In 1952, 150 years after Fletcher implemented
his ‘Entomological Record’, a section on new records of insects in Canada was added. It
included 17 species (5 in Ontario) recorded for the first time either for North America or for
Canada or for a particular province. From 1966—1972, H. W. Goble and others published
articles under the subheading “Review of infestations and other pests” but restricted their
coverage to insects (and nematodes) in Ontario only. Vol. 104 (1973) was the last year such
pest summaries were compiled. Thereafter, relatively more attention was paid to insects not
of economic importance.
In 1954, a symposium on changing faunal ranges was held, in which various
speakers discussed examples (in Lepidoptera, Ephemeroptera, Orthoptera, Hymenoptera,
and Araneae) of Carolinian zone insects that showed evidence of a northward shift in
distribution. Some crop pests already present in southern Ontario or new entries of insects
into the Niagara Peninsula were included. Also exemplified were extensions of faunal
ranges in the Prairie Provinces, species spreading with agriculture, species whose ranges
fluctuate with climatic cycles, and species with annual northern migrations. Northern
shifts in populations of some bird species in the Prairie Provinces and alien pest insect
species introduced from abroad were also listed. The eminent E.M. Walker (1955), of
odonatological fame, summarized things thus: “But looking back over the sixty odd years
since I began to collect insects at De Grassi Point, Lake Simcoe, I have witnessed the
gradual decrease in numbers of some species that were once common, until they vanished
altogether, and I have seen other species, never known in that territory before, arrive there
and in the course of time become firmly established. The species that disappeared were
chiefly northern ones, whereas the newcomers were all from the south. This last statement
suggests a changing climate that is becoming warmer. The problem, however, is not quite
as simple as it seems.” I take the Baker symposium, mentioned in Table 2, and Walker’s
comments as the main post-war starting point for the shift in emphasis on controlling pests
to documenting and understanding the Ontario insect fauna in general, with emphasis on
changing distributions. However, over 150 years previously Webster (1902) noted general
Wi
Huber JESO Volume 144, 2013
trends in insect movements around North America. Fernald (1916) and Felt (1926) also
wrote about distributions and their significance, showing that within about 30 years since
publication of the Annual Reports began entomologists were aware of the importance of
tracking insect distributions.
Almost 100 papers on faunistics were published from 1953-2012 (Table 2), with
a low of 8 papers in the 1950s to a high of 19 in the 1970s. Many of these are species
lists, changes in distributions, or new provincial, country, or continent records. About
4700 arthropod (mostly insect) species in over 20 orders, especially Coleoptera, Diptera
and Hymenoptera, are listed. A wide variety of faunistic topics are covered: insects on
particular substrates, e.g., decaying mushrooms; in particular habitats, e.g., alvars; visitors to
particular species of flowering plants, e.g., Daucus carota L. (Apiaceae); or natural enemies
of particular, non-pest insects, e.g., Bombus spp. (Hymenoptera: Apidae). Almost every
volume included at least one faunistics paper and a few volumes (104, 141, 142) as many
as five. Most papers were restricted to insects of Ontario or parts of Ontario. Occasionally
other provinces (Manitoba, Newfoundland, Nova Scotia, Quebec), or the USA or particular
US states were treated. Sometimes all of Canada, the Nearctic region (usually America
North of Mexico) or the entire New World (an abstract only) was covered.
Conclusions
Up to the 1950s the Annual Reports stressed pest biology and control, and many
detailed papers appeared on their biology often accompanied by excellent line drawings.
The Reports also included a smattering of more general papers discussing distributions
(read faunistics) and taxonomy. Over the past six decades a greater diversity of papers has
appeared, with relatively more emphasis on insects other than pests. On the whole, the
Society’s journal has provided a fair representation of entomological research in Ontario
over most of the past 140 years. This has changed over the past decade. Fewer papers are
published in JESO because of the greater number of competing, electronic journals, often
with more specialized interests. JESO is therefore perhaps a less reliable tracker of entomo-
logical research in the province than previously. Nevertheless, JESO remains a good venue
for publishing information on faunistics and taxonomy of Ontario insects.
SO Volume 144, 2013
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References
Felt, E. P. 1926. The distribution of insects and the significance of extralimital data. Fifty-
sixth Annual Report of the Entomological Society of Ontario, 1925. Pp. 44-47.
Fernald, H. T. 1916. Life zones in entomology and their relations to crops. Forty-sixth
Annual Report of the Entomological Society of Ontario, 1915. Pp. 87-92.
Fletcher, J. 1902. Entomological record, 1901. Thirty-second Annual Report of the
Entomological Society of Ontario, 1901. Pp. 99-109.
Fletcher, J. 1903. Entomological record, 1902. Thirty-third Annual Report of the
Entomological Society of Ontario, 1902. Pp. 97-101.
Glenn, R. 1956. Entomology in Canada up to 1956: a review of developments and
accomplishments. The Canadian Entomologist 88: 290-371.
Holland, G. P. 1956. Systematic Entomology. Pp. 300-304 in Glenn, R. (compiler),
Entomology in Canada up to 1956: a review of developments and accomplishments.
The Canadian Entomologist 88: 290-371.
Walker, E. M. 1955. Summary of symposium. Pp. 37—38 in Baker, A. W. (co-ordinator),
Symposium II. Changing faunal ranges. Annual Report of the Entomological
Society of Ontario 86: 23-39.
Webster, F. M. 1902. The trend of insect diffusion in North America. Thirty-second Annual
Report of the Entomological Society of Ontario, 1902. Pp. 63-67.
26
Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
BIOLOGICAL CONTROL IN ONTARIO 1952-2012:
A SUMMARY OF PUBLICATIONS IN THE
“JOURNAL OF THE ENTOMOLOGICAL SOCIETY OF
ONTARIO”
P.G. MASON
Agriculture and Agri-Food Canada, Research Centre
960 Carling Avenue, Ottawa, ON K1A 0C6
email: peter.mason@agr.gc.ca
J. ent. Soc. Ont. 144: 27-111
Introduction
Biological control involves the manipulation of natural enemies to regulate
populations of pest species. This biologically based approach is key to the successful
management of pest species, and requires a sound understanding of the pest, its associated
organisms and their interactions. A first step is to understand the biology of a target species
which allows determination of such things as number of generations per growing season,
life stages that cause damage, and life stages that are appropriate for control. Knowledge of
the natural enemy community associated with a pest species will provide an indication of the
potential for biological control to suppress and maintain populations below economically
damaging levels. In Ontario, biological control began in 1882 when W. Saunders imported
Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae) from New York state for
release in Ontario gardens to control the Imported Currantworm Nematus ribesii (Scopoli)
(Hymenoptera: Tenthredinidae) (Glen 1962).
The present compilation summarizes the biological control contributions
published in the Annual Report of the Entomological Society of Ontario / Proceedings of
the Entomological Society of Ontario / Journal of the Entomological Society of Ontario
(together, JESO) from 1952-2012 as part of the commemorative activities to celebrate the
150" anniversary of the Entomological Societies of Canada and Ontario. Although most
cover work in Ontario, several (e.g., James 1952; Maxwell and Morgan 1952; Robinson
1952), address studies in other regions. Glen (1956) summarized work in entomology,
including biological control in Canada to 1956 and this should be consulted for information
on studies prior to 1952. It should be noted that studies published in JESO document only
a portion of the work on each species. More comprehensive accounts can be found in the
Biological Control Programmes in Canada series (McLeod et al. 1962; Kelleher et al. 1971;
Kelleher and Hulme 1984; Mason and Huber 2002; Mason and Gillespie 2013).
Several contributions provide general summaries of the knowledge at the time of
their publication. Chant (1957) provided an overview of papers relevant to biological control
Published December 2013
© Her Majesty the Queen in Right of Canada, as represented by the Minister of Agriculture
and Agri-Food Canada.
Di
Mason JESO Volume 144, 2013
presented at the 10" International Congress of Entomology. Cameron (1952) conducted
a review of diseases of insects to 1951 and Cameron (1969) reviewed the problems and
prospects in the use of pathogens for insect control. Putnam (1963) reviewed the biology
and management of codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae). Wallace
and Sullivan (1985) reviewed the status of the white pine weevil, Pissodes strobi (Peck)
(Coleoptera: Curculionidae).
Proverbs (1962) and Van Whervin & Wilde (1970) reported on sterile insect release
for control of codling moth, however this technique falls outside of the definition used
in this summary (1.e., manipulation of natural enemies) as does work with plant extracts
such as that reported for neem, Azadirachta indica A. de Jussieu (Meliaceae), by Lyons et
al. (1996) on the Pine False Webworm, Acantholyda erythrocephala (L.) (Hymenoptera:
Pamphiltidae) and by Li (2000) against Balsam Fir Sawfly, Neodiprion abietis (Harris)
(Hymenoptera: Diprionidae).
In Ontario, more than 75 species have been the subject of studies in which associated
natural enemies have been documented. Introduction of exotic natural enemy species were
implemented for 12 invasive alien arthropods and five exotic weeds. The contributions
published in JESO on these species are varied but can be divided into broad categories,
Pest Life History and Natural Enemy Complexes, General Studies of Natural Enemy
Communities, Natural Enemy Biology, Classical Biological Control of Weeds, Classical
Biological Control of Arthropods, and Inundative Biological Control using Pathogens.
Fundamental to successful biological is correct identification of their natural enemies so
taxonomic studies treating relevant species are therefore also summarized under Natural
Enemy Taxonomy. The approach used here summarizes, under each of the categories
mentioned above, the findings published in JESO for each species studied. The 140 full
length scientific papers, scientific notes and abstracts include those that identified natural
enemies (arthropods and pathogens) associated with a host species and those reporting on
aspects of the biology of natural enemies of pest and beneficial species. A list of the updated
names of natural enemies and known hosts published in JESO from 1952-2012 is provided
in the Appendix.
1. Pest Life History and Natural Enemy Complexes
The development of intensive agriculture brings with it a host of species that exploit
a food source that, grown in large uniform plots, provides one of the optimum conditions that
contributes to exponential population increases. Fundamental to implementing successful
biological control is understanding the biology of target species well and identifying which
natural enemies already present in the system attack the various life stages of the host. In
Ontario, numerous studies have documented the natural enemies of native and non-native
species, usually in response to outbreaks in particular crops or regions. In addition to greater
knowledge of pest biologies these studies have increased knowledge of their natural enemies
present in Ontario. A summary of the findings published in JESO for each species follows.
Apple Maggot, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae) is a native
species that occurs in Ontario and Quebec (Hoffmeister 2002). Monteith (1977; 1978)
studied potential predators of apple maggot, e.g., the sowbug, Porcellio laevis Latreille
(Isopoda: Oniscidae), centipede, Lithobius forficatus (L.) (Lithobiomorpha: Lithobiidae),
earwig, Forficula auricularia L. (Dermaptera: Forficulidae), and beetles, including
28
Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
Calosoma calidum (Fabricius), Harpalus pensylvanicus DeGeer (Coleoptera: Carabidae),
and Staphylinus badipes LeConte (Coleoptera: Staphylinidae), which effectively attacked
and consumed apple maggot larvae. Millipedes, Trachelipus rathkei (Koch) (Polydesmida:
Paradoxosomatidae), attacked puparia. Monteith (1978) also reported on apple maggot
parasitoids, including Diachasma mellea (Gahan), D. lectus Gahan, D. lectoides (Gahan),
D. alloeum (Muesebeck) and D. ferrugineum (Gahan) (Hymenoptera: Braconidae).
Although these parasitoid species survived in wild habitats with apple and Crataegus spp.
(Rosaceae), their numbers were not sufficient to migrate into and reduce apple maggot
populations in managed orchards where even low numbers of this pest could not be tolerated.
Poinar et al. (1978) isolated the potential pathogens, Pseudomonas aeruginosa (Schroeter)
Migula, Bacillus cereus Frankland and Frankland (Bacilliaceae), and Streptococcus sp.
(Streptococcaceae) from larvae and puparia. A nematode, Neoaplectana sp. (Rhabditida:
Steinernematidae) was also associated with puparia. The study suggested that natural
infestation by microorganisms might play an important role in regulating apple maggot
populations.
Armyworm, Mythimna unipuncta (Haworth) (Lepidoptera: Noctuidae), a Nearctic
species, was studied by Goble (1965) during an outbreak in 1964. The nuclear polyhedrosis
virus Betabaculovirus sp. (Baculoviridae) killed 35% of larvae. Parasitoids caused an
additional 25% mortality, particularly two Apanteles spp. (Hymenoptera: Braconidae)
and other Hymenoptera (20% and 3.3% mortality, respectively) as well as Diptera (1%).
Winthemia sp. (Diptera: Tachinidae) was abundant at one site and birds consumed large
numbers of larvae. It was concluded that overall, natural control was of such magnitude that
the population was likely to crash without intervention.
Birch Leaf Edgeminer, Scolioneura betuleti (Klug) (Hymenoptera:
Tenthredinidae), first discovered in Ontario in 1983 near Newmarket, represented a first
record for North America (Nystrom and Evans 1989). They reported 12% parasitism by
three larval parasitoids, Chrysocharis laricinellae (Ratzeburg), Pnigalio minio (Walker),
and Zagrammosoma multilineatum (Ashmead) (Hymenoptera: Eulophidae).
Black Army Cutworm, Actebia fennica (Tauscher) (Lepidoptera: Noctuidae),
a Holarctic species, was studied in black spruce plantations in Newfoundland by West
(1992). Parasitism levels of up to 60% were documented. Tachinomyia panaetius (Walker)
(Diptera: Tachinidae), and Campoletis sp. (Hymenoptera: Ichneumonidae) were reared
from larvae. Gonia sp. (Diptera: Tachinidae), and Enicospilus sp., Ichneumon creperus
Cresson, and Arenetra rufipes Cresson (Hymenoptera: Ichneumonidae) were reared from
pupae. The nematode, Steinernema feltiae (Filipjev) (Rhabiditida: Steinernematidae) also
showed promise as a potential control agent. West (1992) recommended that since only /.
creperus was known to occur in British Columbia, where black army cutworm was also a
problem, relocation of the other spp. may be useful for biological control of A. fennica in
that province.
Cabbage Looper, Trichoplusia ni (Hibner) (Lepidoptera: Noctuidae), is an annual
migrant from the southern USA. Harcourt (1963) determined that 7 ni was significantly
impacted by Copidosoma truncatellum (Dalman) (Hymenoptera: Encyrtidae) but less so
by the polyphagous /toplectis conquisitor (Say), Stenichneumon culpator cincticornis
(Cresson) (Hymenoptera: Ichneumonidae) and Compsilura concinnata (Meigen) (Diptera:
Tachinidae). Polyhedral virus disease frequently killed larvae. Murillo et al. (2012) studied
29
Mason JESO Volume 144, 2013
the larval parasitoids of 7: ni in field tomatoes in southwestern Ontario. Nine primary
parasitoids were reared from 7: ni larvae, including an unidentified Tachinidae, Exeristes
comstockii (Cresson) (Hymenoptera: Ichneumonidae), Copidosoma floridanum (Ashmead)
(Hymenoptera: Encyrtidae), Cofesia marginiventris (Cresson), C. plathypenae (Muesebeck),
Meteorus sp., and Microplitis alaskensis (Ashmead), one unidentified species (Hymenoptera:
Braconidae), and Euplectrus sp. (Hymenoptera: Eulophidae). One hyperparasitoid,
Trichomalopsis viridescens (Walsh) (Hymenoptera: Pteromalidae) was reared from E.
comstockii, the most abundant parasitoid (17.6% and 39.2% parasitism levels in 2005 and
2006, respectively). Although common parasitoids of 7: ni in other parts of North America,
C. floridanum and C. marginiventris occurred in <2% of the host populations in Ontario.
The association of C. plathypenae with T. ni was a new host record.
Corn Aphid, Rhopalosiphum maidis (Fitch) (Hemiptera: Aphididae), is an
important introduced pest of corn. Foot (1974) studied the Coccinellidae (Coleoptera)
community in corn fields in southwestern Ontario. He found that Hippodamia convergens
Guérin-Méneville, H. tredecimpunctata tibilais (Say), and Coleomegilla maculata lengi
Timberlake were the most abundant species. Adalia bipunctata (L.), Cycloneda sanguinea
(L.), H. parenthesis (Say), and Coccinella transversoguttata Faldermann were present but
either not abundant or did not occur in all years. It was concluded that coccinellid numbers
overall were insufficient to control corn aphid as high populations of beetles occurred only
after aphid populations peaked and had damaged the crop.
Diamondback Moth, Plutella xylostella (L.) (Lepidoptera: Plutellidae), first found
in the Ottawa area in 1854, is a global pest of cole crops. Harcourt (1963) determined that
native parasitoids were a major mortality factor, the most important being the larval-prepupal
parasitoid Diadegma insulare (Cresson) (33%), the prepupal-pupal parasitoid Diadromus
subtilicornis (Gravenhorst) (21%) (Hymenoptera: Ichneumonidae), and the larval parasitoid
Microplitis plutellae (Muesebeck) (Hymenoptera: Braconidae). Several species were of lesser
significance including, Oomyzus sokolowskii (Kurdjumov) (Hymenoptera: Eulophidae),
Conura albifrons (Walsh) (Hymenoptera: Chalcididae), Gelis tenellus (Say), Campoletis
sp. (Hymenoptera: Ichneumonidae), Dibrachys microgastri (Bouché), Pteromalus sp.
near phycidis Ashmead, and Trichomalopsis viridescens. According to Harcourt (1963)
predators and diseases did not significantly affect P. xy/ostella populations.
European Red Mite, Panonychus ulmi (Koch) (Trombidiformes: Tetranychidae),
a non-native species, is a serious pest of fruit crops in Canada (Thistlewood et al. 2013).
Herbert (1953) studied the predacious phytoseid mites associated with European red
mite in orchards. More than nine species were collected, including Typhlodromus tilae
Oudemans, 7. rhenanus (Oudemans), 7: pomi (Parrot, Hodgkiss and Shoene), Neoseiulus
fallacis (Garman), T. conspicuous var. herbertae Nesbitt, T: finlandicus (Oudemans), T.
masseei (Nesbitt), Phytoseius macropilis (Banks) and Amblyseius spp. (Trombidiformes:
Phytoseiidae). Abundance and species compositions varied among locations and years.
Populations were denser in the centre of orchards in spring and early summer but increased
at the periphery in midsummer, then decreased as autumn approached. Cadogan and Laing
(1982) surveyed apple orchards in southern Ontario for the European red mite and its predator
Balaustium putnami Smiley (Trombidiformes: Erythraeidae). Two distinct generations of B.
putnami occurred, the 1* generation having an abundance of larvae and the 2" generation
being dominated by nymphs and adults (motile stages). Balaustium putnami coexisted with
30
Biological control in Ontario, 1952—2012 JESO Volume 144, 2013
Phytoseiidae and Stigmaeiidae and fed on both P. u/mi and the twospotted spider mite,
Tetranychus urticae Koch (Trombidiformes: Tetranychidae). Balaustium putnami was also
present in orchards with low volume pesticide application regimes suggesting that spray
regimes and schedules could be designed to preserve natural enemies.
European Skipper, Thymelicus lineola’ (Ochsenheimer) (Lepidoptera:
Hesperiidae), was first collected in 1910 near London, Ontario (Pengelly 1961). He studied
its biology near Bradford, Ontario in 1958. Several native parasitoid species were recovered.
Parasitism of pupae was low at 4.9%, mainly by /toplectis conquisitor. Also reared from
pupae were Pimpla pedalis Cresson and Camposcopus sp. (Hymenoptera: Ichneumonidae).
Larval parasitoids included Meteorus hyphantriae Riley, Rogas sp. and Casinaria sp.
(Hymenoptera: Braconidae). The hyperparasitoid Gelis sp. (Hymenoptera: Ichneumonidae)
was reared from M. hyphantriae. Several Tachinidae were also reared from larvae.
Forest Tent Caterpillar, Malacosoma_ disstria Hibner (Lepidoptera:
Lasiocampidae), a cyclical pest of deciduous trees, was studied by Harmsen and Rose
(1984). They documented differential mortality in wet low-lying and dry higher-ground
habitats. Parasitism by A/leiodes malacosomatos (Mason) (Hymenoptera: Braconidae)
and Phobocampe clisiocampae (Weed) (Hymenoptera: Ichneumonidae) and predation by
unspecified species were lower in the low-lying areas, likely due to limited accessibility
of appropriate sites for pupation and the greater accessibility for predators offered by drier
habitats.
Goldenrod Gall Moth, Epiblema_ scudderiana (Clemens) (Lepidoptera:
Tortricidae) was the subject of a parasitoid survey by Laing and Heraty (1982) who found
the primary parasitoids Apanteles cacoeciae Riley, Macrocentrus pallisteri DeGant, Bassus
binominatus (Muesebeck) (Hymenoptera: Braconidae) and Scambus pterophori Ashmead
(Hymenoptera: Ichneumonidae), and the hyperparasitoid Perilampus fulvicornis Ashmead
(Hymenoptera: Perilampidae), which attacked all the primary parasitoids. Overall parasitism
was 32.4% in 1978-1979, 64.4% in1979—1980, and 76.6% in 1980-1981. Parasitism by M.
pallisteri was the major factor influencing the large annual fluctuations (19.4% in 1978-79,
57.5% in 1979-1980, and 67.5% in 1980-81) in E. scudderiana populations. Perilampus
fulvicornis appeared to be an important regulator of M. pallisteri, preventing it from
drastically reducing E. scudderiana populations.
Horse and deer flies (Diptera: Tabanidae) were the subject of natural enemy
surveys in Churchill, Manitoba by James (1952). The chalcid larval-pupal parasitoid
Diglochis occidentalis (Ashmead) (Hymenoptera: Pteromalidae) was found to parasitize
13.9% of Tabanus spp., including T. affinis Kirby and the T. frontalis-septentrionalis
complex, and 20.8% of Chrysops spp., including C. frigidus Osten-Sacken, and C. furcatus
Walker. Numbers of D. occidentalis that emerged from Tabanus spp. averaged 45.5 while
the smaller Chrysops spp. yielded an average of 16.1.
McDaniel Spider Mite, Zetranychus mcdanieli McGregor, the Apple Mite,
Tetranychus pacificus McGregor, and the Clover Mite, Bryobia praetiosa Koch
(Trombidiformes: Tetranychidae) in Manitoba were the subject of a survey by Robinson
(1952) to document their predators. The following species were collected: Stethocorus
punctum(LeConte), Adalia punctata(L.) (Coleoptera: Coccinellidae), Sti/bus probatus Casey
(Coleoptera: Phalacrididae), Orius insidiosus (Say), Anthocoris musculus (Say) (Hemiptera:
Anthocoridae), Diaphnidia pellucida Uhler, Hyaloides harti Knight, H. vitripennis (Say),
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Plagiognathus obscurus (Uhler) (Hemiptera: Miridae), Nabis ferus (L.) (Hemiptera:
Nabidae), Scolothrips sexmaculatus (Pergande) (Thysanoptera: Thripidae), Aeolothrips
melaleucus Haliday (Thysanoptera: Aelothripidae), Fe/tiella sp. (Diptera: Cecidomyiidae),
Toxomerus geminatus (Say) (Diptera: Syrphidae), Chrysopa carnea (Stephens), C. chi Fitch
(Neuroptera: Chrysopidae), Hemerobius simulans Walker, H. stigmaterus Fitch (Neuroptera:
Hemerobiidae), Zyphlodromus fallacis (Garman), T: longipilus Nesbit (Megostigmata:
Phytoseiidae), and Anystis agilis Banks (Trombidiformes: Anystidae).
Northern Corn Rootworm, Diabrotica barberi Smith and Lawrence (Coleoptera:
Chysomelidae), native to North America, is a minor pest in Ontario. Tyler and Ellis
(1980) studied the importance of ground beetles as its predators. Among the 26 species
collected, Prerostichus melanarius (Illiger), Clivina fossor (L.), Agonum muelleri (Herbst),
Bembidion quadrimaculatum oppositum Say, Poecilus lucublandus (Say), and Harpalus
affinis (Schrank) (Coleoptera: Carabidae) were most numerous. Radioactive labelling trials
indicated that carabids were probably more important as larval than egg predators.
Obliquebanded Leafroller, Choristoneura rosaceana (Harris), the Eyespotted
Bud Moth, Spilonota ocellana (Dennis and Schiffermiiller), and the Pale Apple Budworm,
Pseudexentera mali Freeman (Lepidoptera: Tortricidae) all native species, were present at
all sites surveyed by Hagley and Barber (1992). Although parasitism levels in unmanaged
apple orchards in southern Ontario were low (4—10%), parasitoids reared included 24 species
of Hymenoptera and two species of Diptera. /toplectis conquisitor was the most frequently
reared parasitoid from obliquebanded leafroller and pale apple budworm and Bassus
dimidiator (Nees) (Hymenoptera: Braconidae) was most frequently reared from eyespotted
bud moth. The first records of Colpoclypeus florus (Walker) (Hymenoptera: Eulophidae)
from obliquebanded leafroller and eyespotted bud moth were reported. Colpoclypeus florus
had earlier been introduced from Europe to control the redbanded leafroller (see below).
Highest parasitism levels were found in Coleophora spp. (Lepidopera: Coleophoridae)
(30.2%) and Sparganothis spp. (Lepidoptera: Tortricidae) (62%), primarily due to Scambus
spp. and Orgilus scaber Muesebeck (Hymenoptera: Braconidae) in the former and Triclistus
spp. (Hymenoptera: Ichneumonidae) in the latter.
Pine Shoot Beetle, Zomicus piniperda(L.) (Coleoptera: Curculionidae), a European
species, was first found in the Niagara region in 1993 (Bright 1996). Parasitoids found in
his study included Coeloides pissodis (Ashmead), Spathius sp. (Hymenoptera: Braconidae),
Dinotiscus dendroctoni (Ashmead), Rhopalicus tutela (Walker), Roptrocerus xylophagorum
(Ratzeburg) (Hymenoptera: Pteromalidae), Eupe/mus sp. (Hymenoptera: Eupelmidae) and
Eurytoma sp. (Hymenoptera: Eurytomidae). Predators included Platysoma gracile LeConte
(Coleoptera: Histeridae), Corticeus praetermissus (Fall) (Coleoptera: Tenebrionidae),
Medetera signaticornis (Loew) and M. pinicola Kowarz (Diptera: Dolichopodidae). Most
of the species found are habitat-specific rather than host-specific, thus any bark beetle
encountered under the bark may be a suitable host. A few parasitoid species, e.g., Eupelmus
sp., may be hyperparasitoids. It was concluded that further investigation of the role of native
natural enemies would provide evidence on whether or not there is a need to introduce
exotic natural enemies.
Potato Leafhopper, Empoasca fabae Harris (Hemiptera: Cicadellidae), is a
pest of a variety of field crops such as edible beans, potatoes, alfalfa, peanut and soybean
(Appleton et al. 2004). They concluded that predators and parasitoids were not effective
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Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
regulators of potato leafhopper populations, despite egg parasitism up to 40% by Anagrus
armatus (Ashmead) (Hymenoptera: Mymaridae). Although the fungus Zoophthora radicans
(Brefeld) Batko (Entomophthoraceae) caused epizootics, the narrow environmental
conditions required for this are rare in Ontario; thus it was not considered to be a reliable
control.
Redbanded_ Leafroller, Argyrotaenia velutinana (Walker) (Lepidoptera:
Tortricidae) is a native species that occurs on broad-leaved trees in eastern North America
(Hikichi 1971). In response to increasing outbreaks in apple orchards in Ontario, Hikichi
(1962) studied its mortality factors. Trichogramma minutum parasitized ~2% of the eggs
collected, ~S0% of larvae were infected by a granulovirus and another ~12% of larvae were
parasitized by Phytodietus vulgaris Cresson (Hymenoptera: Ichneumonidae). The study
concluded that disease and drought conditions that reduced foliage quality were the primary
factors contributing to mortality of A. velutinana.
Six-spotted Leafhopper, Macrosteles fascifrons (Stal) (Hemiptera: Cicadellidae),
is an important vector of aster-yellows virus (Miller and De Lyzer 1960). They conducted field
surveys but only a single parasitoid species, Epigonatopus plesius Fenton (Hymenoptera:
Dryinidae) was recovered from adults and levels of parasitism were not considered of
economic importance.
Soybean Aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), native to
eastern Asia, was first reported in Ontario in 2001 (Ragsdale et al. 2004). Bahlai and Sears
(2009) studied the population dynamics of soybean aphid and the predator Harmonia axyridis
(Pallas) (Coleoptera: Coccinellidae) in vineyards in the Niagara region. They found that
high populations of H. axyridis were correlated with substantial numbers of soybean aphid
when aphids occurred early in the season. However, outbreaks of H. axyridis in vineyards
were observed when the numbers of soybean aphid eggs were fewest on overwintering
buckthorn, Rhamnus spp. (Rhamnaceae), plant hosts. The availability of high numbers
of eggs, oviposited by soybean aphid late in the season on the winter host plant, served
to divert H. axyridis from feeding on ripening grapes in vineyards. Thus Bahlai and Sears
(2009) showed that high numbers of aphids in soybean did not result in high numbers of
H. axyridis invading vineyards. They proposed a ‘kick start/distract’ model to explain these
dynamics and provide a basis for integrated management.
Spotted Tentiform Leafminer, Phyllonorycter blancardella_ (Fabricius)
(Lepidoptera: Gracillariidae) an invasive alien pest from the Palaearctic is an important pest
of apples in central Ontario as well as other parts of eastern Canada (Vincent et al. 2013).
Johnson et al. (1977) studied the seasonal occurrence and natural enemies of this pest in
Ontario apple orchards. They reported that the endoparasitic Pholetesor ornigis (Weed)
(Hymenoptera: Braconidae) was the dominant parasitoid (up to 57% parasitism) and
was well-synchronized with the 1“ and 3" host generations. Sympiesis gordius (Walker),
S. sericeicornis (Nees), Pnigalio minio (Walker), P. uroplatae (Howard), Chrysocharis
nepereus (Walker) and Closterocerus sp. (Hymenoptera: Eulophidae) impacted 1‘ and 2"
generation spotted tentiform leafminer, the first three species being most prevalent, although
overall parasitism was at most 24%. Predation was not significant.
Tarnished Plant Bug, Lygus lineolaris (Palisot) (Hemiptera: Miridae) is a
widespread and important pest of vegetable, fruit, greenhouse, and field crops, particularly
those grown for seed (Broadbent et al. 2013). Broadbent et al. (1999) reared five parasitoid
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species, including Leiophron mellipes (Cresson), L. digoneutis (Loan), L. pseudopallipes
(Loan), Leiophron lygivora (Loan), and L. rubricollis (Thomson) (Hymenoptera:
Braconidae). Mason et al. (2011) examined the effect of periodic cutting of alfalfa on
parasitism of tarnished plant bug and alfalfa plant bug, Adelphocoris lineolatus (Goeze)
(Hemiptera: Miridae) by Leiophron spp. Although populations of hosts and parasitoids
declined in cut habitats, they did not go extinct and recolonization by adults sustained
parasitoid populations.
Trefoil Seed Chacid, Bruchophagus platyptera (Walker) (Hymenoptera:
Eurytomidae) is an important pest of alfalfa, clover and trefoil seed crops (Ellis and
Nang’ayo 1992). These authors discovered two parasitoids, Mesopolobus bruchophagi
(Gahan) and Tetrastichus bruchophagi Gahan (Hymenoptera: Pteromalidae) at levels of 8.2
and 11.0%, respectively. Parasitoids were not present in all fields and were more likely to
occur in older fields. They noted that these same species occur elsewhere in North America
where trefoil seed chacid is found.
White Pine Weevil, Pissodis strobi (Peck) (Coleoptera: Curculionidae), native to
North America, is a major pest in pine plantations in most of Canada and the USA (Hulme
and Kenis 2002). Wallace and Sullivan (1985) reviewed its biology, highlighting aspects
that could be exploited to manage the pest. Among major larval and pupal mortality factors
identified were the predator Lonchaea corticis Taylor (Diptera: Lonchaeidae) and the
parasitoids Eurytoma pissodes Girault (Hymenoptera: Eurytomidae) and Dolichotomitus
terebrans nubilipennis (Viereck) (Hymenoptera: Ichneumonidae).
Willow Gall Fly, Rhabdopahaga strobiloides Walsh (Diptera: Cecidomyiidae) was
studied by Judd (1953). In addition to willow gall fly which induces the galls, the inquiline
Dasyneura albovittata Walsh (Diptera: Cecidomyiidae) was reared from these galls, as was
a single female sawfly, Amauronematus sp. (Hymenoptera: Tenthredinidae). Parasitoids
reared from willow gall fly included Copidosoma sp., (Hymenoptera: Encyrtidae), Tridymus
sp. (Hymenoptera: Pteromalidae), and Jorymus cecidomyae (Walker) (Hymenoptera:
Torymidae). Leptacis sp. (Hymenoptera: Platygasteridae), Ceraphron sp. (Hymenoptera:
Ceraphronidae), Tetrastichus sp. (Hymenoptera: Eulophidae) and Jorymus sp. were reared
from cocoons of D.albovittata. Among the remaining parasitoids reared were Adialytus
salicaphis (Fitch) and Aphidius matricariae Haliday (Hymenoptera: Braconidae), known
parasitoids of aphids, and Microgaster hospes Marshall (Hymenoptera: Braconidae) and
Pediobius sp. (Hymenoptera: Eulophidae), parasitoids of Lepidoptera. The hyperparasitoids
Lygocerus sp. (Hymenoptera: Cephronidae) and A//oxysta sp. (Hymenoptera: Alloxystidae)
were reared, probably from A. phorodontis.
2. General Studies of Natural Enemy Communities
Natural enemy surveys that are not pest specific provide a broad perspective of
the complexes present in different habitats. Several studies published in JESO documented
natural enemies associated with particular pests or crop systems, often to evaluate the
impacts of management systems or pesticides on these communities. Other studies appear
to have been opportunistic and documented natural enemies associated with host species
likely encountered fortuitously during field trips focusing on other topics.
Field crop habitats. Ben-Ze’ev and Jaques (1990) surveyed alfalfa fields in
southwestern Ontario for entomopathogens. The invasive Alfalfa Weevil, Hypera postica
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(Gyllenhal) (Coleoptera: Curculionidae) was infected by Erynia phytonomi (Arthur)
Humber, Ben-Ze’ev and Kenneth, Erynia sp. (Entomophthoraceae) and Beauveria bassiana
(Balsamo) Vuillemin (Moniliaceae). Conidiobolus obscurus (Hall and Dunn) Remaudiére
and Keller, C. thromboides Dreschler (Ancylistaceae), Entomophthora planchoniana Cornu,
Erynia neoaphidis Remaudiére and Hennebert (Entomophthoraceae), and Neozygites
fresenii (Thaxter) Remaudiere and Keller (Neozygiotaceae) were associated with a mixed
population of Pea Aphid, Acyrthosiphon pisum (Harris), Black Bean Aphid, Aphis fabae
Scopoli, and the Green Peach Aphid Myzus persicae (Sulzer) (Hemiptera: Aphididae).
Entomophthora muscae (Cohn) Fresen (Entomophthoraceae) complex was associated with
the Seedcorn Maggot, Delia platura (Meigen) (Diptera: Anthomyiidae). Erynia echinospora
(Thaxter) Remaudiere and Keller [or E. dipterigena (Thaxter) Remaudiére and Keller]
was associated with Lauxaniidae (Diptera). Erynia petchii (Ben-Ze’ev and Kenneth) was
associated with the Meadow Spittlebug, Philaenus spumarius (L.) (Hemiptera: Cercopidae)
and Zoophthora radicans (Brefeld) Batko was associated with the Potato Leafhopper,
Empoasca fabae Harris (Hemiptera: Cicadellidae) and Aphididae. The study concluded that
entomopathogens have a role in natural regulation of pest insects and there is potential for
their introduction (e.g., B. bassiana and Erynia spp.) to supplement other biological control
agents to manage H. postica populations.
Orchard habitats. Hagley (1979) studied the effects of insecticides on
natural predator populations in apple, Malus spp. (Rosaceae), orchards. Hippodamia
tridecempunctata tibialis (Say) and Adalia bipunctata (L.) (Coleoptera: Coccinellidae)
were the most abundant predators collected. Phytocoris sp., Deraeocoris fasciolus
Knight and Plagiognathus obscurus (Uhler) (Hemiptera: Miridae) were the main true bug
species encountered, and Chrysopa oculata (Say) (Neuroptera: Chrysopidae), Hemerobius
humulinus (L.) (Neuroptera: Hemerobiidae), Epiodes americanus Wiedemann, Allograpta
obliqua (Say) (Diptera: Syrphidae), Cantharis sp. and Podabrus spp. (Coleoptera:
Cantharidae) commonly occurred. Overall, predator populations were low and insecticide
treatments (phosmet and azinphosmethyl) appeared to reduce eggs and immature stages
of the predators. Most adult predators collected immigrated from outside of treated areas.
Thus, numbers of predators in natural areas was insufficient to provide effective control of
the major pests: codling moth, apple maggot, and Plum Curculio, Conotrachelus nenuphar
(Herbst) (Coleoptera: Curculionidae). Hagley (1979) concluded that augmentation of
predator numbers is required when management practices use insecticides.
Woolhouse and Harmsen (1985) studied the population dynamics of the mite
complex on foliage of a pesticide-free apple orchard. Over a 3-year period, population
dynamics were highly variable but pest species did not reach economic thresholds. Zetzellia
mali (Ewing) (Trombidiformes: Stigmaeiidae) and Phytoseiidae species tracked changes in
prey abundance. Zefzellia mali was more closely linked to eriophyid rust mites, Aculus sp.,
abundance while the Phytoseiidae were linked to tetranychid, i.e., Two-spotted Spider Mite
and European Red Mite, abundance. Ze/ze/lia mali and Phytoseiidae were more abundant
on trees nearer the orchard edge suggesting the acaricide spray programs that focus on
the central parts of an orchard could be less detrimental to predator populations. They
concluded that pest populations tend to be lower, sometimes by an order of magnitude, on
McIntosh and Golden Delicious varieties than on Red Delicious and Empire varieties in a
predator-rich environment.
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Non-crop habitats. Laing and Welch (1963) reported feeding by adults of the
predaceous fly, Dolichopus gratus Loew (Diptera: Dolichopodidae), on larvae of Culex
restuans Theobald (Diptera: Culicidae). Edwards and Pengelly (1966) reported parasitism
of Bombus fervidus (Fabricius) (Hymenoptera: Apidae) by Melittobia chalybii Ashmead
(Hymenoptera: Eulophidae). Loan (1973) reported the first occurrence of parasitism of
adult Notoxus anchora Hentz (Coleoptera: Anthicidae) by Centistes agilis (Cresson)
(Hymenoptera: Braconidae); the level of parasitism was 7%.
3. Natural Enemy Biology
Understanding the biology of natural enemies provides guidance for the
development and conservation of agents to better manage key pests. Since 1952, five JESO
studies described methods to improve rearing of natural enemies useful as biological control
agents while another 18 studied performance of potential biological control agents. Four
other studies described the basic biology of particular natural enemies to better understand
development, behaviours or species interactions. Finally, four studies looked at how
particular pesticides affected the biology of natural enemies.
Rearing of natural enemies. Maybee (1956) described a method for rearing the
exotic parasitoid Basalys tritomus Thomson (Hymenoptera: Diapriidae) on Drosophila
melanogaster Meigen (Diptera: Drosophilidae) in the laboratory. West and DeLong
(1956) studied the biology of and developed a rearing method for Ze/us exsanguis (Stahl)
(Hemiptera: Reduviidae), a generalist predator found in Ontario commonly found feeding
on larvae of the forest tent caterpillar. They successfully reared three generations in the
laboratory; cannibalism appeared to be an important consideration because it affects survival
of newly hatched nymphs.
Corrigan et al. (1990) studied the pupal orientation and emergence success
of Horismenus puttleri (Grissell) (Hymenoptera: Eulophidae), imported from Central
America for biological control of Colorado Potato Beetle, Leptinotarsa decemlineata (Say)
(Coleoptera: Chrysomelidae). Because H. puttleri is unable to overwinter in temperate North
America, mass production for inundative releases was considered as the best option to use
this agent. Location of host eggs on leaf surfaces influenced parasitoid pupal orientation
and emergence. When egg masses faced down (i.e., underside of leaf) 98% of parasitoids
pupated with their head down and 89% of adult H. puttleri emerged successfully. In contrast,
when egg masses faced up (1.e., upper side of leaf) 63% of H. puttleri individuals faced
down (head faced the leaf surface) and 66% of adult parasitoids emerged successfully.
Corrigan and Laing (1992) studied an improved method for producing small,
consistent samples of hosts for presentation to the egg parasitoid, 7richogramma minutum.
They described a new sampling strip to decrease preparation times and reduce damage
to host Ephestia kuehniella Zeller eggs (Lepidoptera: Pyralidae). Corrigan et al. (1994)
studied the feasibility of delaying emergence of 7. minutum and subsequent effects on adult
longevity and fecundity. Adult longevity of individuals reared at 16°C increased in direct
proportion to the length of time they were held as pre-adults at this temperature, compared
to 25°C. However, offspring production was reduced when reared at 16°C, although
reproductive potential was not affected by length of time at 16°C or 12L:12D conditions.
The results indicated that under the conditions studied emergence from E. kuehniella eggs
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Biological control in Ontario, 1952—2012 JESO Volume 144, 2013
was not sufficiently delayed and rearing at lower temperatures (i.e., 16°C) adversely affected
reproductive performance.
Performance of biological control agents. James (1959) studied egg development,
hatching and prey consumption in several habitats by Mantis religiosa L. (Orthoptera:
Mantidae), introduced from Europe in the early 1900s. Egg development differed among
habitats but did not affect hatching. He found that prey abundance, primarily field crickets,
influenced number of and size of egg masses indicating the importance of this prey for
maintaining local populations of M. religiosa.
Loan (1964) studied the biology of Centistes ater (Nees) (Hymenoptera:
Braconidae), an internal parasitoid of adult Sitona spp. (Coleoptera: Curculionidae), as a
biological control agent of S. /inee//us (Bonsdorff) in Canada. In the field, C. excrucians is
well synchronized with the univoltine S. /ineellus. The parasitoids overwintered as mature
larvae in adult weevils, emerging the following spring in late April or May, depending on
temperature, to pupate in the soil. Adult C. ater emerged in late June—early July when the
summer-emerged adult S. /inee/lus were present.
Loan (1965) described the life cycle and development of Leiophron mellipes
(Cresson) in five Miridae (Hemiptera) hosts in southern Ontario. Adults were present
from May to September. Immature stages were found in Labops hirtus Knight (late May
to mid-June, 20% parasitism), Leptopterna dolobrata (L.) (mid-May to end of June, 42%
parasitism), Adelphocoris lineolatus (Goeze) and A. rapidus Say (June, 49% and 60%,
respectively), and Lygus lineolaris (Hemiptera: Miridae) (June-July, 46% parasitism, and
August-September, 12% parasitism). A single generation of L. me/lipes occurred in each host
species, although each of the two distinct generations of L. /ineolaris were parasitized.
Griffiths (1972) studied the discrimination ability of the parasitoid Pleolophus
basizonus (Gravenhorst) (Hymenoptera: Ichneumonidae) introduced from 1939-1949 for
biological control of the invasive European Pine Sawfly, Neodiprion sertifer (Geoffroy)
(Hymenoptera: Diprionidae). Although unable to detect hosts containing eggs of conspecifics,
P. basizonus were able to detect hosts containing later developmental stages. Pleolophus
basizonus was also recovered from two other introduced and seven native sawflies.
Reid and Harmsen (1975) studied the biology of Trihabda borealis Blake
(Coleoptera: Chrysomelidae) on goldenrod, Solidago canadensis (Kirby) (Asteraceae).
They determined 7: borealis is of major importance as a phytophage on S. canadensis in
southeastern Ontario, although serious defoliation was rare.
Ramey (1990) studied the host identification and oviposition behaviour of Eurytoma
obtusiventris Gahan (Hymenoptera: Eurytomidae), a parasitoid of Eurosta solidaginis
(Fitch) (Diptera: Tephritidae) that live in galls of goldenrod (Solidago spp.). Female E.
obtusiventris preferred stems of Solidago altissima L. infested with E. solidaginis but also
explored S. altissima stems without fly larvae, although females only oviposited in plants
containing host larvae. He also showed that E. obtusiventris females prefer S. altissima
infested plants over infested S. canadensis plants.
George (1979) studied the potential of Dugesia tigrina (Girard) (Tricladida:
Duegesiidae) for control of mosquitoes in Ontario. Field tests showed that D. tigrina
reduced populations of Culex restuans and C. pipiens L. (Diptera: Culicidae) by 17 times
(4/dip versus 69/dip in control treatments). Low oxygen levels and toxins such as turpentine
and paint were important mortality factors of D. tigrina in catch basins (George 1984).
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Loan (1982) developed a field technique to study the interaction of the fungi
Zoophthora spp. (Entomophthoraceae) and the parasitoid Perilitus colesi (Drea)
(Hymenoptera: Braconidae) both of which attack larvae of the alfalfa weevil. Foliar
applications of the fungicide captafol protected weevil larvae from attack by Zoophthora
spp. The study confirmed earlier findings that peak attack by M. co/esi occurred after disease
epizootics caused by Zoophthora spp. began to subside.
Bolter and Laing (1984) studied competitive interactions between Diadegma
insulare and Microplitis plutellae for larvae of diamondback moth. Development of both
parasitoids was synchronized with that of the host. At 23°C average fecundity was 814
eggs per female for D. insulare and 316 eggs per female for M. plutellae. Degree-day
development from egg to adult was 282 above 6.6°C for D. insulare and 218 above 9.2°C
for M. plutellae. Diadegma insulare avoided superparasitism and multiple parasitism of
larvae already parasitized by M. plutellae. In contrast, M. plutellae avoided superparasitism
but could not detect eggs of D. insulare for at least 12 h after they were oviposited in the
host. When eggs of both species were oviposited at the same time, 1‘ instar M. plutellae was
intrinsically superior to 1*' instar D. insulare. However, 2™ and 3" instar D. insulare were
superior to 1“ instar M. plutellae.
Clements (1989) studied the role of the stigmaeid mite, Z. mali in orchards.
Zetzellia mali fed on the European red mite and the Apple Rust Mite, Aculus schlechtendali
(Nalepa) (Trombidiformes: Eriophyidae) but did not interfere with the phytoseiid mite
Typhlodromus caudiglans (Schuster) (Mesostigmata: Phytosetidae), either by intraguild
predation or competition.
Whistlecraft and Lepard (1989) studied the effect of flooding on the survival
of the Onion Maggot, Delia antiqua (Meigen) (Diptera: Anthomyiidae), and two of its
parasitoids, Aphaereta pallipes (Say) (Hymenoptera: Braconidae) and Aleochara bilineata
Gyllenhal (Coleoptera: Staphylinidae). Survival of A. pallipes was greater than or equal
to that of its host while survival of A. bilineata was less, even at temperatures below the
developmental threshold (1°C). This suggested that flooding of fields to control D. antiqua
may lead to elimination of 4. bilineata populations. Whitfield et al. (1981) developed a
computer model to simulate the interaction between onion maggot and A. pallipes, a larval
parasitoid. The model determined that 4. pallipes reduced 2™ and 3" generation maggot
populations, resulting in a 70% profit gain. As well, the model provided guidance on when
spray applications would least affect parasitoids.
Wang and Laing (1989) studied the reproductive biology of the introduced
Ageniaspis testaceipes (Ratzeburg) (Hymenoptera: Encyrtidae), an egg-larval parasitoid,
and its effect on the spotted tentiform leafminer. Potential fecundity of H. testaceipes
was 25 eggs per female and an average of 9.1+3.4 broods were produced over an average
lifespan of 7.5+2.7 days. Although newly oviposited host eggs were preferred, eggs up to 5
days old were successfully parasitized and parasitoid development took 35—37 days at 25°C.
Development of parasitized H. testaceipes was delayed and these individuals were larger
than unparasitized individuals. The longer feeding period and larger size of parasitized
spotted tentiform leafminer larvae suggest that H. testaceipes may consume more foliage,
however, this may also increase the size and/or number of parasitoids. They concluded that
the attribute that female H/. testaceipes may oviposit into host eggs of any age provides a
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larger window of opportunity for oviposition, facilitating synchronization with its host and
improve efforts to establish H. testaceipes in North America.
Song (1990) studied the potential for Gelis tenellus, a hyperparasitoid, to influence
parasitism of gypsy moth by Cotesia melanoscela (Ratzeburg) (Hymenoptera: Braconidae).
Gelis tenellus produced significantly more eggs when hosts were available on a daily
basis versus every third day. Nealis and Bourchier (1995) compared the vulnerability to
hyperparasitism of different European and Asian strains of Cotesia melanoscela, a biological
control agent of gypsy moth. Rates of predation and hyperparasitism were not related
to cocoon morphology but were dependent on length of time cocoons were exposed to
hyperparasitism in the field. The nondiapause characteristics of the Asian strain decreased
its exposure time and therefore reduced vulnerability to hyperparasitism. Thus, inundative
releases of nondiapause strains early in the season were likely to minimize exposure of
C. melanoscela to hyperparasitism, which currently is 95% over the summer. They also
concluded that diapause of already established local strains of C. melanoscela could be
manipulated by varying photoperiod during larval development, thus release of additional
exotic strains would not be required.
Villaneuva and Harmsen (1996) studied the ecological interactions of tarsonemid
mites inapple orchards. Dendroptus n. sp. near suskii Sharonov and Livshitz(Trombidiformes:
Tarsonemidae) was identified as a predator of apple rust mite and contributed to the mid-
summer decrease of this pest.
Jones et al. (2006) studied the influence of greenhouse microclimate on predation
of Western Flower Thrips, Franklinella occidentalis (Pergande) (Thysanoptera: Thripidae)
by Neoseiulus cucumeris (Oudemans) (Mesostigmata: Phytoseiidae). Leaf temperature
was positively correlated with predation and oviposition by N. cucumeris, suggesting that
seasonal adjustments in release of this biological control agent could be made.
Development, behaviour and species interactions. Vander Hoek (1971)
described the larval instars of Aphidius nigripes Ashmead (Hymenoptera: Braconidae), a
common parasitoid of the pea aphid, Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae).
Five instars were documented based on changes in cuticular structure observed at 24 h
intervals.
Bennett (2004) studied the host location behaviour of Pe/ecinus polyturator (Drury)
(Hymenoptera: Pelecinidae) a common endoparasitoid of june beetles, Phyllophaga spp.
(Coleoptera: Scarabeidae). Host location consisted of wandering on the surface until the
antennae ceased moving and the distal abdominal segments appeared to touch the surface.
Then a series of movements would push the distal segment into the soil, penetrating up to 5
cm. The procedure lasted about 145 seconds.
Macfarlane and Pengelly (1978) studied Brachioma spp. (Diptera: Sarchophagidae),
and the eulophid Melittobia chalybii Ashmead (Hymenoptera: Eulophidae), parasites of
the brood of Bombus spp. in southern Ontario. They reported Brachioma setosa Coquillett
as a parasite of Bombus for the first time and found that 2—3 generations occurred each
season. As well, M. chalybii attacked both Bombus spp. and B. setosa. They found that these
parasites attacked larvae of Bombus spp. and infested colonies had fewer workers and died
out more quickly than unparasitized colonies.
Wright and Laing (1979) reported on the effects of temperature on development,
adult longevity and fecundity of Coleomegilla maculata lengi Timberlake (Coleoptera:
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Coccinellidae) and its parasitoid Dinocampus coccinellae (Shrank) (Hymenoptera:
Braconidae). A total of 198.8 degree-days (above 13.8°C) were required for development of
C. maculata lengi from egg to adult while 180.5 degree-days (above 11.2°C) were required
for D. coccinellae development. Coleomegilla maculata lengi produced an average of 191.5
eggs per female and longevity averaged 82.3 days. Dinocampus coccinellae survived for 5
days when continuously exposed to hosts and produced an average of 66.8 eggs per female
at 25°C. Earlier studies had estimated potential fecundity of D. coccinellae at 200-400 eggs
per female, thus it appeared that realized fecundity was limited by the ability of females to
find hosts. Wright (1979) observed the copulatory behaviour of C. maculata lengi. The male
mounted females from behind and assumed the dorsal position as is normal for braconids.
Copulation lasted for 18-20 min, considerable longer than the <1 min known for other
braconids.
Effects of pesticides on natural enemies. Robinson (1953) described the biology
of Stethorus punctum (LeConte) (Coleoptera: Carabidae) and determined that DDT and
methoxychlor were lethal to adults, killing 47.1 and 60%, respectively, in laboratory
experiments. Fisher (1988) reported on the effects of pesticides on Pholetesor ornigis
(Weed) and P. pedias (Nixon) (Hymenoptera: Braconidae), parasitoids of the spotted
tentiform leafminer. The number of days to 50% mortality (LT50) of Pholetesor pedias was
higher than for P. ornigis when exposed to azinphosmethyl and permethrin.
Hagley and Laing (1989) studied the effect of pesticides on parasitism by 7. minutum
and 7: pretiosum Riley (Hymenoptera: Trichogrammatidae) of eggs of codling moth. The
insecticides azinphosmethyl, difluibenzuron, permethrin, and methomyl were toxic, as was
the acaricide cyhexatin. Triflumuron, a lower rate of diflubenzuron (1/3 of recommended
dose), and the fungicides captan, dodine and polyram did not affect parasitism.
Wang and Laing (1990) studied the toxicity of methomyl, permethrin,
azinphosmethyl and phosmet to adult Ageniaspis testaceipes, an introduced egg-larval
parasitoid of the spotted tentiform leafminer. At the time, these insecticides were used to
control spotted tentiform leafminer, the plum curculio, codling moth and apple maggot in
Ontario orchards. They concluded that understanding tolerance levels of A. testaceipes to
pesticides used is essential for integrating this biological control agent into management
programmes. For example, methomyl and permethrin residues caused higher mortality of
A. testaceipes than azinphosmethy! and phosmet, although responses of individual females
were highly variable to the latter two products.
4. Classical Biological Control of Weeds
Studies on classical biological control of weeds reported in JESO have been few and
none are comprehensive. A great deal of the work in this area relevant to Ontario has been
published elsewhere (e.g., The Canadian Entomologist). In JESO there are publications on
various aspects of exotic phytophagous-feeding insects introduced for biological control of
five non-native weed species (Table 1). Three species are on the Noxious Weeds in Ontario
list (Anonymous 2013) and all are treated in the Ontario Weeds guide (Alex, 1998). The
publications summarized here report on the status, at the time of publication, of introduced
species.
Canada Thistle, Cirsium arvense (L.) Scopoli (Asteraceae), is a noxious and
widespread weed in Ontario, most abundant in southern areas (Moore, 1975). Urophora
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cardui L. (Diptera: Tephritidae) was introduced for its biological control (Laing 1978). Three
years after initial releases in 1975, 40% of host plants around the release site contained galls
of U. cardui and the agent had spread to plants several hundred meters from the release
site.
Leafy Spurge, Euphorbia esula L. (Euphorbiaceae), is a noxious and widespread
weed in Ontario (Best etal. 1980). LeSage (1996a) reported that populations of the introduced
biological control agent, Aphthona nigriscutis Foudras (Coleoptera: Chrysomelidae)
increased significantly in 1994 and 1995 but did not damage leafy spurge. The survey also
yielded specimens of A. flava Guillebeau, a related exotic species that had not been approved
for release, suggesting that some individuals in the released population were misidentified.
Nodding Thistle, Carduus nutans L. (Asteraceae), is a noxious and widespread
weed in Ontario where it is most abundant, although it occurs throughout Canada
(Desrochers et al. 1988). Laing and Heels (1979) reported that three years after 1975
releases, Rhinocyllus conicus Frélich (Coleoptera: Curculionidae) was well established
around Guelph. Infestation levels up to 95% (24-95%) were recorded. Thistle seed heads
with 7+ pupal cells of R. conicus produced significantly reduced amounts of seed than those
seed heads with 0-6 pupal cells.
Purple Loosestrife, Lythrum salicaria L. (Lythraceae), is highly abundant in the
Great Lakes Basin and along the St. Lawrence River (Mal et al. 1992). Corrigan et al.
(1998) conducted a study on potential non-target feeding by Neogalerucella calmariensis
(L.) (Coleoptera: Chrysomelidae) introduced for biological control of this invasive plant.
The study was initiated based on the field observations of G. calmariensis feeding on
cuttings of Swamp Loosestrife, Decodon verticillatus (L.) Elliott, and egg masses on
Winged Loosestrife, Lythrum alatum Pursh (Lythraceae), at the Ontario Royal Botanical
Garden where large populations of NV. calmariensis were present. Monitoring of all three
plant species through two generations of the beetle revealed that L. salicaria plants sustained
moderate to complete defoliation in all areas monitored. Several D. verticillatus and L.
alatum plants were slightly damaged by N. calmariensis feeding and about 15 egg masses
were found when several hundred of these non-target plants were examined. No late instar
larvae were found on either D. verticillatus or L. alatum. The results suggested that the
minimal feeding and few egg masses represent a ‘spill-over’ effect that occurred when large
numbers of N. calmariensis were dispersing from locations where L. salicaria populations
had been significantly reduced.
St. John’s Wort, Hypericum perforatum L. (Hyperiaceae) is found in the Great
Lakes-St. Lawrence regions of Ontario (Crompton et al. 1988). LeSage (1996b) reported on
the presence in the Gatineau area of Quebec of Chrysolina hyperici (Férster) (Coleoptera:
Chrysomelidae), introduced for biological control of St. John’s wort. The agent had dispersed
145 km from the release site in Belleville, Ontario, at an estimated rate of 6 km per year.
5. Classical Biological Control of Arthropods
Introduction of exotic species for the biological control of arthropods has been
reported in JESO for 15 invasive species (Table 2). All but one of these papers (Maxwell
and Morgan 1952) treated pests of agriculture crops or trees in Ontario. The JESO studies
summarized here for each target species report on the status of introduced biological
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control agents, i.e., whether established or not, and document native natural enemy species
associated with the targets at the time of publication.
Alfalfa Blotch Leafminer, 4gromyza frontella (Rondani) (Diptera: Agromyzidae),
invaded Ontario in the mid 1970s. Coote and Ellis (1987) studied the parasitoids of
alfalfa blotch leafminer near Guelph in 1983-1984. Four parasitoids, Diglyphus begini
(Ashmead), D. intermedius (Girault), D. pulchripes (Crawford) and Pnigalio maculipes
(Crawford) (Hymenoptera: Eulophidae) were reared from larvae. Cyrtogaster vulgaris
Walker (Hymenoptera: Pteromalidae) and Chrysocharis giraulti Yoshimoto (Hymenoptera:
Eulophidae) were reared from pupae. Overall parasitism was low, averaging 3.4% due in
part to poor synchrony of the parasitoids with the host. One additional species, Diglyphus
isaea (Walker) (Hymenoptera: Eulophidae) was collected from alfalfa plants. Parasitoids
emerged later in the spring than alfalfa blotch leafminer, thus parasitizing only 2™ and 3"
generation hosts. Diglyphus intermedius was the most abundant of the larval parasitoids and
the three Diglyphus spp. accounted for 75% of the parasitoids reared from hosts. Diglyphus
isaea and C. vulgaris are exotic parasitoids and were reported for the first time in Ontario
and in association with alfalfa blotch leafminer. All but one of the pupal parasitoids was C.
vulgaris and parasitism levels were low, averaging <1% but were highest at 3.3% in the 3
generation, although sampling included only the few pupae on plants and not those in soil
where alfalfa blotch leafminer normally pupates. It was concluded that the existing parasitoid
complex was unlikely to maintain alfalfa blotch leafminer below economic thresholds and
exotic species already established in the USA would be suitable for introduction. Harcourt
et al. (1987) reported that the European larval-pupal endoparasitoid Dacnusa dryas (Nixon)
(Hymenoptera: Braconidae), first released in1979 near Ottawa, became well established in
most counties of southern Ontario, with rates of attack averaging 84% (65-95%). Dispersal
from nursery plots and natural spread from release sites and life table data indicated that
alfalfa blotch leafminer populations declined less than three years after release of D.
dryas.
Alfalfa Weevil, Hypera postica (Gyllenhal) (Coleoptera: Curculionidae), of
European origin, was first reported in the Great Lakes region in the early 1960s. Abu and
Ellis (1976) studied Bathyplectes curculionis (Thomson) (Hymenoptera: Ichneumonidae) a
larval parasitoid of alfalfa weevil and found that although spring emergence of B. curculionis
was synchronized with that of alfalfa weevil larvae, parasitism levels were low early in
the season (6.3—33.3%) when host populations were highest, increasing later in the season
(60-68%) when host populations were declining. High rates of diapause in 1* generation
parasitoid larvae were thought to be responsible for the lower initial parasitism. Several
hyperparasitoids, Gelis sp., Trichomalopsis viridescens, Pteromalus sp. and Eupelmella
vesicularis (Retzius) (Hymenoptera: Eupelmidae) were reared from 24% of B. cuculionis.
Harcourt et al. (1980) studied the distribution of the European exotic Perilitus aethiops
Nees (Hymenoptera: Braconidae), a parasitoid that attacks adults of alfalfa weevil. First
released in Ontario in 1970-1971, P. aethiops became widely established in southern Ontario
by 1979, where parasitism levels of the spring generation of alfalfa weevil averaged 60%
(13-92%). Harcourt et al. (1982) conducted a survey for Perilitus colesei (Drea), a larval
parasitoid introduced in 1970 and found that P. colesi was present in 39 of 41 counties, with
parasitism levels averaging 13% (152%). Although two fungal pathogens also attack alfalfa
weevil larvae, M. colesi emerges from cocoons in late May or early June and attacks larger
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host larvae, likely after epizootics have subsided and therefore it is able to coexist with the
disease agents. The widespread distribution of M. colesi is probably the result of dispersal
from the USA into southwestern Ontario and dispersal from the release site in Prince Edward
County in eastern Ontario. Harcourt and Ellis (1992) determined that the larval parasitoid
Bathyplectes anurus (Thomson) (Hymenoptera: Ichneumonidae), introduced in 1970, had
become widespread in southern Ontario and had displaced B. curculionis as the main larval
parasitoid of H. postica. Abundance of this parasitoid was influenced by the fungal! pathogen
Zoophthora phytonomi (Arthur) Batko (Entomophthoraceae), which dominated during wet
periods while B. anurus increased during successive dry springs.
Carrot Rust Fly, Psi/la rosae (Fabricius) (Diptera: Psilidae) was introduced
around 1885. Releases of Chorebus posticus (Haliday) (Hymenoptera: Braconidae), a larval
parasitoid, and Basalys tritoma, a pupal parasitoid, were made from 1949-1953 in Ontario,
British Columbia, and Prince Edward Island (Maybee 1954). Although recoveries were
made in the year of release neither C. posticus nor B. tritoma were collected the following
winter.
Cereal Leaf Beetle, Oulema melanopus (L.), (Coleoptera: Chrysomelidae), was
first found in southwestern Ontario in 1965 and became established in 1967 (McClanahan
et al. 1968; Bierne 1971). McClanahan et a/. (1968) reported that while no natural enemies
were present in southwestern Ontario during the study, predators, parasitoids and diseases
had been reported elsewhere in parts of North America invaded by this European pest.
Ellis et al. (1979) reported that Tetrastichus julis (Walker) (Hymenoptera: Eulophidae),
introduced into southern Ontario in 1974 as a biological control agent for cereal leaf beetle,
had by 1977 expanded its range into the area north of Lake Huron and parasitism levels
from 19-90% were documented. In areas where T. ju/is had been established since 1976,
parasitism averaged 65%, indicating that it can maintain populations even when host
densities are low. This successful biological control continued until an outbreak occurred
in the central tobacco growing area of Ontario (Ellis et al. 1989). In a 1987 survey they
reared a single Anaphes sp. from eggs of cereal leaf beetle and parasitism by T. julis was
nil, despite high levels (~75%) of parasitism in other parts of the province. It was concluded
that tillage, which kills 95% of overwintering 7: julis, probably accounted for the absence
of this agent in areas where crop rotations were practiced.
Codling Moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), of southeastern
European origin, was present in Ontario by 1858-1860 and a major apple pest by 1868
(Putnam 1963). In a review of the status of C. pomonella Putnam (1963) included what was
known at the time about natural enemies. 7richogramma minutum was the only egg parasitoid
associated with C. pomonella, while larval parasitoids included Scambus pterophori
Ashmead (Hymenoptera: Ichneumonidae), Dibrachys microgastri (Bouché) (Hymenoptera:
Pteromalidae), Hymenochaonia delicata (Cresson), Macrocentrus ancylivora Rohwer, M.
instabilis Muesebeck, Phanerotoma fasciata Provancher (Hymenoptera: Braconidae),
Mastrus carpocapsae (Cushman), Temelucha minor (Cushman), Cryptus albitarsis
(Cresson), Glypta sp., Aritranis sp. (Hymenoptera: Ichneumonidae), and the adventive
Ascogaster quadridentata Wesmael (Hymenoptera: Braconidae). Dibrachys microgastri
was also found to be a hyperparasitoid of A. quadridentata as were Perilampus fulvicornis
Ashmead, P. tristis Mayr and Perilampus sp. (Hymenoptera: Perilampidae), sometimes
at levels of 72%. Pupal parasitism was negligible but included D. microgastri, Eupelmus
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cyaniceps Ashmead (Hymenoptera: Eupelmidae), Pimpla annulipes Brullé, Itoplectis
conquisitor, and Eurytoma sp. a hyperparasitoid. Eupelmus cyaniceps also parasitized the
larval parasitoids Macrocentrus spp. and the pupal parasitoid P. annulipes. Liotryphon
caudatus (Ratzeburg) and Nippocryptus vittatorius (Jurine) (Hymenoptera: Ichneumonidae)
were introduced from France from 1941—1945 but failed to establish. E/odia tragica (Meigen)
(Diptera: Tachinidae) and Pristomerus vulnerator (Panzer) (Hymenoptera: Ichneumonidae)
were introduced from England in 1943-1944. Ascogaster quadridentata, L. caudatus and
N. vittatorius were introduced into British Columbia but only 4. quadridentata became
established. The most important insect predators were the trogositid borer Tenebroides
corticalis Melsheimer (Coleoptera: Trogossitidae), Chrysopa carnea (Stephens) and C.
rufilabris Burmeister (Neuroptera: Chrysopidae), the egg feeding Haplothrips faurei Hood
and Leptothrips mali (Fitch) (Thysanoptera: Phlaeothripidae), and the mite Anystis agilis
Banks (Trombidiformes: Anystidae). Downy, Dendrocopos pubescens (L.), and hairy, D.
villosus (L.) woodpeckers (Piciformes: Picidae) were important predators of codling moth.
Several diseases have been isolated from codling moth, including Bacillus cereus from
the Niagara Penninsula, Beauveria bassiana from Nova Scotia, and Hirsutella subulata
Petch (Ophiocordycipitaceae) from the USA. Mermis sp. and Neoaplectana n. sp. (DD136)
(Mermithidae) nematodes were also found infecting codling moth. Hagley (1970) studied
codling moth to assess the importance of biotic and abiotic factors in regulating populations.
He determined that disease (34.4—65.1%) and parasitism (31.9-80%) could be significant,
although they were not uniform across orchards. Predation by birds was as high as 90%.
Hagley (1987) surveyed the Trichogramma spp. in apple orchards in southern Ontario after
inundative releases of 7. pretiosum and T. minutum. Only T: pretiosum was recovered from
sentinel codling moth eggs set out in 1982 and 1983. Parasitism ranged from 2.2—11.9% and
parasitoids were recovered in both unsprayed and sprayed orchards. In 1984, 7. minutum was
the only species recovered in unsprayed orchards. The results indicated that Trichogramma
spp. migrated into orchards from alternative hosts and occurred in low numbers early in the
season. This and overall low natural parasitism suggested that augmentative releases and
management of parasitoid populations could improve the success of biological control of
coding moth.
Cranberry Fruitworm, Acrobasis vaccinii Riley (Lepidoptera: Pyralidae), in New
Brunswick was the subject of a study showing that eggs were parasitized by Phanerotoma
franklini Gahan (Hymenoptera: Braconidae) and that Cryptus albitarsus albitarsus (Cresson)
(Hymenoptera: Ichneumonidae) emerged from overwintered larvae (Maxwell and Morgan
1952):
European Pine Shoot Moth, Rhyacionia buoliana (Denis and Schiffermiiller)
(Lepidoptera: Tortricidae), was introduced adventively from the USA into Ontario near
Windsor in 1925 (Pointing and Green 1962). Coppel and Arthur (1954) provided an update
on parasitoids introduced in Ontario to control it. From1928—1953 nine species, including
Campoplex difformis (Gmelin), Sinophorus turionum (Ratzeburg), Temeluca interruptor
(Gravenhorst), Exeristes ruficollis (Gravenhorst), Pimpla turionellae (L.), an unidentified
Pimpla sp. (Hymenoptera: Ichneumonidae), Orgilus obscurator (Nees) (Hymeoptera:
Braconidae), Copidosoma filicorne (Dalmen) (Hymenoptera: Encyrtidae), and Baryscapus
turionum (Hartig) (Hymenoptera: Eulophidae) were released. Among the species recovered
during post-release surveys, C. interruptor and O. obscurator accounted for more than 2/3
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of all parasitoids. Overall parasitism was |.96—10.86% and the native species, Campoplex
sp., /toplectis conquisitor, Itoplectis sp., Scambus hispae (Harris) (Hymenoptera:
Ichneumonidae), Eurytoma appendigaster (Swederus) (Hymenoptera: Eurytomidae),
Habrocytus sp. (Hymenoptera: Pteromalidae), Hyssopus thymus Girault (Hymenoptera:
Eulophidae), the tachinid Exeristes comstockii, and an undetermined species were reared
from European pine shoot moth. Individuals of the introduced 7. turionum, C. difformis,
and P. turionellae were also reared; however, no recoveries of C. geniculatum, C. rufifemur
and £. ruficollis were made during the survey. Pointing and Green (1962) determined that
the 21 native and introduced parasitoids had negligible impact on the host and only four, O.
obscurator, T. interruptor, P. turionellae and B. turionum, of the 13 species introduced had
established. Among these, O. obscurator was the most abundant in Ontario and Quebec.
Gypsy Moth, Lymantria dispar (L.) (Lepidoptera: Erebidae), was first reported in
Ontario on Wolf Island near Kingston in 1969, spreading to the mainland and throughout
eastern Ontario by 1971 (Griffiths 1977). A survey in 1974-1975 by Griffiths (1977)
reported that among the four parasitoid species recovered, Cotesia melanoscela, Compsilura
concinnata and Parasetigena agilis (Robineau-Desvoidy) (Diptera: Tachinidae) are
exotic introductions, none of which targeted gypsy moth, while Pimpla pedalis Cresson
(Hymenoptera: Ichneumonidae) is native. Also reported was the native Gelis tenellus as a
hyperparasitoid of C. melanoscela. Cotesia melanoscela was the most widely distributed
while C. concinnata was the most abundant. Nealis and Quednau (1996) documented
releases and overwintering survival of the European Ceranthia samarensis (Villeneuve)
(Diptera: Tachinidae) introduced for biological control of gypsy moth. Releases of gravid
female adults, parasitized larvae and parasitized pupae were made from 1991—1996. In each
year of release, evidence of successful parasitism by field-released females was observed.
All progeny retrieved were in diapause and overwintering studies indicated that survival
of pharate adults was expected to be high. Because of the low fecundity of C. samarensis
ongoing monitoring was recommended to determine if successful establishment had
occurred.
Imported Cabbageworm, Pieris rapae (L.) (Lepidoptera: Pieridae), was first
reported in eastern Ontario in 1871 and throughout southwestern Ontario by 1876 (Harcourt
1963). Parasitoids are important mortality factors of P- rapae (Harcourt 1963), principally
Cotesia glomerata (L.) (Hymenoptera: Braconidae) which attacks larvae. Later instars
are attacked by Phryxe vulgaris (Fallén) (Diptera: Tachinidae) and pupae are attacked by
Pteromalus puparum (L.) (Hymenoptera: Pteromalidae). Generalist species associated with
P. rapae include C. concinnata, Helicobia rapax (Walker) (Diptera: Sarcophagidae) and
Madremyia saundersii (Williston) (Diptera: Tachinidae). Although invertebrate predators
and birds are present they did not have a significant impact, unlike granulosis virus which
caused up to 94% mortality. Corrigan (1983) conducted a survey for Cotesia rubecula
(Marshall) (Hymenoptera: Braconidae) introduced from British Columbia as a biological
control agent. Recovery of C. rubecula near Ottawa 10 years after its release indicated that
this agent had established in eastern Canada and was tolerant of winter conditions. Up to
1982, no progeny of C. rubecula released near Guelph and Harrow in 1978-1979 were
recovered in the years after release and it was thought that C. rubecula had been negatively
impacted by hyperparasitoids. Carter and Laing (1997) reported on recoveries of a Chinese
strain of C. rubecula released in 1991-1992. Three years after releases C. rubecula was
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found in the release area, although the hyperparasitoids Catolaccus sp. (Hymenoptera:
Pteromalidae), Mesochorus vittator (Zetterstedt) (Hymenoptera: Ichneumonidae) and
Baryscapus galactopus (Ratzeburg) (Hymenoptera: Eulophidae) were reared from C.
rubecula cocoons. Parasitism levels ranged from 15—21%.
Larch Casebearer, Coleophora laricella (Hiibner) (Lepidoptera: Coleophoridae),
was introduced into Ontario from 1935-1941. Graham (1958) studied the effectiveness of
parasitoids of larch casebearer and confirmed establishment of Chrysocharis laricinellae
(Ratzeburg) (Hymenoptera: Eulophidae) and Agathis pumila (Ratzeburg) (Hymenoptera:
Braconidae). Parasitism by the widely established 4. pumila ranged from 41% south of
43° north latitude to 67% between 44—4S° north and it was present in areas of low and
discontinuous host populations. In contrast, C. /aricinellae had spread only 42 miles from
the release point and spread appeared to be dependent on high host populations.
Oriental Fruit Moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae),
was first reported in Ontario in 1925 (McLeod 1962). Boyce and Dustan (1954) compared
parasitism of G. molesta in a young peach orchard and a mature orchard, before and after
pesticides (DDT and parathion) came into use. The most prevalent parasitoids recovered were
the introduced Macrocentrus ancylivora, and the native Hymenochaonia delicata, Enytus
obliteratus (Cresson), Glypta rufiscutellaris Cresson (Hymenoptera: Ichneumonidae) and
Temelucha minor. Overall, M. ancylivora populations increased since insecticide use began
while those of G. rufiscutellaris and H. obliteratus decreased. Hymenochaonia delicata
a common parasite of the ragweed borer, Epiblema strenuana (Walker) (Lepidoptera:
Tortricidae), continued to be abundant. Dustan and Boyce (1966) assessed parasitism
of G. molesta from 1956-1965 and found that average parasitism by M. ancylivora was
43.2-64.5% in 1‘ and 2™ generations, respectively, in the Niagara region and 10.3 and
12.4% in the Essex county region. Parasitism of 2"4 generation oriental fruit moth by G.
rufiscutellaris was 1.4% in Niagara and 28.6% in Essex. Among the other parasitoids,
T. minor was reared from larvae of the I“ and 2™ generations, and Enytus obliteratus
(Cresson) (Hymenoptera: Ichneumonidae) and H. delicata were reared from larvae of the
I generation. The abundance of M. ancylivora in the Niagara region was attributed to the
presence of strawberry plantings which support populations of Ancylis comptana (Frolich)
(Lepidoptera: Tortricidae), an alternate host of M. ancylivora. Phillips (1969) found that M.
ancylivora was the most abundant of eight parasitoids reared from Oriental fruit moth in pear
orchards. Between 40 and 50% of 1‘ and 2™ generation fruit moth larvae were parasitized
from 1964-1966. In 1967, parasitism of 1‘ and 2™ generation fruit moth larvae increased to
61-74%, respectively, a positive response to increasing host numbers. Increased parasitism
of 2™ generation larvae led to low adult emergence.
Pea Aphid, Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae), an invasive
species believed to be of Palaearctic-Oriental origin, was first reported in the Ottawa
area about 1898 (Mackauer 1971). Mackauer and Bisdee (1965) reported on the status of
Aphidius smithi Sharma and Subba Rao (Hymenoptera: Aphidiidae) introduced to control
pea aphid. Their southern Ontario survey revealed Praon pequodorum Viereck and Aphidius
nigripes Ashmead (Hymenoptera: Aphidiidae) to be the principal parasitoids of A. pisum,
with Praon sp. and Aphelinus semiflavus Howard (Hymenoptera: Aphelinidae) of secondary
importance. Although not released in Ontario, A. smithii was found in areas adjacent to
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Lake Ontario and it was concluded that the populations present were the result of dispersal
from releases made in the USA (New Jersey, Delaware, Pennsylvania) in the late 1950s.
Pear Psylla, Cacopsylla pyricola (Férster) (Hemiptera: Psyllidae), a European
invader was first reported in Ontario in 1894 (McMullen 1971). Wilde (1965) studied the
biology of C. pyrico/a and noted that the nymphal parasitoid 7rechnites insidiosus (Crawford)
(Hymenoptera: Encyrtidae) was abundant before the widespread use of insecticides came
to dominate control strategies. However, the predators Chrysopa spp., Hippodamia sp.,
Cycloneda sp., Ceratomegilla sp., Anthocoris sp. and Orius sp. were abundant during the
study period. Anthocoris melanocerus Reuter (Hemiptera: Anthocoridae) from British
Columbia was released in southwestern Ontario (Wilde 1965). Philogene and Chang
(1979) reported new records of parasitic chalcidoids of pear psylla in Ontario. Trechnites
insidiosus, Pachyneuron sp. and Coccidencyrtus sp. were found for the first time parasitizing
C. pyricola.
Potato Stem Borer, Hydraecia micacea (Esper) (Lepidoptera: Noctuidae), an
invasive pest of European origin, established in southern Ontario in the 1960s, becoming a
pest in eastern Ontario in 1968 (Deedat et al. 1983). West et al. (1984) studied the parasitoids
of H. micacea, in southern Ontario and Europe. In Ontario, the tachinid Lydella radicis
(Townsend) (Diptera: Tachinidae) was reared from 61% of the host larvae collected. Other
parasitoids recovered were Diadegma sp., Campoletis sp., Glypta sp., and Pterocormis sp.
and Therion sp. (Hymenoptera: Ichneumonidae), although parasitism levels were low (0.5—
6%). European parasitoids imported into quarantine included Lydella stabulans (Meigen)
(Diptera: Tachinidae), Macrocentrus blandus Eady and Clark (Hymenoptera: Braconidae),
Exephanes occupator Gravenhorst (Hymenoptera: Ichneumonidae) and an unidentified
mermithid nematode. Comparison of the biologies of L. radicis and L. stabulens suggested
that these species may coexist in the field. Lydella stabulens has a lower developmental
threshold (6.7°C) and develops faster (159 Degree days) than L. radicis (13.5 °C and 113
Degree days), suggesting that the latter species would attack overwintering potato stem
borer larvae earlier in the season than the former species. Developmental studies of M.
blandus suggested that it may require an alternate host in order to produce a 2™ generation
in summer. Small numbers of L. stabulans and M. blandus were released near Guelph
(43.7167°N 80.4000°W).
Red Clover Casebearer, Coleophora deauratella Lienig and Zeller (Lepidoptera:
Coleophoridae), was discovered in Ontario in 1989 at New Liskard (Ellis and Bjornson
1996). This European native is a threat to red clover, Trifolium pratense L. (Fabaceae), seed
crops. Ellis and Bjornson (1996) studied the biology and biological control of red clover
casebearer. Based on a successful biological control program in New Zealand, individuals
of the European native Neochrysocharis formosus (Westwood) (Hymenoptera: Eulophidae)
were imported from the population established in New Zealand and released in Ontario.
However, no recoveries of N. formosa were made, although several other parasitoids,
including the native Bracon pygmaeus Provancher (Hymenoptera: Braconidae), were
reared. There are some taxonomic issues relating to whether the New Zealand specimens
released in Ontario were indeed N. formosa, known to be Holarctic, or a distinct more host-
specific population of N. formosa, or the related European N. trifolii Erdés (Hymenoptera:
Eulophidae).
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6. Inundative Biological Control of Arthropods with Pathogens
Entomopathogens, like other natural enemies, are important agents for reducing
populations of pest arthropods, particularly insects. Among these, Bacillus thuringiensis
Berliner (Bacilliaceae) is the most studied and this is evident in the JESO publications
summarized here. In addition, papers in JESO have evaluated several other entomopathogenic
organisms for their potential for inundative biological of pest insects (Table 3).
Cameron (1952) conducted a review of diseases of insects to 1951. Included
was information on the fungi, Beauwveria bassiana, Anisoplia austraca Herbst,
Metarhizium anisopliae (Metchnikoff) Sorokin (Clavicipitaeae), Aspergillus flavus Link
(Trichocomaceae) and /saria larinosa (Holmskiold) Fries (Moniliaceae), the bacteria
Enterobacter aerogenes Hormaeche and Edwards, Bacillus subtilis (Ehrenberg), B. proteus
(Bach), B. thuringiensis, and Paenibacillus popilliae Dutkey (Bacilliaceae), as well as
polyhedroviruses and granuloviruses. The main conclusion was that better understanding
of the biology and pathogenesis of the organisms should be a priority, before practical
application as biopesticides could be considered. Later, Cameron (1969) reviewed the
problems and prospects in the use of pathogens for insect control. He reported that B.
thuringiensis and B. papillae were the most practical and most developed pathogens at
the time. Other pathogens reported on in JESO include Nosema species (Microsporida),
viruses, and pathogenic nematodes.
Bacillus thuringiensis. Angus and Heimpel (1960) reviewed the potential
of bacteria for insect control. Among the several species mentioned, strains of Bacillus
thuringiensis Berliner (Bt) were considered to be promising and the authors concluded
that bacterial pathogens can be used to advantage in certain situations but they will never
entirely replace chemical insecticides.
Angus (1965) studied the post-larval mortality of Bt on forest tent caterpillar, the
Grey Midget, Nycteola cinereana Neumoegen and Dyar (Lepidoptera: Nolidae), and the
Mourning Cloak, Nymphalis antiopa (L.) (Lepidoptera: Nymphalidae). Results showed that
while most larvae were killed by Bt in the larval stage, some individuals of each species
pupated; however, these did not survive and they contained Br cells. Stewart et al. (1992)
studied the factors affecting the efficacy of Bt serovar. San Diego against larvae of the
Colorado potato beetle. They determined that young larvae are most susceptible and should
be targeted when using this agent. Morris (1980) isolated microbial pathogens from the
Maize Weevil, Sitophilus zeamais Motchulsky (Coleoptera: Curculionidae), including a
Bacillus sp. from adults and two Pseudomonas spp. from larvae and pupae.
Tripp (1972) reported on field trials of Bt applications to control Eastern Spruce
Budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae). Application
rates of 3.6 and 4.0 billion International Units (BIU) / US gal / acre effected mortalities of
96-99% on balsam fir and 80-86% on white spruce 33 days after spraying, although the
occurrence of frost shortly after spraying may have influenced mortality.
Cadogan et al. (1987) evaluated a formulation of Br on Jack Pine Budworm,
Choristoneura pinus pinus (Freeman) (Lepidoptera: Tortricidae). Futura®, a new Br
formulation effectively suppressed populations of C. pinus pinus and prevented serious
defoliation of host tress when applied at a rate of 30 BIU/ha. Cadogan (1993) showed that
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Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
C. pinus pinus larvae that survived Br applied at 30 BIU weighed significantly less than
controls and Bt applied at 20 BIU, suggesting that weights of surviving larvae could be used
as an additional criterion for assessing efficacy of Bz.
Nosema species — Wilson (1978) determined that incidence of the microsporidian
Nosema fumiferanae (Thompson) (Nosematidae) infections increased from 35.9-69.0%
over a five-year period of outbreak of its host, eastern spruce budworm. Wilson (1981)
looked at the effects of N. fumiferanae on rearing stock of this host. Synthetic diets allowed
the host to cope better with infection by N. fumiferana. Wilson (1985a) studied transmission
and effects of N. fumiferanae and Pleistophora schubergi ZwGlfer (Pleistophoridae) on
eastern spruce budworm. Males infected with either N. fumiferanae or P. schubergi did
not transmit spores to uninfected females through mating. However, P. schubergi infection
reduced pupal weight (about 30%) and adult longevity of females by 2.5 days. Wilson
(1985b) studied dose mortality response of P. schubergi on eastern spruce budworm and
found that a dose of 5 x 10° spores/larva caused >80% mortality of larvae and a dose of
5x10’ spores/larva caused 100% mortality. Higher doses resulted in decreased survival times
of infected larvae. Wilson (1987) found that Vairimorpha necatrix (Kramer) (Nosematidae)
caused high mortality: a dose of 5 x 10* spores/needle caused 100% mortality. Large doses
caused mortality by gut damage and bacterial septicemia, whereas low doses caused death
by microsporidiosis usually just before pupation.
Wilson and Burke (1979) documented the presence of three microsporidians,
Nosema cerasivoranus Thomson (Nosematidae), Pleistophora sp. (Pleistophoridae)
and Thelohania sp. (Thelohaniidae) from larvae of the Ugly Nest Caterpillar, Archips
cerasivoranus (Fitch) (Lepidoptera: Tortricidae). Levels of parasitism varied between years
and among species, Pleistophora sp. at 335.9% was the most prevalent, followed by N.
cerasivoranae at 0-28%, and Thelohania at 0-2.3%. Wilson (1980) examined the effects of
Nosema disstriae Thompson (Nosematidae) on the forest tent caterpillar, M. disstria, finding
that this microsporidian adversely affected pupal weights, adult fecundity and longevity.
Laing and Jaques (1985) studied Microsporidia associated with the European
Corn Borer, Ostrinia nubilalis (Hibner) (Lepidoptera: Pyralidae) over a 7-year period.
Applications of Nosema pyrausta (Paillot) (Nosematidae), V. necatrix, Bt and Autographa
californica nuclear polyhedrosis virus (ACNPV) (Baculoviridae) had little or no effect on
reducing crop damage compared to insecticides. However, Microsporidia infection levels
of field collected corn borer larvae (17-40%) and adults (10-24%) did not result in reduced
damage to the current crop but these authors concluded that infection levels may, over the
longer term, reduce viability of populations of the pest.
Viruses — Cunningham et al. (1987) found the nuclear polyhedrosis virus
Lecontvirus (Baculoviridae) to be highly effective against the Redheaded Pine Sawfly,
Neodiprion lecontei (Fitch) (Hymenoptera: Tenthredinidae). A dose of 5 x 10° polyhedral
inclusion bodies (PIB) per ha in spray volumes of 2.49.4 L/ha provided consistent control
when applied to 1‘-3" instar larvae. The virus can be cheaply produced (50 infected larvae
can produce enough concentrate for the 5 x 10° PIB/ha dose) at about $2.50/ha in 1985
dollars and applied using water alone. Evaluation of 100 trees, each with one redheaded pine
sawfly colony and scoring colonies as healthy, diseased or dead, allowed reliable monitoring
of epizootic progress. It was registered in Canada and was being used by Ontario.
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Jaques (1971) studied the potential for use of viruses to control cabbage insect
pests. Natural epizootics of the nuclear polyhedrosis viruses, Tricoplusia ni NPV (TnNPV)
and Pieris rapae GV (PrGV) (Baculoviridae) contributed substantially to control of T. ni
and P. rapae in the latter part of the season. Natural epizootics were the result of virus
accumulations in the soil, TnNPV residues being found in 60% and P. rapae GV in 19%
of samples taken. Application of the viruses to plants resulted in control as effective as or
better than that provided by chemical pesticides.
Bird et al. (1973) studied the possible use of a nuclear polyhedrosis virus (NPV) and
entomopoxvirus (EPV) (Poxyviridae) to control eastern spruce budworm. Both viruses were
isolated from eastern spruce budworm and the Two-year-cycle Budworm, Choristoneura
biennis Freeman (Lepidoptera: Tortricidae) from British Columbia. EPV was more effective
on white spruce than on balsam fir in early season applications, while late spray of NPV was
more effective. Virus carryover from 1971—1972 occurred.
Cunningham et al. (1996a) evaluated Disparvirus, nuclear polyhedrosis virus, and
Bt serovar. kurstaki (Btk) applied as aerial sprays on mortality of gypsy moth. Average
egg mass reductions from Disparvirus were 76% and 80% at a rates of 5.0 and 2.5 L/ha,
respectively, and 96% for Btk at 50 billion International Units (BIU) in 4.0 L/ha. Cunningham
et al. (1996b) reported on impact of Disparvirus and Btk one year after application. Gypsy
moth, larvae were 20.4% positive for NPV in plots treated with Disparvirus at 5.0 L/ha,
14.6% positive for NPV in plots treated at 2.5 L/ha, and 8.0% positive for NPV in plots
treated with Brk, and 9.2% positive for NPV in control plots. Negligible foliage damage was
reported and fall egg mass numbers were low indicating that in the treated area, the gypsy
moth population had collapsed, suggesting that NPV was a contributing factor.
Nematodes — Welch (1962) reviewed the status of nematodes as agents for insect
control. In nature, nematodes are generally not significant mortality factors, although
under some conditions they may be significant regulatory factors. Mermithidae have the
greatest potential as biological control agents because of their size and similarity to insect
parasitoids. Neoaplectanidae also show potential because of their high rate of reproduction.
Allantonematidae and Aphelenchoidea are best suited to environmental manipulation.
Moisture, moderate temperatures and high host density are important factors for successful
control.
Welch and Briand (1962) evaluated a neoplectanid nematode for control of Colorado
potato beetle, cabbage root maggot, European corn borer and the imported cabbage worm.
Use of the nematode was most promising for control of cabbage root maggot and European
corn borer where the soil environment provides conditions suitable for nematode survival.
Briand (1960) reported the occurrence of the nematode Howardula beninga
Cobb (Tylenchida: Allantonematidae) in the Striped Cucumber Beetle, Diabrotica vittata
(Fabricius) (Coleoptera: Chrysomelidae). Parasitism was 7.6% and 2.5% in surveys
conducted in 1958 and 1959, respectively. Parasitism was nil in the secondary host D.
unidecimpunctata howardi (Barber) even though this species was common in the southern
Ontario study area.
Wright (1972) reported a new Canadian record for the adventive nematode
Heterotylenchus autumnalis Nickle (Nematoda: Sphaerulariidae) as a parasite of the Face
Fly, Musca autumnalis DeGeer (Diptera: Muscidae). Heterotvlenchus autumnalis is widely
distributed in Ontario but incidence was <2% and unlikely to contribute significantly to
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Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
natural control. Gregory and Wright (1973) released irradiated female face flies parasitized
with H. autumnalis and found that doses of 1.0 and 2.5 krad did not sterilize the nematodes
and parasitized face fly females produced progeny with high levels of parasitism. Release
of sterile flies that were parasitized was considered better than the release of sterile flies
alone.
Welch (1958) evaluated the nematode Neaplectana chresima Steiner (Rhabdidita:
Steinernematidae) for biological control of Colorado potato beetle. Application of ~20,000
cultured nematodes resulted in an approximate 14% reduction in beetle numbers although
abiotic factors, 1.e., significant rainfall, had an impact on the nematodes.
7. Natural Enemy Taxonomy
Taxonomy is essential to biological control and a few studies on groups relevant to
biological control have been published in JESO. These studies, while clarifying taxonomic
status, unfortunately also demonstrate just how poorly the biology of parasitoids is
understood. Six taxonomic studies published in JESO that are relevant to biological control
treat taxa within the Hymenoptera familes Braconidae (2), Eucharitidae (1) and Mymaridae
(3).
Braconidae. Loan (1970) described the new species, Leiophron pseudopallipes
Loan and Leiophron lygivora (Loan) (Hymenoptera: Braconidae) reared from tarnished
plant bug, Lygus lineolaris, in Ontario. Leiophron pseudopallipes is ecologically distinct,
attacking 2™ generation L. /ineolaris, from the related L. mellipes (Cresson) which attacks
the 1* generation. Leiophron lygivora also attacks 2"! generation L. lineolaris. Loan
and New (1972) reviewed the taxonomy of the Euphorine (Hymenoptera: Braconidae)
genus Leiophron, subgenus Euphoriella Ashmead and redescribed L. (E.) sommermanae
(Muesebeck), L. (E.) incerta (Ashmead), and L. (E.) pacifica (Muesebeck). Leiophron (E.)
nixoni (Loan and New), L. (E.) kaladarensis (Loan and New), L. (E.) solidaginis (Loan
and New), L. (E£.) foutsi (Loan and New), L. (E.) pallidifacia (Loan and New), L. (E.)
hyalopsocidis (Loan and New) and L. (E.) criddlei (Loan and New) were newly described.
Leiophron (E.) hyalopsocidis was the only species associated with a host and it was reared
from the psocid Hyalopsocus striatus (Walker) (Psocoptera: Psocidae).
Sharkey (2007) revised the Neotropical Braconidae (Hymenoptera) genus
Trachagathis Viereck. Among the 3 species treated, Trachagathis rubricincta (Ashmead) is
associated with the lesser cornstalk borer, Elasmopalpus lingosellus (Zeller) (Lepidoptera:
Pyralidae), from sugarcane and the biologies of the other two species are unknown.
Eucharitidae. Heraty (1985) revised the Nearctic Eucharitinae (Hymenoptera:
Eucharitidae), providing keys to the 5 genera and 16 species. Species of Eucharitidae are
specialized ant parasitoids. Among the species treated, only the host of Pseudometagea
schwarzii (Ashmead), the ant Lasius neoniger Emery (Hymenoptera: Formicidae), is
known.
Mymaridae. Huber (1992) studied the subgenera and species groups of Anaphes
(Hymenoptera: Mymaridae), and reviewed the described Nearctic species of the fuscipennis
group of Anaphes s.s. and the described species of Anaphes (Yungaburra). Anaphes spp.
are mostly parasitoids of Curculionidae and Chrysomelidae. Among the 9 species of the
Anaphes fuscipennis group treated, hosts have been associated with Anaphes fuscipennis
Haliday [Sitona humeralis Stephens, Hypera postica (Gyllenhal) and H. punctata
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Mason JESO Volume 144, 2013
(Fabricius) (Coleoptera: Curculionidae)], 4. io/e Girault [Lygus spp. and Pseudatomoscelis
sp. (Hemiptera: Miridae)], 4. byrrhidiphagus Huber [Lioon simplicipes (Mannerheim) and
Lioligus nitidus (Motschulsky) (Coleoptera: Byrridae)], and Anaphes flavipes (Forster)
[Oulema melanopus (L.), O. gallaeciana (Heydon), O. collaris (Say), Lema trilineata
Oliver, L. trilineata trivittata (Say), L. lichenis Voet. and L. cyanella (L.) (Coleoptera:
Chrysomelidae)]. Anaphes flavipes was imported for biological control of O. melanopus.
Hosts are unknown for the six species of the Anaphes (Yungabura) group.
Huber (2006) reviewed the described species of the Anaphes crassicornis group,
important in biological control with the aim to improve identification of the species. Among
the 13 species treated hosts are known for Anaphes calendrae (Gahan) [Sphenophorus spp.
(Coleoptera: Curculionidae)], A. conotracheli Girault | Conotrachelus geminatus (LeConte),
Hypera nigrirostris (Fabricius) (Coleoptera: Curculionidae)], 4. cotei Huber [Listronotus
oregonensis (LeConte) (Coleoptera: Curculionidae)], A. diana (Girault) [Sitona hispidulus
(Fabricius), S. humeralis Stephens, S. lineatus (L.) (Coleoptera: Curculionidae)], A.
gerrisophagus (Doutt) [Gerris sp. (Hemiptera: Gerridae) and Lestes sp. (Odonata: Lestidae)],
A. listronoti Huber [L. oregonensis], A. luna (Girault) [Hypera spp., and in North America,
H. postica (Gyllenhal) and H. eximia (LeConte) (Coleoptera: Curculionidae)], A. pallipes
(Ashmead) [Cylindrocopturus adspersus (LeConte) (Coleoptera: Curculionidae) and
Rhagoletis pomonella Walsh (Diptera: Tephritidae)|, A. pullicrurus (Girault) [Chaetoctema
denticulata (Illiger) (Coleoptera: Chrysomelidae)], A. sordidatus |Tyloderma foveolatum
(Say) (Coleoptera: Curculionidae)]|, and A. victus Huber [L. oregonensis]. Anaphes luna and
A. diana were imported and released in the USA as biological control agents.
Huber (2012) revised the Ooctonus spp. (Hymenoptera: Mymaridae) in the Nearctic
region. Among the 15 species described, hosts are known for O. aphrophorae Milliron
[on Aphrophora saratogensis (Fitch) (Hemiptera: Cercopidae)|], and O. vulgatus Haliday
[on Philaenus spumarius (L.) (Hemiptera: Cercopidae)]. Although white pine weevil was
recorded as a potential host for O. guadricarinatus Girault the record is incorrect (J. Huber,
personal communication).
Conclusions
Over the years, the Journal of the Entomological Society of Ontario has been an
important venue for dissemination of scientific results on biological control of pest arthropods
and weeds in Ontario. Included are studies on natural enemy assemblages, biology of natural
enemies, releases of exotic species as agents for biological control, entomopathogens for
use in reduced risk management strategies, and taxonomy of groups important to biological
control. In recent years, competition with an ever increasing number of specialized journals
with high impact factors, many of which have no page charges, has led to a decline in
submissions to JESO. However, there are unfilled niches for which JESO can provide a
good opportunity to publish: documenting the status and distribution of natural enemies
intentionally released as biological control agents, documenting associations among natural
enemies and hosts, and assessing changes in natural enemy assemblages over time.
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Acknowledgements
Andrea Brauner assisted in the task of verifying taxonomic names and JESO editor, John
Huber invited this review. Two reviewers provided constructive recommendations to
improve the manuscript.
References
Abu, J. F. and Ellis, C. R. 1976. Biology of Bathyplectes curculionis (Thomson) (Hymenoptera:
Ichneumonidae) a parasitoid of the alfalfa weevil in Ontario. Proceedings of the
Entomological Society of Ontario 106: 8-12.
Alex, J. F. 1998. Ontario Weeds. Ontario weeds: descriptions, illustrations and keys to their
identification. Ontario Ministry of Agriculture, Food and Rural Affairs Publication
505, Guelph, Ontario, Canada. Available from http://www.omafra.gov.on.ca/english/
crops/pub505/p505order.htm [Accessed 1 November 2013].
Angus, T. A. and Heimpel, A. M. 1960. The bacteriological control of insects. Proceedings of the
Entomological Society of Ontario 90: 13-21.
Angus, T. A. 1965. Mortality due to Bacillus thuringiensis in post-larval stages of some
Lepidoptera. Proceedings of the Entomological Society of Ontario 95: 133-134.
Anonymous. 2012. Noxious weeds in Ontario. Available at: http://www.omafra.gov.on.ca/
english/crops/facts/noxious_weeds.htm [Accessed 1 November 2013].
Appleton, E. S. B., Gillard, C. and Schaafsma, A. W. 2004. Biology and management of the
potato leafhopper, Empoasca fabae Harris (Homoptera: Cicadellidae) on field crops in
Ontario. Journal of the Entomological Society of Ontario 134: 3-17.
Bahlai, C. A. and Sears, M. K. 2009 Population dynamics of Harmonia axyridis and Aphis
glycines in Niagara peninsula soybean fields and vineyards. Journal of the Entomological
Society of Ontario 140: 27-39.
Beirne, B. P. 1971. Pest insects of annual crop plants in Canada: Part I, Lepidoptera; II, Diptera;
III, Coleoptera. Memoirs of the Entomological Society of Canada 103, Supplement
S78: 1-124.
Bennett, A. M. R. 2004. Host location behaviour in Pelecinus polyturator (Hymenoptera:
Pelecinidae). Journal of the Entomological Society of Ontario 135: 131—134.
Ben-Ze’ev, L. S. and Jaques, R. P. 1990. Entomopathogenic fungi in insects in alfalfa fields
in southwestern Ontario. Proceedings of the Entomological Society of Ontario 121:
71-78.
Best, K. F., Bowes, G. G., Thomas, A. G. and Maw, M. G. 1980. The biology of Canadian weeds.
39. Euphorbia esula L. Canadian Journal of Plant Science 60: 651-663.
Bird, F. T., Cunningham, J. S. and Howse, G. M. 1973. The possible use of viruses in the control
of spruce budworm. Proceedings of the Entomological Society of Ontario 102: 69—
a:
Bolter, C. J. and Laing J. E. 1984. Competition between Diadegma insulare (Hymenoptera:
Ichenumonidae) and Microplitis plutellae (Hymenoptera: Braconidae) for larvae of the
diamonback moth, Plutella xylostella (Lepidoptera: Plutellidae). Proceedings of the
Entomological Society of Ontario 114: 1-10.
Briand, L. J. 1960. The nematode Howardula benigna Cobb, 1921, parasite of the cucumber
beetles in Ontario. Proceedings of the Entomological Society of Ontario 90: 53.
53
Mason JESO Volume 144, 2013
Bright, D. E. 1996. Notes on native parasitoids and predators of the larger pine shoot beetle,
Tomicus piniperda (Linnaeus) in the Niagara region of Canada (Coleoptera: Scolytidae).
Proceedings of the Entomological Society of Ontario 127: 57-62.
Broadbent, A. B., Goulet, H., Whistlecraft, J. W., Lachance, S. and Mason, P. G. 1999. First
Canadian record of three parasitoid species (Hymenoptera: Braconidae: Euphorinae)
of the tarnished plant bug, Lygus lineolaris (Hemiptera: Miridae). Proceedings of the
Entomological Society of Ontario 130: 109-111.
Broadbent, A. B., Haye, T., Gariepy, T. D., Olfert, O. O. and Kuhlmann, U. 2013. Lygus lineolaris
(Palisot), tarnished plant bug (Hemiptera: Miridae). Pp. 221-227 in Mason P. G. and
Gillespie, D. R. (eds), Biological Control Programmes in Canada 2001—2012. CABI
Publishing, Wallingford, Oxfordshire United Kingdom, 518 pp.
Boyce, H. R. and Dustan, G. G. 1954. Parasitism of twig-infesting larvae of the oriental fruit
moth, Grapholita molesta (Busck) (Lepidoptera: Olethreutidae), in Ontario, 1939—
1953. Annual Report of the Entomological Society of Ontario 84: 48-55.
Cadogan, B. L. 1993. Field weights of jack pine budworm larvae (Lepidoptera: Tortricidae)
surviving aerial applications of Bacillus thuringiensis and two chemical insecticides.
Proceedings of the Entomological Society of Ontario 124:189-196.
Cadogan, B. L. and Laing, J. E. 1982. A study of Balaustium putnami (Acarina: Erythraeidae)
in apple orchards in southern Ontario. Proceedings of the Entomological Society of
Ontario 112: 13-21.
Cadogan, B. L., Zylstra, B. F., Nystrom, C., Ebling, P. M. and Pollock, L. B. 1987. Evaluation
of a new Futura” formulation of Bacillus thuringiensis on populations of jack pine
budworm, Choristoneura pinus pinus (Lepidoptera: Tortricidae). Proceedings of the
Entomological Society of Ontario 117: 59-64.
Cameron, J. W. M. 1952. A survey of investigations on the diseases of insects. Annual Report of
the Entomological Society of Ontario 83: 7-14.
Cameron, J. W. M. 1969. Problems and prospects in the use of pathogens in insect control.
Proceedings of the Entomological Society of Ontario 99: 73-81.
Carter, N. J. and Laing, J. E. 1997. Cotesia rubecula (Marshall) (Hymenoptera: Braconidae)
recovered three years after release in southern Ontario. Proceedings of the Entomological
Society of Ontario 128: 113-115.
Chant, D. A.1957. Biological control. Annual Report of the Entomological Society of Ontario
87: 54-58.
Clements, D. R. 1989. The role of stigmaeids (Acari) in the orchard acarine system. Proceedings
of the Entomological Society of Ontario 120: 74-75.
Coote, L. D. and Ellis C. R. 1987. Parasites of the alfalfa blotch leafminer, Agromyza frontella
(Diptera: Agromyzidae), near Guelph, Ontario. Proceedings of the Entomological
Society of Ontario 117: 21—27.
Coote, L. D. and Ellis, C. R. 1987. Diglyphus intermedius (Hymenoptera: Eulophidae), in
indigenous parasite of the alfalfa blotch leafminer, Agromyza frontella (Diptera:
Agromyzidae). Proceedings of the Entomological Society of Ontario 117: 83-85.
Coppel, H. C. and Arthur, A. P. 1954. Notes on introduced parasites of the European pine shoot
moth, Rhyacionia buliana (Schiff.) (Lepidoptera: Tortricidae), in Ontario. Annual
Report of the Entomological Society of Ontario 84: 55—S8.
Corrigan, J. E. 1983. Cotesia (Apanteles) rubecula (Hymenoptera: Braconidae) recovered in
Ottawa, Ontario ten years after its release. Proceedings of the Entomological Society
of Ontario 113: 71.
54
Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
Corrigan, J. E. and Laing, J. E. 1992. An improved method for producing small, consistent
samples of hosts for presentation to the egg parasitoid, Trichogramma minutum.
Proceedings of the Entomological Society of Ontario 122: 103-104.
Corrigan, J. E., Lashomb, J. H. and Kingsley, P. C. 1990. Pupal orientation and emergence success
of the egg parasitoids Edovum puttleri (Hymenoptera: Eulophidae). Proceedings of the
Entomological Society of Ontario 121: 95—100.
Corrigan, J. E., Laing, J. E. and Caddick, G. 1994. Feasibility of delaying emergence of
Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae) and subsequent
effects on adult longevity and fecundity. Proceedings of the Entomological Society of
Ontario 125: 81—86.
Corrigan, J. E., Mackenzie, D. L. and Simser, L. 1998. Field observations of non-target feeding
by Galerucella calmariensis {Coleoptera: Chrysomelidae], an introduced biological
control agent of purple loosestrife, Lythrum salicaria |Lythraceae]. Proceedings of the
Entomological Society of Ontario 129: 99-106.
Crompton, C. W., Hall, I. V., Jensen, K. I. N. and Hildebrand, P. D. 1988. The biology of
Canadian weeds. 83. Hypericum perforatum L. Canadian Journal of Plant Science
68: 149-162.
Cunningham, J. C., deGroot, P. and Kaup, W. J. 1987. A review of aerial spray trials with
Lecontvirus for control of redhead pine sawfly, Neodiprion lecontei (Hymenoptera:
Diprionidae), in Ontario. Proceedings of the Entomological Society of Ontario 117:
65-72.
Cunningham, J. C., Payne, N. J., Brown, K. W., Fleming, R. A., Burns, T., Mickle, R. E. and
Scarr, T. 1996a. Aerial spray trials with nuclear polyhedrosis virus and Bacillus
thuringiensis on gypsy moth (Lepidoptera: Lymantriidae) in 1994. I. Impact in the year
of application. Proceedings of the Entomological Society of Ontario 127: 21-35.
Cunningham, J. C., Brown, K. W., Scarr, T., Fleming, R. A. and Burns, T. 1996b. Aerial spray
trials with nuclear polyhedrosis virus and Bacillus thuringiensis on gypsy moth
(Lepidoptera: Lymantriidae) in 1994. II. Impact one year after application. Proceedings
of the Entomological Society of Ontario 127: 37-43.
Deedat, Y. D., Ellis, C. R. and West, R. J. 1983. Life History of the Potato Stem Borer (Lepidoptera:
Noctuidae) in Ontario. Journal of Economic Entomology 76: 1033-1037.
Desrochers, A. M., Bain, J. F. and Warwick, S. I. 1988. The Biology of Canadian weeds. 89.
Carduus nutans L. and Carduus acanthoides L. Canadian Journal of Plant Science
68: 1053-1068.
Dustan, G. G. and Boyce, H. R. 1966. Parasitism of the oriental fruit moth, Grapholitha
molesta (Busck) (Lepidoptera: Tortricidae) in Ontario, 1956-1965. Proceedings of the
Entomological Society of Ontario 96: 100-102.
Edwards, C. J. and Pengelly, D. H. 1966. Melittobia chalybii Ashmead (Hymenoptera: Eulophidae)
parasitizing Bombus fervidus Fabricius (Hymenoptera Apidae). Proceedings of the
Entomological Society of Ontario 96: 98-99.
Ellis, C. R., Harcourt, D. G. and Dubois-Martin, D. 1979. The current status in Ontario of
Tetrastichus julis (Hymenoptera: Eulophidae), a parasitoid of the cereal leaf beetle.
Proceedings of the Entomological Society of Ontario 109: 23-26.
Ellis, C. R., Kormos, B. and Guppy, J. C. 1989. Absence of parasitism in an outbreak of the
cereal leaf beetle, Oulema melanopus (Coleoptera: Chrysomelidae), in the central
tobacco growing area of Ontario. Proceedings of the Entomological Society of Ontario
119: 43-46.
55
Mason JESO Volume 144, 2013
Ellis, C. R. and Nang’ayo, F. L. O. 1992. The biology and control of the trefoil seed chalcid,
Bruchopagus platypterus (Walker) (Hymenoptera: Eurytomidae), in Ontario.
Proceedings of the Entomological Society of Ontario 123: 111-122.
Ellis, C. R. and Bjornson, S. 1996. The biology, importance, and biological control of Coleophora
deauratella (Lepidoptera: Coleophoridae), a new pest of red clover in North America.
Proceedings of the Entomological Society of Ontario 127: 115-124.
Fisher, P. 1988. The effect of pesticides on two species of parasitoids, Pholetesor ornigis and
Pholetesor pedias (Hymenoptera: Braconidae). Proceedings of the Entomological
Society of Ontario 118: 171.
Foott, W. H. 1974. Observations on Coccinellidae in corn fields in Essex County, Ontario.
Proceedings of the Entomological Society of Ontario 104: 16-21.
George, J. A. 1979. The potential of a local planarian, Dugesia tigrina (Tricladida, Turbellaria),
for the control of mosquitoes in Ontario. Proceedings of the Entomological Society of
Ontario 109: 23.
George, J. A. 1984. Survival of the mosquito predator Dugesis tigrina (Tricladida: Turbellaria),
over three years in catch basins. Proceedings of the Entomological Society of Ontario
114: 83-85.
Glen, R. 1962. Preface. Pp. 1x—x in McLeod, J. S., McGugan, B. M. and Coppel, H. C. (eds),
A review of the biological control attempts against insects and weeds in Canada.
Technical Communication Commonwealth Institute of Biological Control 2, Franham
Royal, Bucks, England, 216 pp.
Glen, R. 1965. Entomology in Canada up to 1956. Areview of developments and accomplishments.
The Canadian Entomologist 58: 290-371.
Goble, H. W. 1965. The armyworm Pseudaletia unipuncta (Hawthorn) in Ontario in 1964.
Proceedings of the Entomological Society of Ontario 95: 11-14.
Graham, A. R. 1958. Effectiveness of two introduced parasites of the larch casebearer,
Coleophora laricella (Hbn.) (Lepidoptera: Coleophoridae), in Ontario. Annual Report
of the Entomological Society of Ontario 88: 37-41.
Gregory, D. and Wright, R. E. 1973. Irradiation of the female face fly, Musca autumnalis DeGeer,
parasitized by the nematode, Heterotylenchus autumnalis Nickle. Proceedings of the
Entomological Society of Ontario 102: 104-107.
Griffiths, K. J. 1972. Discrimination between parasitized and unparasitized hosts by Pleolophus
basizonus (Hymenoptera: Ichneumonidae). Proceedings of the Entomological Society
of Ontario 102: 83-91.
Griffiths, K. J. 1977. A preliminary report on the gypsy moth and its parasites in southeastern
Ontario. Proceedings of the Entomological Society of Ontario 107: 79-84.
Hagley, E. A. C. 1970. The distribution and survival of overwintering codling moth larvae in
southern Ontario. Proceedings of the Entomological Society of Ontario 100: 40-47.
Hagley, E. A. C. 1979. Integrated pest management — insecticides and natural predator
populations on apple. Proceedings of the Entomological Society of Ontario 109:
9-21.
Hagley, E. A. C. 1987. Occurrence of Trichogramma spp. (Hymenoptera: Trichogrammatidae)
in apple orchards in southern Ontario. Proceedings of the Entomological Society of
Ontario 117: 79-82.
Hagley, E. A. C. and Barber, D. R. 1992. Foliage-feeding Lepidoptera and their parasites
recovered from unmanaged apple orchards in southern Ontario. Proceedings of the
Entomological Society of Ontario 122: |—7.
56
Biological control in Ontario, 1952—2012 JESO Volume 144, 2013
Hagley, E. A. C. and Laing, J. E. 1989. Effect of pesticides on parasitism of artificially distributed
eggs of the codling moth, Cydia pomonella (Lepidoptera: Tortricidae). Proceedings of
the Entomological Society of Ontario 120: 25-33.
Harcourt, D. G. 1963. Biology of cabbage caterpillars in eastern Ontario. Proceedings of the
Entomological Society of Ontario 93: 61—75.
Harcourt, D. G. and Ellis, C. R. 1992. Distribution in southern Ontario of Bathyplectes anurus
(Hymenoptera: Ichneumonidae), a larval parasitoid of the alfalfa weevil. Proceedings
of the Entomological Society of Ontario 122: 41-45.
Harcourt, D. G., Ellis, C. R. and Guppy, J. C. 1980. Distribution of Microctonus aethiopoides, a
parasitoid of the alfalfa weevil (Coleoptera: Curculionidae) in Ontario. Proceedings of
the Entomological Society of Ontario 110: 35-39.
Harcourt, D. G., Guppy, J. C. and Ellis, C. R. 1982. Distribution of Microctonus colesi
(Hymenoptera: Braconidae), a new parasite of the alfalfa weevil in Ontario. Proceedings
of the Entomological Society of Ontario 112: 33-37.
Harcourt, D. G., Guppy, J. C. and Ellis C. R. 1987. Establishment and spread of Dacnusa dryas
(Hymenoptera: Braconidae), an exotic parasite of the alfalfa blotch leafminer in
Ontario. Proceedings of the Entomological Society of Ontario 117: 29-33.
Harmsen, R. and Rose, M. R. 1984. Habitat effects on larval mortality in the forest tent caterpillar,
Malacosoma disstria (Lepidoptera: Lasiocampidae). Proceedings of the Entomological
Society of Ontario 114: 87-89.
Heraty, J. M. 1985. A revision of the Nearctic Eucharitinae (Hymenoptera: Chalcidoidea:
Eucharitidae). Proceedings of the Entomological Society of Ontario 116: 61—103.
Herbert, H. J. 1953. Progress report on predacious mite investigations in Nova Scotia (Acarina:
Phytoseiidae). Annual Report of the Entomological Society of Ontario 84: 27-29.
Hikichi, A. 1962. Notes on mortality factors affecting the red-banded leaf roller, Argyrotaenia
velutinana (Wlkr.), (Lepidoptera: Tortricidae) in an unsprayed apple orchard in Ontario.
Proceedings of the Entomological Society of Ontario 92: 180-182.
Hikichi, A. 1971. 3. Argyrotaenia velutinana (Walker), red-banded leafroller (Lepidoptera:
Tortricidae). Pp. 10—12 in Kelleher et al. (eds), Biological control programmes against
insects and weeds in Canada 1959-1968. Technical Communication Commonwealth
Institute of Biological Control 4, 266 pp.
Hoffmeister T. S. 2002. Rhagoletis pomonella (Walsh), apple maggot (Diptera: Tephritidae). Pp.
238-241 in Mason, P. G. and Huber, J. T. (eds), Biological Control Programmes in
Canada 1981—2000. CABI Publishing, Wallingford, Oxon, United Kingdom, 583 pp.
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.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. 2012. Revision of the genus Ooctonus (Hymenoptera: Mymaridae) in the Nearctic
region. Journal of the Entomological Society of Ontario 143: 15-105.
Hulme, M. A. and Kenis, M. 2002. Pissodes strobe (Peck), white pine weevil (Coleoptera:
Curculionidae). Pp. 221—228 in Mason, P. G. and Huber, J. T. (eds), Biological Control
Programmes in Canada 1981-2002. CABI Publishing, Wallingford, Oxon, United
Kingdom, 583 pp.
Sf
Mason JESO Volume 144, 2013
Jaques, R. P. 1971. Control of cabbage insects by viruses. Proceedings of the Entomological
Society of Ontario 101: 28-34.
James, H. G. 1952. Natural control of Tabanidae (Diptera) in the region of Churchill, Manitoba.
Annual Report of the Entomological Society of Ontario 83: 70-74.
James, H. G. 1959. Egg development, hatching, and prey taken by the European mantis, Mantis
religiosa L., in several habitats. Annual Report of the Entomological Society of Ontario
89: 50-SS.
Jones, T., Shipp, J. L., Scott-Dupree, C. D. and Harris, C. R. 2006. Influence of greenhouse
microclimate on Neoseiulus (Amblyseius) cucumeris (Acari: Phytosetidae) predation
on Frankliniella occidentalis (Thysanoptera: Thripidae) and oviposition on greenhouse
cucumber. Journal of the Entomological Society of Ontario 136: 71-83.
Johnson, E. F., Laing, J. E. and Trottier, R. 1977. The seasonal occurrence of Lithocolletis
blancardella (Gracillariidae), and its major natural enemies in Ontario apple orchards.
Proceedings of the Entomological Society of Ontario 107: 31-45.
Judd, W. W. 1953. Diptera and Hymenoptera reared from pine cone willow galls caused by
Rhabdophaga strobiloides (Diptera: Itonididae). Annual Report of the Entomological
Society of Ontario 84: 34-42.
Kelleher, J. S. et al. 1971. Biological control programmes against insects and weeds in Canada
1959-1968. Technical Communication Commonwealth Institute of Biological Control
4, Franham Royal, Slough, England, 266 pp.
Kelleher, J. S. and Hulme, M. A. (eds) 1984. Biological Control Programmes agains insects and
weeds in Canada 1969-1980. Commonwealth Agricultural Bureaux, Franham Royal,
Slough, England, 410 pp.
Laing, J. E. 1978. Establishment of Urophora cardui L. (Diptera: Tephritidae) on Canada thistles
in southern Ontario. Proceedings of the Entomological Society of Ontario 108: 2.
Laing, D. R. and Jaques, R. P. 1985. Microsporidia of the European corn borer (Lepidoptera:
Pyralidae) in southwestern Ontario: natural occurrence and effectiveness as microbial
insecticides. Proceedings of the Entomological Society of Ontario 115: 13-17.
Laing, J. E. and Welch, H. E. 1963. A dolichopodid predacious on larvae of Culex restuans
Theob. Proceedings of the Entomological Society of Ontario 93: 89-90.
Laing, J. E. and Heraty, J. M. 1982. The parasite complex of the overwintering population of
Epiblema scudderiana (Lepidoptera: Olethreutidae) in southern Ontario. Proceedings
of the Entomological Society of Ontario 112: 59-66.
Laing, J. E. and Heels, P. R. 1979. Establishment of an introduced weevil, Rhinocyllus conicus
(Coleoptera: Curculionidae) for the biological control of nodding thistle, Carduus
nutans (Compositae) in southern Ontario. Proceedings of the Entomological Society
of Ontario 109: 3-8.
LeSage L. 1989. Survival of Aphthona species in Ontario. Biocontrol News 2: 1.
LeSage, L. 1996a. Suivi, pour les années 1994 et 1995, des altises de l’euphorbe (Coleoptera:
Chrysomelidae: Aphthona spp.) relachées a Marshall Bay (Ontario) contre l’euphorbe
(Euphorbia esula L.). Proceedings of the Entomological Society of Ontario 127: 125—
126.
LeSage, L. 1996b. Expansion de l’aire de répatition de Chrysolina hyperici (Forster) depuis son
introduction en Ontario (Coleoptera: Chrysomelidae). Proceedings of the Entomological
Society of Ontario 127: 127-130.
Li, S. Y. 2000. Larval susceptibility of balsam fir sawfly (Hymenoptera: Diprionidae) to neem.
Proceedings of the Entomological Society of Ontario 110: 139-143.
58
Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
Loan, C. C. 1964. Observations on the biology of Centistes excrucians Haliday (Hymenoptera:
Braconidae). Proceedings of the Entomological Society of Ontario 94: 56-61.
Loan, C. C. 1965. Life cycle and development of Leiophron pallipes Curtis (Hymenoptera:
Braconidae: Euphorinae) in five mired hosts in the Belleville district. Proceedings of
the Entomological Society of Ontario 95: 115-121.
Loan, C. C. 1970. Two new parasites of the tarnished plant bug in Ontario: Leiophron
pseudopallipes and Euphoriana lygivora (Hymenoptera: Braconidae, Euphorinae).
Proceedings of the Entomological Society of Ontario 100: 188-195.
Loan, C. C. 1973. Parasitism of adult Notoxus anchora Henz. (Coleoptera: Anthicidae)
by Syrrhizus agilis (Cresson) (Hymenoptera: Braconidae). Proceedings of the
Entomological Society of Ontario 102: 76.
Loan, C. 1982. Suppression of the fungi Zoohthora spp. by capitol: a technique to study interaction
between disease and parasitism of the alfalfa weevil Hyperica postica (Coleoptera:
Curculionidae). Proceedings of the Entomological Society of Ontario 112: 81-82.
Loan, C. C. and New T. R. 1972. An account of the North American species of Eurphoriella
Ashmead with descriptions of seven new species (Hymenoptera: Braconidae,
Euphorinae). Proceedings of the Entomological Society of Ontario 102: 92-108.
Lyons, D. B., Helson, B. V., Jones, G. C., McFarlane, J. W. and Scarr, T. 1996. Systemic activity
of neem seed extracts containing azadirachtin in pine foliage for control of the pine false
webworm, Acantholyda erythrocephala (Hymenoptera: Pamphiliidae). Proceedings of
the Entomological Society of Ontario 127: 45-55.
MacFarlane, R. P. and Pengelly, D. H. 1978. Brachicoma spp. (Sarcophagidae) and Melittobia
chalybii (Eulophidae) as parasites of the brood of Bombus spp. (Apidae) in southern
Ontario. Proceedings of the Entomological Society of Ontario 108: 31-35.
Mackauer, M. 1971. 2. Acyrthosiphon pisum (Harris), pea aphid (Homoptera: Aphididae).
Pp. 3-10 in McLeod, J. S., McGugan, B. M. and Coppel, H. C. (eds), Biological
control programmes against insects and weeds in Canada 1959-1968. Technical
Communication Commonwealth Institute of Biological Control 4, 266 pp.
Mackauer, M. J. P. and Bisdee, H. E. 1965. Aphidius smithi Sharma and Subba Rao (Hymenoptera:
Aphidiidae) a parasitoid of the pea aphid new in southern Ontario. Proceedings of the
Entomological Society of Ontario 95: 121-124.
Mal, T. K., Loverr-Doust, J., Lovett-Doust. L. and Mulligan, G. A. 1992. The biology of Canadian
weeds. 100. Lythrum salicaria. Canadian Journal of Plant Science 72: 1305-1330.
Mason, P. G. and Huber, J. T. (eds) 2002. Biological Control Programmes in Canada 198 1—2000.
CABI Publishing, Wallingford, Oxon, United Kingdom, 583 pp.
Mason P. G. and Gillespie, D. R. (eds) 2013. Biological Control Programmes in Canada 2001-
2012. CABI Publishing, Wallingford, Oxfordshire, United Kingdom, 518 pp.
Mason, P. G., Goulet, H. and Bostanian, N. 2011. Effect of harvest on Euphorine (Hymenoptera:
Braconidae) parasitism of Lygus lineolaris and Adelphocoris lineolatus (Hemiptera:
Miridae) in alfalfa. Journal of the Entomological Society of Ontario 142: 3-10.
Maybee, G. E. 1954. Introduction into Canada of parasites of the carrot rust fly, Psila rosae
(F.) (Diptera: Psilidae). Annual Report of the Entomological Society of Ontario 84:
58-62.
Maybee, G. E. 1956. Observations, life-history, habits, immature stages, and rearing of Loxotropa
tritoma (Thoms.) (Hymenoptera: Proctotrupoidea) a parasite of the carrot rust fly, Psila
rosae (F.) (Diptera: Psilidae). Annual Report of the Entomological Society of Ontario
86: 53-58.
59
Mason JESO Volume 144, 2013
Maxwell, C. W. and Morgan, G. T. 1952. Life-history studies of the cranberry fruitworm,
Mineola vaccinii (Riley), in New Brunswick (Lepidoptera: Pyralidae). Annual Report
of the Entomological Society of Ontario 83: 2\—25.
McClanahan, R. J., Boyce, H. R. and Code, W. R. 1968. The cereal leaf beetle — A new insect in
Ontario. Proceedings of the Entomological Society of Ontario 98: 21-26.
McLeod, J. H. 1962. Part I — Biological control of pests of crops, fruit trees, ornamentals, and
weeds in Canada up to 1959. Pp. 1-34 in McLeod, J. S., McGugan, B. M. and Coppel,
H. C. (eds), A review of the biological control attempts against insects and weeds in
Canada. Technical Communication Commonwealth Institute of Biological Control 2,
Franham Royal, Bucks, England, 216 pp.
McLeod, J. S., McGugan, B. M. and Coppel, H. C. (eds) 1962. A review of the biological
control attempts against insects and weeds in Canada. Technical Communication
Commonwealth Institute of Biological Control 2, Franham Royal, Bucks, England,
216pp.
McMullen, R. D. 1971. 16. Psylla pyricola Forster, pear psylla (Hemiptera: Psyllidae). Pp. 33—
38 in Kelleher, J. S. et al. (eds), Biological control programmes against insects and
weeds in Canada 1959-1968. Technical Communication Commonwealth Institute of
Biological Control 4, Franham Royal, Slough, England, 266 pp.
Miller, L. A. and De Lyzer, A. J. 1960. A progress report on studies of biology and ecology
of the six-spotted leafhopper, Macrosteles fascifrons (Stal), in southwestern Ontario.
Proceedings of the Entomological Society of Ontario 90: 7-13.
Monteith, L. G. 1976. Laboratory feeding studies of potential predators of the apple maggot
Rhagoletis pomonella (Diptera: Tephritidae) in Ontario. Proceedings of the
Entomological Society of Ontario 106: 28-33.
Monteith, L. G. 1977. Field studies of potential predators of the apple maggot Rhagoletis
pomonella (Diptera: Tephritidae) in Ontario. Proceedings of the Entomological Society
of Ontario 107: 23-30.
Monteith, L. G. 1978. Additional records and the role of the parasites of the apple maggot Rhagoletis
pomonella (Diptera: Tephritidae) in Ontario. Proceedings of the Entomological Society
of Ontario 108: 3-6.
Moore, R. J. 1975. The biology of Canadian weeds. 13. Cirsium arvense (L.) Scop. Canadian
Journal of Plant Science 55: 1033-1048.
Morris, G. W. 1980. Microbial isolations from Sitophilus zeamais (Coleoptera: Curculionidae).
Proceedings of the Entomological Society of Ontario 110: 93—96.
Mulligan, G.A. and C. Frankton. 1954. The plumeless thistles (Carduus spp.) in Canada.
Canadian Field Naturalist 68: 31—36.
Murillo, H., Hunt, D. W. A. and Vanlaerhoven, S. L. 2012. Larval parasitoids of the cabbage
looper, Trichoplusi ni (Lepidoptera: Noctuidae), in field tomato crops in southwestern
Ontario. Journal of the Entomological Society of Ontario 143: 115-119.
Nealis, V. G. and Bourchier, R. S. 1995. Reduced vulnerability to hyperparasitism in nondiapause
strains of Cotesia melanoscela (Ratzeburg) (Hymenoptera: Braconidae) Proceedings
of the Entomological Society of Ontario 126: 29-35.
Nealis, V. G. and Quednau, F. W. 1996. Canadian field releases and overwinter survival of
Ceranthia samarensis (Villeneuve) (Diptera: Tachinidae) for biological control of the
gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae). Proceedings of the
Entomological Society of Ontario 127: 11-20.
60
Biological control in Ontario, 1952-2012 JESO Volume 144, 2013
Nystrom, K. L. and Evans, H. J. 1989. Biological notes on a birch leaf edgeminer, Scolioneura
betuleti (Hymenoptera: Tenthredinidae), new to North America. Proceedings of the
Entomological Society of Ontario 120: 17-24.
Pengelly, D. H. 1961. Thymelicus lineola (Ochs.) (Lepidoptera: Hesperiidae) a pest of hay and
pasture grasses in southern Ontario. Proceedings of the Entomological Society of
Ontario 91: 189-197.
Phillips, J. H. H. 1969. Parasitism of a population of the Oriental fruit moth, Grapholitha molesta
Busck, in an unsprayed peach orchard. Proceedings of the Entomological Society of
Ontario 99: 81.
Philogene, B. J. R. and Chang, J. F. 1979. New records of parasitic chalcidoids of pear psylla
(Homoptera: Psyllidae) in Ontario, with observations on the current world status of its
parasitoids and predators. Proceedings of the Entomological Society of Ontario 109:
53-63.
Poinar, G. O. Jr., Thomas, G. and Prokopy, R. J. 1978. Microorganisms associated with Rhagoletis
pomonella (Diptera: Tephritidae) in Massachusetts. Proceedings of the Entomological
Society of Ontario 108: 19-22.
Pointing, P. J. and Green, G. W. 1962. A review of the history and biology of the European pine
shoot moth, Rhyacionia buoliana (Schiff.) (Lepidoptera: Olethreutidae) in Ontario.
Proceedings of the Entomological Society of Ontario 92: 58-69.
Proverbs, M. D. 1962. Progress on the use of induced sexual sterility for the control of the
codling moth Carpocapsa pomonella (L.) (Lepidoptera: Olethreutidae). Proceedings
of the Entomological Society of Ontario 92: 5—11.
Putnam, W. L. 1963. The codling moth, Carpocapsa pomonella (L.) (Lepidoptera: Tortricidae):
A review with special reference to Ontario. Proceedings of the Entomological Society
of Ontario 93: 22-60.
Ragsdale, D. W., Voegtlin, D. J. and O’Neil, R. J. 2004. Soybean aphid biology in North America.
Annals of the Entomological Society of America 97: 204-208.
Ramey, C. A. 1990. Host identification and oviposition behaviour of a parasitic wasp, Eurytoma
obtusiventris (Hymenoptera: Eurytomidae). Proceedings of the Entomological Society
of Ontario 121: 127.
Reid, D. G. and Harmsen, R. 1975. Trirhabda borealis Blake (Coleoptera: Chrysomelidae) a
major phytophagous species on Solidago canadensis L. (Asteraceae) in south-eastern
Ontario. Proceedings of the Entomological Society of Ontario 105: 44-47.
Robinson, A. G. 1952. Annotated list of predators of tetranychid mites in Manitoba. Annual
Report of the Entomological Society of Ontario 83: 33-37.
Robinson, A. G. 1953. Notes on Stethocorus punctum (LEC.) (Coleoptera: Coccinellidae), a
predator of tetranychid mites in Manitoba. Annual Report of the Entomological Society
of Ontario 84: 24-26.
Sharkey, M. 2007. Revision of the Neotropical genus Trachagathis Viereck (Hymenoptera:
Braconidae: Agathidinae). Journal of the Entomological Society of Ontario 137: 5\—
61.
Song, S. 1990. The effect of host encounter rate on fecundity of Gelis tenellus (Hymenoptera:
Ichneumonidae). Proceedings of the Entomological Society of Ontario 121: 129.
Stewart, J. G., Lund, J. E. and Thompson, L. S. 1992. Factors affecting the efficacy of Bacillus
thuringiensis var San Diego against larvae of the Colorado potato beetle. Proceedings
of the Entomological Society of Ontario 122: 21-25.
61
Mason JESO Volume 144, 2013
Thistlewood, H. M. A., Bostanian, N. J. and Hardman, J. M. 2013. Panonychus ulmi (Koch)
European red mite (Trombidiformes: Tetranychidae). Pp. 238-243 in Mason, P. G. and
Gillespie, D. R. (eds), Biological Control Programmes in Canada 2001-2012. CABI
Publishing, Wallingford, Oxfordshire, United Kingdom, 518 pp.
Tripp, H. A. 1972. Field trials to control spruce budworm, Choristoneura fumiferana (Clem.),
through aerial application of Bacillus thuringiensis. Proceedings of the Entomological
Society of Ontario 102: 64-69.
Tyler, B. M. J. and Ellis, C. R. 1980. Ground beetles in three tillage plots in Ontario and
observations on their importance as predators of the northern corn rootworm, Diabrotica
longicornis (Coleoptera: Chrysomelidae). Proceedings of the Entomological Society of
Ontario 110: 65-73.
Vander Hoek, G. 1971. The larval instars of Aphidius pulcher Baker (Hymenoptera: Aphidiidae).
Proceedings of the Entomological Society of Ontario 101: 55—59.
Van Whervin, L. W. and Wilde, W. H. A. 1970. An analysis of the possibilities of the sterile male
release technique in the eradication of the surgarcane moth borer, Diatraea saccharalis
(F.), in Barbados West Indies. Proceedings of the Entomological Society of Ontario
100: 111-113.
Villanueva, R. and Harmsen, R. 1996. Ecological interactions of tarsonemid mites in apple
orchards: Predation of apple rust mite and use of Phyllonorycter blancardella mines.
Proceedings of the Entomological Society of Ontario 127: 99-106.
Vincent, C., Huber, J. T., Gibson, G. A. P. and Goulet, H. 2013. Phyllonorycter blancardella
(Fabricius), spotted tentiform leafminer (Lepidoptera: Gracillariidae). Pp. 244-247
in Mason, P. G. and Gillespie, D. R. (eds), Biological Control Programmes in Canada
2001-2012. CABI Publishing, Wallingford, Oxfordshire, United Kingdom, 518 pp.
Wallace, D. R. and Sullivan, C. R. 1985. The white pine weevil, Pissodes strobe (Coleoptera:
Curculionidae): a review emphasizing behavior and development in relation to physical
factors. Proceedings of the Entomological Society of Ontario 116 Supplement: 39-62.
Wang, T. and Laing, J. E. 1989. Reproductive biology of Holcothorax testaceipes (Hymenoptera:
Encyrtidae) and its effect on development of the host, Phyllonorycter blancardella
(Lepidoptera: Gracillariidae). Proceedings of the Entomological Society of Ontario
120: 35-41.
Wang, T. and Laing, J. E. 1990. Toxicity of selected insecticides to adult Holcothorax testaceipes
(Hymenoptera: Encyrtidae), an imported parasitoid of Phyllonorycter blancardella
(Lepidoptera: Gracillariidae). Proceedings of the Entomological Society of Ontario
121: 119-122.
Welch, H. E. 1958. Test of a nematode and its associated bacterium for control of the Colorado
potato beetle Leptinotarsa decemlineata (Say). Annual Report of the Entomological
Society of Ontario 88: 53—S4.
Welch, H. E. 1962. Nematodes as agents for insect control. Proceedings of the Entomological
Society of Ontario 92: 11-19.
Welch, H. E. and Briand, L. J. 1961. Field experiment on the use of a nematode for the control
of vegetable crop insects. Proceedings of the Entomological Society of Ontario 91:
197-202.
West, A. S. and DeLong, B. 1956. Notes on the biology and laboratory rearing of a predatory
insect, Zelus exsanguis (Stahl) (Hemiptera: Reduviidae). Annual Report of the
Entomological Society of Ontario 86: 97-101.
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West, R. J. 1992. Notes on the biology and control of black army cutworm, Actebia fennica
(Lepidoptera: Noctuidae), in black spruce plantations. Proceedings of the Entomological
Society of Ontario 122: 53-63.
West, R. J., Laing, J. E. and Marshall, S. A. 1984. Parasitoids of the potato stem borer, Hydraecia
micacea (Lepidoptera: Noctuidae). Proceedings of the Entomological Society of
Ontario 114: 69-82.
Whistlecraft, J. W. and Lepard, I. J. M. 1989. Effect of flooding on survival of the onion fly
Delia antiqua (Diptera: Anthomyiidae) and two parasitoids, Aphaereta pallipes
(Hymenoptera: Braconidae) and Aleochara bilineata (Coleoptera: Staphylinidae).
Proceedings of the Entomological Society of Ontario 120: 43-47.
Whitfield, G. H., Drummond, F. A. and Haynes, D. L. 1981. A simulation model for the survival
and development of the onion maggot Hylemya antiqua (Diptera: Anthomyiidae).
Proceedings of the Entomological Society of Ontario 111: 39-55.
Wilde W. H. A. 1965. The pear psylla, Psylla pyricola Foerster, in Ontario (Homoptera:
Chermidae). Proceedings of the Entomological Society of Ontario 95: 5-10.
Wilson, G. G. 1978. Observations on the incidence rates of Nosema fumiferanae (Microsporidia)
in a spruce budworm, Choristoneura fumiferana, (Lepidoptera: Tortricidae) population.
Proceedings of the Entomological Society of Ontario 108: 144-145.
Wilson, G. G. 1980. Effects of Nosema disstriae (Microsporida) on the forest tent caterpillar,
Malacosoma disstria (Lepidoptera: Lasiocampidae). Proceedings of the Entomological
Society of Ontario 110: 97-99.
Wilson, G. G. 1981. Effects of Nosema fumiferanae (Microsporida) on rearing stock of spruce
budworm, Choristoneura fumiferana (Lepidoptera: Tortricidae). Proceedings of the
Entomological Society of Ontario 111: 115-116.
Wilson, G. G. 1985a. The transmission and effects of Nosema fumiferanae and Pleistophora
schubergi (Microsporida) on Choristoneura fumiferana (Lepidoptera: Tortricidae).
Proceedings of the Entomological Society of Ontario 115: 71—75.
Wilson, G.G. 1985b. Dose-mortality response of Choristoneura fumiferana (Lepidoptera:
Tortricidae) to a microsporidium, Pleistophora schubergi. Proceedings of the
Entomological Society of Ontario 115: 93-94.
Wilson, G. G. 1987. The effects of Vairimorpha necatris (Microsporida) on the spruce budworm,
Choristoneura fumiferana (Lepidoptera: Tortricidae). Proceedings of the Entomological
Society of Ontario 117: 91-93.
Wilson, G. G. and Burke, J. M. 1979. Microsporidian parasites of Archips cerasivoranus (Fitch)
in the district of Algoma, Ontario. Proceedings of the Entomological Society of Ontario
109: 84-85.
Woolhouse, M. E. J. and Harmsen, R. 1985. The mite complex on the foliage of a pesticide-
free apple orchard: population dynamics and habitat associations. Proceedings of the
Entomological Society of Ontario 115: 1-11.
Wright, E. J. 1979. Observations on the copulatory behaviour of Perilitus coccinellae
(Hymenoptera: Braconidae). Proceedings of the Entomological Society of Ontario
109: 22.
Wright, E. J. and Laing, J. E. 1979. The effects of temperature on development, adult longevity
and fecundity of Coleomegilla maculate lengi and its parasite Perilitus coccinellae.
Proceedings of the Entomological Society of Ontario 109: 33-47.
Wright, R. E. 1972. Nematode parasite of the face fly in Ontario. Proceedings of the Entomological
Society of Ontario 102: 168-175.
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69
JESO Volume 144, 2013
Mason
(8/861) SUA 2 9005 proyisesed deplpeulolaig eioydouswA}] JOH[LA\ SLDB/NA 19JSPBO]IAD
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IUIIIJIY=—- PYOIU SUIPIIy INTO | 1IPAO AUWOUD [BANJEN 1SOH]
“penuyuos V XIGNdddV
70
144, 2013
JESO Volume
Biological control in Ontario, 1952-2012
(696|) UoIoOWR)
(996]) 29A0g 2 uRIsnq
(82861) SHIA 2 23005
(8L861) SHIA 2 91005
(BL861) SHIA 3005
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(BL861) SHIA 93005
(82861) SHIA 2 93005
(82861) SHIA 2 91005
(8861) SIA 2 91005
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““penuyuos VW XTGNadddV
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JESO Volume 144, 2013
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(Z661) Joqieg 2 Aa[sey proyisesed oeprulyoey, viaidiq yayinbod vj9a1a psav0104g
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Biological control in Ontario, 1952—2012
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13
JESO Volume 144, 2013
Mason
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1SOH
““penuyuos VW XIGNaddV
74
144, 2013
SO Volume
4
4h
JI
Biological control in Ontario, 1952—2012
(6L6|) SueyD 2 auasojiyg
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““penunuos V XIGNaddV
75
JESO Volume 144, 2013
Mason
(6261) Suey 2 euesojiyd
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““penuljuos VW XIGNdAddV
76
144, 2013
‘SO Volume
4
4
JI
Biological control in Ontario, 1952-2012
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““penuljuos V XTGNdddV
(el
JESO Volume 144, 2013
Mason
(Z66 |) Joqieg 2p Ao[seyH ployisered oepneuueidsoyouy eiojdouawiAY AQ] RY] WYNUIUL DUD ABOYIIAT,
(7661) Joqieg 2 Ase prouseied aepluowmnouy9s] eiojdouawAY BINGIZILY SNLMVIISAAA SAQUIDIG
(7661) Jaqueg 2 Aa[seyH proyiseied oepruoumnouyo] piajdouswAY Jayouraolg sijpnbav vjduig
(Z661) Joqueg 2 Aasey proqseied oepruoumMauysy] viajdouawAY UOSSAID SLUDSINA snJalpojAyd
(7661) Joqieg 2 Aa[seyH prousesed aepruowmouyo] viojdouswAY ‘ds snjaipojAyg
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[(Youaty) suadsaplldl SNAJUAIOAIDY =]
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(7661) taqueg 2 Aq[sey plowseied oepruoumnsuyo] eiaydouswAY (Aes) “ojisinbuos sysajdoqy
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(7661) taqueg 2 Aq[sey prouseied aeprydojng eiojdousw AY ‘ds snjsayov] gy
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(7661) Joqueg 2 Aajsey prousesed oepruryoe | elaidiq ueling vIdnsiajul DIOP
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““penuyuos V XIGNaddV
78
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duatIay aydiu SsuIpsoq AqTUue JIp1O AWIIUD [BNIB 1SOH
~penunioes v7 XTGNaddVv
Biological control in Ontario, 1952—2012
79
JESO Volume 144, 2013
Aason
(€96]) weuing Joyepoid aepnsduy SoULOjIpIquioly, syurg sijisp sysdup
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““penuyuos V XIGNdddV
82
‘SO Volume 144, 2013
4
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Biological control in Ontario, 1952—2012
(O861) SUI 2 JQJAL Joyepoid oepiqeie) viajdoajop (Aes) snouysna snjAjopposiup
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““penunuos V XIGNaddV
83
JESO Volume 144, 2013
Mason
(961) BuO Taq 7 ISAM
(9561) s0qhey
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(WOYyIOUURI\\) DUAL) DINOAGDIG
1SOH
““penuyuos WV XIGNHddV
84
SO Volume 144, 2013
JE
Biological control in Ontario, 195
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1SOH
““ponurjuos VW XIGNdAddV
85
JESO Volume 144, 2013
Mason
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(P61) Te 19 ISAM
(PR61) “1B 19 ISAM
(F861) Te 19 ISOM
(ZL61) MON 29 ULOT
(ZL61) MON 7B UOT
SO Volume 144, 2013
(ZL61) MON 7 ULOT
JE
(TL61) MON 2 ULOT
(ZTL6O1) MON 2 UOT
(ZL61) MAN 2F ULOT
(TL61) MON 2 UOT
(ZL61) MON 29 UROT
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IIUITIJIY
Biological control in Ontario, 1952—2012
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viajdouswiAy
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eiajdouswiA}]
eiajdouswiAp]
viojdouswiA}{
viajdouswiAy
viay}douswiA}]
viajdouswA}]
eiajdouswA}{
epuipqeyy
viojdouswiA}{
eiajdouswiA}]
eiaj}douswiAy]
IIPAO
‘ds pusapviq
JNVILD WNASNIO] Wau DAOpOsjUaD
‘ds syajodup)
[yoaqasanA] apupusauuos vyjal4oydny=|
(paqasany\\) apupuaUos UoLydo1aT
[MON 2 UOT s1uIsHpI/OS DjJaL4oydny=|
(MON 2 UvOT) SIuIsDpI/OS UoAYdolaT
[MON a UOT YIDv{ipyjod vjjal4oydny=|
(MON 2 URO7) BIOv{ipyjpd UuosydolaT
[yoaqasany| vayiond pjyjaisoydng=|
(yoaqasony\) DaYiond uosydo1aT
[MON a uvoT voxIU pjjalsoydng=|
(MON 2 URvOT) 1OxXIU UOsYydo1aT
[MON 29 UOT SisuasppH] Dy Pjjal1oydny=|
(MON 2 UROT) SisuadppHjpy UosAYydo1aT
[peouysy pi4a0u1 pyjalsoydng=|
(peowysy) pjaoul uorydolaT
[MON
w uro’7T siploosdojpdy vjjal4oydny=|
(MON 2 URO'T) sIpL20sdojpAy UoAydolaT
[MON a uvOT 1synNof Pjjal4oydny=|
(MON 2p UROT) Isinof UoAydolaT
[MON
3 uLvOT lajppl49 vjjalsoydng=| (MAN 2
ueo7) lajppl4o (pyjatdoydng) uoaydoiaT
(9€1dq) ‘ds ‘u punjrajdpoay
Ad] IY WNSOladd DULUDABOYIIA
[AoyouRAoIg snpnoop
pIsajo)=| (AYIURAOIg) DPNDID VISAIOD
YODA, SISMOPLIBIU SNAJUBIOAID,
(oepinjooN :esojdopida7)
(dodsq) paavoim pIaapApAL]
87
(oepio0sq :e.1ajdoo0sq)
(4oy[BA\) SNIDLYS SNIOSdoO]PAL]
(aepinjoon :esojdopida7)
[(sniotiqey) suaasauia siysoyapy=|
(snioigeq) suaasadia DdAaAolay]
(oepinjooN
:e1aydopidaq) ‘dds srysorayy
(aeprloyioy :v.1aidopidaq)
(YoMeB}]) DUdAAfiqnu DApAaL]
(aeprlopoy :e1ojdopida7)
(49]|[97Z) Diasouolys vApaT]
JSOH
“penunuos VY XTIGNdddV
JESO Volume 144, 2013
Mason
(7661) SUA
unosieH (9/61) SHIA 2 NQV
(Z661) SIA 2 Noose Hy
(7661) 19qnH
(9007) 4990H
(7661) 4990H
(7661) 19q0H
(9007) 4940H
(7661) 9q0H
(9007) 9q0H
(9007) 49qnH
(7661) 49q0H
(9007) 42q0H
(F861) ‘Te 19 ISAM
(F861) Te 19 ISAM
(P8361) Te 19 ISAM
(F861) ‘Te 19 ISOM
(F861) ‘Te 19 ISAM
(F861) ‘Te 19 ISAM
(P61) ‘Te 19 ISAM
(F861) Te 19 ISAM
(F861) ‘Te 19 ISAM
(F861) ‘Te 19 ISAM
(F861) ‘Te 19 ISOM
Jud1IJOY
proyisesed
proysesed
proytsesed
prousesed
proqrseied
proqiseied
proqrsesed
proqrsesed
proyiseied
prowsesed
proyrsesed
plowseied
proyiseied
proqseied
proqisesed
proqsesed
proqisesed
prowsesed
plowsesed
plouseied
proyseied
proysesed
ploysesed
dYoiu SUIpIIy
oepruowmnouys]
aepruownouysy
oepuewdyy
oepuewdAy|
oepuewAyy
oepluewAyy
oepuewdAy
oeplewAyy
oeplewAyy
oeplewdAyy
oepuewAy\
oepuewdyy
depHeUIWeISOYSIL,
aepruowmouyd]
SEPIUOI[A9G
aepruowinouys]
aepiuoorig
oepluoovig
aepruryor |,
seprulyoe
depruowimnouyd]
oepruouimnouydy
aepruownouys]
INTO ae
viajdouswAy
viajdouswiAY
eiojdouswAH
pia}douswiAY
eiajdouswiAy
eiajdouswAY
eiajdouswiAp
eiajdouswiAy
eiajdouswAY
eiojdouawAY
eiajdouswiA}
eiajdouswAY
elajdouswApY
elajdouswiAY
eiajdouswiAY
eia}douswAY
eiajdouswAY
eiajdouswiAY
viaqdiq
elaqdiq
eiojdouswAp
eiajdouswAy
eiajdouawAp
Reyne)
(uoswioy]) sivolnoind sajsajddyog
(uoswioy |) Snanup sajoajddyjog
yNeIH snpjaisiu saydoupy
(nei) puny saydoup
Aepey siuuadiosnf saydoupy
yNeIH snjjaisiu saydpup
yNeIIH yayov.ouos saydoup
yNeILD snjjaisiu saydvupy
(ney) vuny saydpup
(ney) puny saydpupy
JNCILD snjjaisiu saydpup
(ney) puny saydoup
(ney) Wnpi4ts10Jad DUWDABOYILAT,
‘ds uolsay
‘ds snuiouajay
‘ds snuisor01a}
(SOON) SNWAYUL SNAJUBIOAIDV,
yle[D 2 Apeq snpunjg snajuas0.ovyy
(uadlay)) suDjngnys KpjapAaT
(pudsuMO]) S1OIpY4 B]JAPAT
‘ds nid}.
JSIOYUDARID 4OJDdnNII0 saubydaxy
‘ds snuidq
AW9Ud [BANJEN,
(aepiuolnoimy :e1aj}doajoD)
({yeua|[AD) voysod piadAy]
(oepiuolnoinyD :e1ajdoajog)
Jaw vjooipnjod p1iadAH
(aepruolnoin|D :e1ajdoajoD)
(SnIoLiqe{) SLysouisiu d1adap]
(aeprluolnoiny :e1ajdoajoD)
(a4UOge7T) DIMIXAa DiadAL
(aepiuolnoinyg :e1ajdoajo))
[uewysog snyuiuo snuog=|
(ueWlayog) smjiui4o vAadAL
(oepluolnoinyD :e1ajdoajo))
(Aes) piduoo piaday
(aepluolnoin, :e1a}doaj02)
(uewayog) s1uuadiauunsg DiadAy
1SOH
““penunuos ¥ XIGNaddV
88
144, 2013
SO Volume
4
vi
JE
Biological control in Ontario, 1952-2012
(9661) 143g
(9661) 14811
(9661) 1UsUg
(P861) TE 19 ISAM
(49661) a8eS2T
(9007) 49q0H
(9007) 49q0H
(9007) 42qnH
(9007) 49q0nH
(7661) 4990H
(Z861) ueOT
(T6611) SHIA 2 Nose
(0661) sanber 2p Aa,a7z7-ueg
(Z861)
uBOT (7861) ‘Je 19 UNOsIeY
(ZT86]) ueOT
(T8611 :O861) ‘Te 19 UNosIeH
(0661) senber 2 A2,97-uag
(0661) senber 2 Aa,a7-uag
(O61) Je 19 uNOSIe}]
(0661) Sanber 2 Aa,az7-uag
(0661) sanber 2p Aad,a7-uag
ERTIENCIEDS|
ployiseied
proyseied
proqiseied
proqisesed
aseydoyAyd
ploytseied
proqisesed
ployisesed
proyseied
ploytseied
uasoyjed
uosoyjed
uasoyjed
proytsesed
ployiseied
uosoyjed
uasoyjed
uosoyjed
uasoyjed
uasoyjed
dyoiu SUIPIdy
depl[ewlold}g
oepiuoorig
oepluooeig
oepluoseig
oepljowosAry)
aepuewdyy]
oepliewAyy
oepluewAyy
oepluewAyy
oepuewAy\
aevoorloyydowougq
ovaovioyjydowojuq
seaovioyjYydowojuq
aepiuoseig
oepiuoorig
aeaovioyjydowojuq
aevaovioyjydouojuq
aeoorioyiydouojuq
aevaovioyydowojuq
avaoeI [UOJ
jruey
eiajdouswiAy
piajdouswiAy
viajdouswAY
eiajdouswiAY
eiaidoajod
eiajdouswAY
eiajdouswiAy
eiaj}douswiAp
piajdousw AH
eiajdouswiAy
eiajdouswAY
piajdouswiAY
IPO
(4oy[BA\) Pjainy snoypdoyy
(peowysy) sipossid sapioja0y
(peawysy) sipossid sapioja0)
Ye] 2 Apeg snpunjqg snsjuaso.vp
(49}8104) 1o14addy purjosdy)
(Qy[neIy) pun; saydouy
Qyneiy) puny saydpup
QynetD) puny saydoupy
(Q[neiD) bun) soydoupy
Aepley siuuadiosn{ saydoup
‘dds paoyjydooz
omeg (anyy) uouojdyd vsoyjydooz
TysMmazoer 1uoUuojAYyd wniydLvdy,
ROI 18a]09
SNUOJIOAIY =| (WIIG) 18A]0I sn4jip1dad
ueo7] sapiodoiyjap
SNUOJIOAIY =| S99N Sdolyjav snjyi4ag
‘ds piudsq
YOUU] 2? AI,9Z
-uog “Joquiny (anyry) muouojAyd viudag
Inyy 1uouojdyd psoyjydowojuq
DISMOGIeH
pywjound (wniyaiavny ) pAoysydowojugq
UIWAT[INA (OWRS|eg) VUDISSDg DIdaAnDag
Auwoud [B.1NjeN)
(osepruolnoiny
:e19}d0a]09) (Aes) mid sdy
(aepruolnoin, :e1ajdoa]o))
(yoyyoig) s1jjooipunss sdj
(sepruolpnoing :e1a}doa]0))
(aeULIOD) s1ydp1s81jp9 sdy
(oepInjooN :e1ajdopida7)
Aepalqnog siispjad pizavsdyy
(apaopiiadAy])
“7 wnypsofaad wnoaisaday
(oepluolnoinZ :e19}do0a]0))
yodoog snj1oz viadAy]
(oepluolnoin, :e1ajdoajoD)
ISQIOH SIjigo1ivAa DAéadAy]
(sepruolpnoing :v1a}d0a]0D)
weysieyl Vypauipia) DAadAL]
(sepruorNoinD :e1ajdoajog)
(snioliqe.) vipjound paadAy]
1SOH
~"penulyuos V XTANdaddV
89
JESO Volume 144, 2013
Mason
(9007) 2eqnH proyisered aepliewAy| viajdouswiAY (qnoq) vsnydosisias saydoup (aepysay :ejeuopd) “ds sajsaT
[suing sadyjod
uoaydoiaT= (suing) sadyjod snuajsiag
jou ‘(UOssaID) Sadijjau snuajsi4ag=| (sep :e1aydrwioy})
($96) uROT ployisesed oepluoorig piajdouswAY (uossaiD) sadyjau uoiydo1aT (J) vipaqojop vusajdojdaT
(7961) YIP
(1961) pueug 2 YoTEM ayiseied IEPNLUWISUIIUII}S epuipqeyy (9¢1dq) ‘ds ‘u vunj2ajdpoan
(S61) U91EM ayisered = 9ePRUWI9UIOUTI}$ epuipqeyy JauIa}g DUIsa4yo DUD_JIa]dvOaN,
[[[@ssuH lapynd
(0661) ‘[e 19 URSILUOD proyisered oepiydojng eigjdouswiAY = wnaopy=] (]]OSSuD) Mapund snuawus14opy
(aeprpawiosAry, :e1a}do0a]0_)
(7661) Je 19 UeEMAIS uasoujed avaoeyloeg Joulpiag sisuaisuniny) snjyjloog — (Keg) pipauljwacap vsavjoundaT
(aeprjowiosAry, :e1a}doajo))
[(Aes) DypyiALy DyDaUIp1A] DUAT=|
($661) J0qnH ployiseied oepuewAyy eiojdouawiAY (4938104) Sadiapyf saydvupy (Ae) DIDYIALA] DIDI1ALL] DUT
(aeprjauosAryS :e1a}do0a[0D)
(€661) JoqnH proyiseied oepluewAy piajdouawA}] (19}8104) sadiavyf saydoup (IAQ) DIDAUIILA] DUAT
(aepljauosAryD
(7661) 20qnH ployisered oepuewAyy piojdouswAH (19}810,4) Ssadianyf saydoup —:8.19\d0a]0D) 0A SIMaYyoI] DUAT
(aepljauosAry)
(7661) 49qnH proysered aepuewAyy| piajdouawAY (4018104) Sadiapyf saydvup — :e1aydoajod) (°7) pyjaundo vuaT
(aeprjawosAry, :e1a}doaj0D)
[(sntolige q) DIDYIA DIYOAGvIG=|
(0961) pueug oyisered = oepyeuoUOjURI[y epryouajAL, qqog vs8uiuag DjnpinMoy (SNIOLIge J) DIDI1A DIGAT
(oeprlolwu0 jf
(S861) Aier9H ploseied oepnieyong viojdouawAY (peowysy) 1zZioMyos vaspvjawopnasg :e1ajdouswiAp) “ds snisp7
(Qeprorwi0; :e1a}doawA})
(S861) AeI0H proyiseied oepniueyong piajdouswiAy (peowysy) uzioMyos vpaspjaumopnasd AIOWIY 4931UOIU SNISDT
[sing sadyjod
uosydolaT= (sing) sadyjod snuajsisag
jou {(uossaiD) sadijjau snuajsisag=| (aepliyyy
(S96) ueOT ployisesed oepluosrig piajdousw A} (uossaiD) sadiyjau uosydoiaT :e1aydiwiay) wystwy snjiy sdoqvT
(aepruolnoing
(9661) 1UsLIg proysesed depl[RUIOIA} eiojdouswiAY (prowysy) 1Moj201puap snosyoulq :e1ajdoajop) ‘dds sdy
VUdIIJIY=—-- YIU SuIpIoq INTO | JIP1O AWOUd [B.1NjBN }SOH
““ponuyuos V XIGNdddV
90
SO Volume 144, 2013
vie
JE
Biological control in Ontario, 1952-2012
(6661) ‘Je 19 JUaqpeolg
([L0Z) ‘Je 19 UOse|
(6661) ‘Je 19 JUaqpRo1g
(1107) ‘Ie 19 UoseW -(6661)
‘ye 19 uaqpeolg {(S96]) ULOT
(TLOZ) [2 39 Uose|
(6661) ‘JB 10 JUaqpRo1g
(T10Z) ‘Je 18 UoseYy
(6661) ‘12 19 Jaqpeoig
( I 107) ‘ye Jo UOSe|Y]
(1107) ‘Te 19 Uose
*(6661) ‘Te 12 JUaqpeoig
(T10Z) ‘Je 19 Uose
(L107) ‘[e 19 uoseyy
(110Z) ‘Je 19 Uoseyy
({10Z) ‘Je 39 UOse|
( 1107) “ye Jo UOSR|Y
(9007) 4990H
(4/861) SHIA 2 91005
(4/861) SHIA 2 91005
(7661) 49q0H
(7661) 1990H
IIUIAIJIY
proysesed
proyisesred
proqseied
proysesed
proyisered
proysered
proyiseied
Joyepoid
Joyepaid
Joyepoid
Joyepoid
Joyepoid
proyrtsesed
proyisesed
proysesed
proyiseied
proyisesed
dyotu SUIpIdy
aeprluoorig
oepruoovig
aepiuoorig
oepiuoorig
oepluoorig
aepiuoorig
oepluoovig
aeploooyUy
SEDIGEN|
SEDIGEN
aeplIod0ayH
aeplo0s0aH
oepluewAyy
aeprydojng
aeprydojng
oepluewAyy
oeplewAy|
Ayre y
piajdouswAY
eiajdouswiAY
eiajdouswiAY
elajdouswiAY
plojdouswiAY
piajdouswAY
eioydousuiAY
vio diay
eloidiwioy
playdiwioy]
era diwioy]
eiaidiwiaH
piojdouswiAY
eiajdouswiAy
piajdouswAY
viajdouswAY
vio}douswAyY
(ueyeo) siusofiun uosydoiaT
[ysiey] Ipp1uod snuajsl4ag=
(UOSWIOYL) S1//OI14gn4 snualsi4ag=|
(uosuIOY) s17joaluqna Uuosydo1aT
[sung sadyjod
uosydolaT= (Suing) sadyjod snuajsiiag
jou ‘(UOssa) sadijjawm snuajsisaq=|
(uossai)) sadyjau uosydolaT
(ueo7T) D10A18A] UOLYdoIaT
[uvoT] synauosip
snuajsi4ag=] (ueo'q) sinauosip uosydolaT
hejnoy
1App Snuajsi14aq=] (aN) Mop uosydoiaT
ueo7T sadyjodopnasd snuajsisag=|
(ueo7) sadijjodopnasd uosydoiaT
(UY) 40/091S814] SIAC
(uoAeIe)) snsafOI14alUD SIGDN
Aayysieg smjpu.iayjD SIGDN
(Aes) sadyound s11080a5)
[yes suappod si1090aH)
JNBIIL) 17AYIDAJOUOID saydpu V
(Qyjnemy) smipausajur snyddjsiq
Qynemy) saipausajur snydapsiq
Joqny saspydipiysidg saydpup
Jaqgny snspydipiyssdg saydpup
I3piO
AUWI9Ud [B.INJEN,
[(SloAnvag) s1“pjOaUT] S110901T=|
(Qosijed) sipjoaul) SNBAT
(oepliryy :e1oydiw9}])
IYsIuy sniadsay snBAT
(aepruorlnoin)
:B10}d09]09) (a3U0927T)
SISUDUOBALO SNJOUOAISIT
(oepizAwoisy :e1ajdiq)
[asouadg winspajofia) DZAWOLT=|
Joouads winivarjofid] DZAUWOLALT
(oepizAwoisy :e1ojdiq)
preyouryg apaos pzdwolsT
(oepiuAg :viaj}doajo_)
(ulaysouury|) sadiouduas uoory
(oeprludg :eiajdoajoD)
(Ays[nyos}oy]) Sp sns1]01T
}S0H
“penuryuos V XTGNAddV
|
oa)
i=)
= (9S61) BUD TAQ 7 ISOA\ Joyepoid oepnueyy piajdoyuo ({yRIS) siInsuDsxa snjaz
SE episeyd
—Te .
St (L861) UOSTIM, uosoyjed aepryeulasoN -o]deyrporsossiq (Jawery) x14jp2au DydsouLADs
o
E (F861) SOY 2 uasuLeH poniseied oepluowmouys] eiajdouswAY (paay) avdiunosoisya adups0qoyd
4 (O861) UOSTIA uosoujed aepleUlasON uosdwoy |] aviussip DUasSON
> (S96|) sn3uy uasoyjed avaor][loeg JOUT[IOg SisuaIsulLiny) snjploog
© [uose] SospwosoonjpU spsoy—| (aepidwesoise7 :e1ajdopida7)
is (P86) asoy 2 UssULIe}H proysesed oepiuoorig eia}dousw AY (uOsRI\)) SOJMWIOSODD]DUL Sapolaly Jouqny} VISsIp DUIOSOIDI DIV
= (aeprjepeoiy :ezajdiway} )
(0961) 49247 9d 2 J2TIIN proyriseied oepiulAiq viojdouswAY uojuay snisajd sndojpuosidq ([B1S) suosiospf sajajsosoDy
(aepluooRig
(€96]) Weujng proysesediadAy oeprujadng via}douswAY peowysy sdasiupds snujadng :e1aydouswA}) ‘dds snajuarosovyy
[(]) Sisuatapuppo DjjaonsajvH=|
(661) ‘[e 19 URSILIOD asvydojAyd oepljowosAiy) eiaydoajo) (J) SisualapUjDd Dj]JAIMNABJVBOIAN (@DAIDAYIJAT) “TT DILADIIOS UNAYIAT
(BSQ6]) UOSTIMA uasouied aeploydojsia}d JayJOMZ Iduaqnyos vsoydojsiajg
(6L61) SHIWJUD proysesed depruownouys] viajdouawAY uossaly sijppad vjduig
(2261) SYD proqrseied aeprlulyory viaidiq (Aploasaq-neouigoy) siji3p DUuasasvADg
(6861) sure] 2 Agpsey proyisesed oepiyidouq viajdouswAY (paeMOH]) avuvany snj4AoUua0—
[(Sinqazyey) smjaosouvjau sajajuvdy=|
(9461) SWIWGUD prowsesed aepiuoorig piajdouswAp] (Bnqazyey) Vjaasounjau vIsajoy
(S66|) Jaryoinog 2 sI[RaN proqseied aepluoorig viajdouawAY (Singazjey) Vjaasouvjau vIsajo)
(LL61) SwUJUDH ployisesed oepruryoe viaidiq (uasiay) DipUUIDUOD DAN[ISdUOD
(9661) Nepuand) x sITeaN proqisesed aepluryor |, viaidiq (QANSUDTIIA ) SISUAIDLUDS DIYJUDAID
(oepiqoiq :erajdopida7)
1yDISANY [J todsip vi.yayliog=)
(q ‘e966]|) ‘Je 19 WeysuluUND uasoujed avaor[loeg “IPAOIOS IOUT[IOG SISUAIBULINY) SNpPLODg (-q) todsip v1quowuArT
[sung sadyjod
uosydoiaT= (suing) sadyjpd snuajsisag
jou {(UOSsaID) Sadijawu snuajsiiag=|
(S96) UeOT proyiseied oepruoorig elajdouswAY (uossai)) sadyjau uosydoiaT
(7661) Joqny proqiseied aeplewdAyl viojdouswAY yNeRIIH ajor soydoup — (aepiayy :esajdiway) ‘dds snsa7
Udy —- BYU Sulpoay ATI IIP1O AWOUd [B.1NjB 1S0H
D
= ““penuyuos V XIGNdddV
92
‘SO Volume 144, 2013
4
4
JI
Biological control in Ontario, 1952-2012
(696) uoraUIeD
(TL61) SHUGUD
(696|) UoroWIeD)
(L861) ‘Je 19 WeysiuunD
(L661) O8epty 2 Jeqny
(0661) Senber 2p Aa,a7z7-uag
(S961) 21905
(S961) 21995
(S961) 21995
(ZL61) WSU
(€L61) USM 2p Alo3eI1H
(ZL61) SUM
(EL61) BUA 2 AIOSaIN
(ZL61) WS
(1961) Ajjesued
(7961) YIP
(686) Sule] 2 Suen
dIUIIIJOY
uasoyjed
prouseied
uasoyjed
uasoyjed
prowsesed
uosoyjed
proytseied
aseasip
prousesed
oyisesed
uasoyyed
oysesied
oisesed
proyserediadAy
oyiseied
proysesed
dou SUIpIey
appllAaojnavg
oepluowmnouyd]
aDplAlAO]NIDG
aDpldlAaojnoavg
aeplewAy
avoovlouYydowojwgq
SUMMA
appllAaojnang
aepruoseig
oepiiueynioevyds
avaovioyjydouojug
oepriuiepnioeyds
oepiiiepnioeyds
oepruowmoauydy
oepiyiAouq
Ayre
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eiojdouswiAY (Js1OYUSARIH) snuozispg snydojoalg
sndla sisoaspayAjod
(dN 4yMps auid pappaypas) SnAlAajuoraT
viodouswiAY (SuDyIag) 1/01Anpas sapouvydajs
nulog vuviuoyounjd vsoyjydowojugq
vlaqdiq ‘ds piuayluiy
[ds snarapurjassog=| “ds snaaojnopqvjag
eia}douswiAY ‘ds sajajupdp
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‘ds psoyjydowojug
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eiajdouswiAH (Qnemy) sadisojozg sidspiuasp
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(Aoypood) 4afijsas uolsdipoan
(aepruolidiq :e1o}douswiA})
[pupisyupg uolddipoan=] 1aMYOr
apupisyung yjpsd uoldipoan
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(YoI.4) 1a/U02a] uolsdipoan
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IeULIayH simsofisddd siqoN’
(aepiprydy :e1ajdiwiop)
(daz[NS) avaisiad snzApy
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pjoundiun pyajppnasd=|
(yIOMeH]) BJouNdiun DUAYIAY
(oeplosnyy :B1aqdiq)
JOY[eA\ DUISSIINJaA DISNW
(oeplosnyy :e1ajdiq)
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(ug][e@ 4) WnsO,4OY DILJ]2AAOPy
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[Aayry viguoyddy snsoajapy=|
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(aepliepioe1H :e1ajdopida7q)
uneig Dj/OI1UIXDAL DADULADIN.
}sOH
““penuyuos V XTANaHddV
93
JESO Volume 144, 2013
Mason
(oeprlolyoL
(7661) Joqleg 2 Ag[sey ployisesed oepruoumMauysy] piajdouswAY ‘ds snjsyotsy e1aydopidaq) ds sruwapung
(6861 °6L61) ‘18 19 SULA proysered oepiydojng pia}douawAY (tayyeM) synl snyoysp.ay
(7661) (aeprjawiosAry, :v1a}doajo))
Jogny ‘(6861) ‘Te 3° SIA proyiseied oepuewdy piajdousw A} (49}S104) sadianyf saydpup (J) sndouvjau pwajno
(aeprjawiosAryD :e1ajdoajo))
(7661) 29qnH ploysesed oepliewuAy| eiajdouswAH (49}819,4) sadianyf saydpup (uopAa}]) DUDIJaD]]Ds DUAa]NC
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(7661) J0qnH prouseied oeplewdAy eiajdouswAY (49}s104) Sadiavyf saydpup (Aes) siunjjoo pwuajng
(S861) sanboer 2 Sure] uasoyjed oepleuwloson (dower) XILp2au pYd.oULAiD
(Sg6|) sanboer 2p Sure] uasoujed oeplyeuUlasoN (o]ieg) vIsnvadd pulason,
(BSR6]) UOSTIM, uasoujed oepnewuason Jopualds 2 zn] sipiyjoljay DULAaSON,
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(aepiquies) :e1ajdopida7)
(S$Q6|) sanboer 2p Sure] uasouyjed aDpldlAaojnovg (AdNOV) AdN Dalusofijp2 pydbisonpy (aa9uqnH) S1/ppignu piul.ysCQ
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(aeproeqeseds :e1ajdoajo))
(7961) YO192M ayiseied = depIRUIDULIOUIAIS epuipqeyy Jautals Lasp]s punjoajdvoan’ (-q) sosaz0uiy.t S2JIAIC
(aepiquieid :e1aj}dopida7)
[aagping xajduus snqup1iyj=|
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(aepryeydwAy :esajdopida7)
(S96]) sn3uy uasoujed avaoryjloeg JOUIpIOg sisualsuLiny) snyplopg (-q) vdoyun sijpyduany
(aepl[ON :eraydopida7q)
[aevAq 2 uassowMNaN vUba/aUII
sndltyjoslswg =] 1ekQ 29
(Sg6|) sn3uy uosoyjed aevaory[loeg JOUI[og Sisuaisuliny] snpplopg UasaoWINaN VUYALIUII DJOAIIAN
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(C161) ‘[e 19 pug uasoyjed appldlaojnovg AdN Snala sisospaydjod
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# [suewiapng
= snupuayd (Snjniasoany) snwospolyday =|
= (€S6]) UeqiaH Joyepoid oepriasoyAyg RILUISIISOSaI (suewiapnd) snupuays snuos1polyday
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(€S61) WeqiaH Joyepaid oepiiasoyAyd PIBUISISOSa/\ (suewapng) snoipunjuy snuospojyday
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(€S61) HeqiaH Joyepoid oepliasoyAyd BILUIBIJSOSO[\ (uewuRed) snnaidsuod snwuospojydd],
(686) squawayD Jojepoid oepilasojyAyg RILLUSISOS|\ (Jajsnyos) supjsipnvs snuospojyday
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(€S61) HeqisH Joyepaid oepriasoiAyg RILWISISOSJ\| (syueg) siidosopu sniasojAyg
[(uewueED) sropjppf snuospojyday =|
(€S61) HeqiaH Joyepoid oepiiasoyAyg PILLS ISOS (uewueD) sropjjof snjniasoany
[suewiapng si4awnons snuos1polyday] =|
(€S6|) HeqiayH Jojyepoid aepliasoyAyd RILUISIISOS]A] (suewlapngd) sizawnond snjniasoayy
(S861)
USSULIe}{ A IsNoyjOoM, Joyepoid seploriyJAIq SouUOojIpIquio1y ‘ds wnysnpjpog
(726) Sure7 2 uesoped Joyepaid seploriyJAIq SouojIpIquMio1y, Ad|IwWg mupujnd wnysnojpg
(€S61) HeqieH Joyepaid oepliasoyAyg RILUIDISOSa ‘dds sniasdjqup
(oeplyoAuenay, :SOULOJIpIquiol| )
[(ydoy) 1ujn snyodub.4ajvjay=|
(6261) AqiseH Joyepoid depl[[euI9905 elajdoajoa (-q) bipjaundig vyopy (Yoox) Mujn snyodtuouvng
dUaIIJIYy«=—- BYU SuIpsaq Aqrure JIIP1O AWIOUD [B.INJe NY 1SOH
““penuyuos VW XTGNAddV
Biological control in Ontario, 1952—2012
95
JESO Volume 144, 2013
Mason
(LL6|) ‘[e 19 UOsuYyor proyisesed oeprydojng eiojdouswAY (JOx[VA\) SNINYS snigoIpag
[(poomjsany ) DsOMOf DjjadmYyI0sAyop=|
(LL6|) ‘Je 19 UosuYyor proyisesed oeprydojng viojdousuiApY = (poomysay) SMSOULLOf SIADYIOSAAYIOAN
[Qao19J\) Ssnyjaunjyyd snyoysp.ay—|
(LL61) ‘Je 19 UOsuYyoOL ployisesed oeprydojng eiajdouswiAY (JaosaJA\) Snyjaunjojd snyoysp4jajoulpy
[(Bingozyey) sdoyjupx snyoysp.ajay =|
(LL6|) ‘Je 19 UosuyoL proyisesed oeprydojng viojdouswAY (S99N) SIPUOL SNYISD.AJ2]OUL,
[(Bingazyey) /ajspsojoAo snyoysv.a] =|
(LL6|) ‘Je 19 UOsuYoL proyisesed oepiydojng eia}douswAY (SOON) SIppJUOL SNYIYSDAJAJOUY,
(686) Sule] 2 Suey proyisesed oepridouq eiajdouswAy (uewyed) wnyjajpouns Duosopidoy
(LL6|) ‘Je 12 UosuYyor proyisesed oeprydojng eiajdousw AY ‘ds sn4a90.4a]so].)
[(49}819,4) DIDIOSpfiun vapiowosodo}p=|
(LL61) ‘Je 18 UosuYyor ployisesed aeprydojng eiaydouswAY Joyem snoudy snpidsosuy
[4dyJeA\ SNIUd] Daplowosodojp=|
(LL61) ‘Je 19 UOsuYoL proyisesed oeprydojng piajdouswAY Joye snoudy snyidsosdy
(LL61) [eB 19 UOsuYor proyisesed oeprydojng eiajdousw ApH Ppoomysam SnUissyUDsaja SnjIdso1dy
(LL61) ‘[e 19 UOSsUYor ployseied oepiydojng eiajdouswiAY tysonjaq suagnd siupyoosdy)
[ojowlysoX “asspsopidsno siapysosduy)=|
(LL6|) ‘[e 19 UOsuYor ployiseied oeprydojng eiajdouawAp (dayx[eA\) SNasadau sLADYyIOSAAYD
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(LL6|) ‘[e 19 UOsUYoL proyisesed oeprydojng piajdouswAy (S9dN) Snaovjolo1siu sndvosdupg
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(686) Sule] 2 Surry proyiseied oepryidouq eiajdousw Ap (Binqazyey) sadiaanisaj sidspiuasp
(oepluey[lovsy :e1ajdopida7q)
[snioiqe| pyjapapoun)g
suayjor0yn T=] (sniotigeJ)
(LL6|) ‘[e 18 UOsuYor prowseied oepiydojng eiajdouswiAY [ds pusdpugq=] ‘ds saprospyoosAayopy DIJAaPADIUD]G AAJIAAOUO]]AY
(686|) Sule] a suey proyrseied oepiuoorig viajdouswAH ueyey apapuisoyjyd snuojay)
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(Z10Z) 4oqnH proytsesed oepluewAyy viajdouswiAY AepljeH smpsjna snuojo0—Q,
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(0661) sanber a Ao,a7-u9g uadoyjed ovoorsoyiydowojug (yjauUay 2 Ad,97-UNg) WYyaJad DIUAAT ((q) sniapunds snuavjiyd
(oepineyg :esaidoyud)
(9961) Aljasuag 2 sprempy ployisesed oeprydojng eiajdousw Ay peowysy uqdjoyo vIgoayy ((7q) puvoliaup pjauv)disad
dually: aYdtu Surpsaq ATIUB 1Ip1O AWI9UD [BANE] 1SOH
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=? (6861) Suey 2 Sur °(b861) (C1) srpsawmojs
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(L161) ‘Te 19 Uosuyor proqiseied oeprydojng eiajdouswAY (S99N) S1UAOIIAIIAAS SISAIdWUAG
(L161) ‘[e 19 UOsuyor proqseied aeprydojng viojdouswAY Joye SMipso38 sisaidudg
[aJNeIID sisuapunjAspu
(LL61) ‘Je 19 UOsUYoL ployisesed oeprydojng elajdouswAY = sisaiduiAg=] (194[eA\) SNIPs0S sisaidudg
(LL6]) ‘Te 19 UOsuyor proyisesed oeprydojng eiojdouswAY peowiysy “ajspsoouop sisaiduag
(LL6]) ‘Je 19 UOsuYyor proqtseied oeprydojng viojdouswAY (paeMmoyp) avinjdoin oypsiug
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(S861) URATTING 7 9eET|EA
(9661) 18g
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[(ys]eM) suosiqyp
(€96]|) UNoOdIe}Y provisesed oepiployeyD viojdouswiAy = szaypyooyidg=| (ys|eA\) suosfiqyp pAnuoy
(oepryjoin|_ :esoidopida7)
[(suang) simuadiynovu
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DUdAIIJIY «=: 9YTU Surpsay [rue yy rey re) WUD [B.1Nje 1SOH
~“penuryuos VW XTGNaddV
Biological control in Ontario, 1952-2012
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sIowi[eYyD pure rueyjayseD
(LL6|) ‘[e 19 JeUlod uosoyjed avaderidajoRqo1aUq ( e[NBIA\) 709 pIyoLaYyIST
(LL6|) ‘[e 19 JeUlod uosoyjed avaorliajoRqoiojUuq ‘ds 4ajopqosajugq
[ueyen snazjawu snidQ=|
(2261) yoo proytsesed oepluoovig eiojdouswAY (ueyen) vayjau pydiowmuspyovig
[ueyen snauisnssaf
(8261) yoo proqisesed aepluoorig viajdouswiAy snidcQ=] (ueyen) wnaursnisaf Duspyovig
[yooqasonyy snaojjD
(8261) yaiuop\ proyseied oepluoorig viajdouswiAY smid¢Q=] (yoaqasony\\) Winaoj]p DiuspYyovig
(LL61 9L61) WoO Joyepoid oepiqele) eiaidoajop (snioliqeq) wnapijp2 DULOSO]D.)
(LL6|) ‘[e 19 JeUloOd uasoyjed avaor][loeg pueyyuely wz purpyuesy svaiao snpplovg
(9007) 49qnH proyiseied oepluewdAyy viojdouswiAY] (peowysy) sadijjod saydpup
(9007) 4oqnH proyiseied oepluew Ay viajdouswA} yNeIID yayop.sjouos saydoupy
(6L61) Ae[8eH Joyepoid oepilasoyAyd RILLUDISOSO\ URWIRD s1onjjpf sniasdjquyp
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a QUdIIJIY-« AYOIU SuIpIody INTO 1IPIO AUWIOUD [BANE 1SOH
““ponuyuos V XIGNdAddV
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Biological control in Ontario, 1952—2012
(-L61) Noo
(PLol) HOoY
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1SOH
“ponuyuos V XIGNdddV
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(aeproAquieia, :v1a}doajo))
(9961) Ajjasuad 2 sprempy ployisesed oeprydojng piajdouaw Ay peowysy gdjoyo vigou1ayy sniolige| ppipuvs ppiadvg
Z96|) U2dIH 2 Sunulog [GQsioyuoaRInH) 4osdnssajul snjspuiad =|
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($S6|) anyuy 2 jaddoy proyiseied aepruoumnouyo] viajdouswAY ‘ds pjduig
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($S6|) anyuy 2 jaddoa proyseied depruoumMouys] pla}douswiAY ‘ds syoajdoqy
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(pS61) anyuy 2 jeddop proytseied depl[euUlolag eia}douswAY ‘ds snjAo0igoy]
[Qsioyusavin) sijjooyns saypoiydq—=|
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(PS6|) anyuy a2 jeddoy proqiseaed oepidouq viojdousw AY (uewyeq) ausoony puosopido)
($S6|) anyuy 2 jaddoy ploysesed oepluowimnouyos] viajdousw AY “ds xajdoduin)
[(uaIsWOH) syigninu xajdodup)=|
(pS6|) anyuy 2 jeddoy proqyisered aepruoumnouyds] piajdousw Ay (uljowin) sruoffip xajdodun)
(Z96|) Ua2ID 2 Suynulog [(S41ey) wnuoiiny snyousp.ay =|
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(OeploloL
sesajdopida) saypnuuayypiyos
(Z96|) Uad1D 2 Buynulog proyisesed ovaoe][loeg JOUIPIIG SIsualsUuLiny) SN[LIVG 3 studg PUDI/ONG DIUOLoDAYY
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(0661) Sanber 2 Ad,a7-uNg uosoyjed svaovioyyydowowuq nulog puviuoyouvyid vsoysydowojugq (J) ped wnydisojpdoyy
auUatIjYy =: AYDIU Bulps9oy ApH By 4JIPAO AUWIIUD [BANJEN 1SOH
“-panuuos VY XIGNAddV
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(SL6]) UasWURH 2 ploy aseydoyAyd aeprjowiosAry) elajdoajoa (Aqity]) Sisuappuvs DpqvylAL
(apa9D4A]SP)
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(661) ‘Te 19 URSLUIOD ployisered oepyeureisoyoiy, eiajdouswiAH Aap hy winsoiasd DUMDABOYIIA
(aepiyoayay :e1aj}dopida7)
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(aepruolnoing :e1a}do0a]0_)
(0861) SLO uasoyjed avooryjioeg ‘ds snyiovg — Aysjnyosyoy] srpuvaz snpiydojig
(aepruolnoing :e1a}doajo_)
(O861) SILO uasoyjed avaor[loeg ‘ds snpplovg (J) sniapuvas snjiydojig
(796) ueOT ployrseied oepruoorig eiajdouswAY (SOON) snyva]of snjojsoBAg
[uoseyyy apuojis
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[Aepljey supionsoxa [Aeg suouissios puojig=]
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(9007) 49qnH ployisered oepliewAy piajdouswiAY Qneig) vuvip saydoup snoelyey syjoolspurjAo DUuOjIS
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(7961) UIOM oyisered aeployouajaydy Ppoyewioan, wyny munypjo snyouajaydvjisping (wieysieyA]) SNIDLYSIjNU snjAjoos
(6861) Sueagq 29 WoNsAN ployisered seprydojng eiojdouswiAY (peowysy) wnjnauijijniu DUOSOWWDASDZ
(686|) SuBAq 27 WoNSAN proyiseied oeprydojng eiajdouswiAY (Joye) O1UIW O11DBIUG
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adUIIIJIY«=—- BYU Sulpsaq ATLUe JIP1O AWI9Ud [B.1NJeN] }soH]
““penunuos V XIGNuddV
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(eeprolyoy,
:erajdopidaq) (sa];NWUayyIyIS
(Z661) Jaqieg 2 Ag[sey proyisered oepiuoorig plojdouswiAy (S99N) 4OJDIpIuIp snsspg 2p sluaq) VUD]]290 Pjouopids
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(aepruornoin, :e1ajdoajop)
(9007) 49qnH ployiseied oepliewAy eiojdouswAY (ueyed) apipuajpo saydoupy (Aes) snypuaa snaoydouaydy
(aepruolnoin, :e1ajdoajo))
(900Z) J2qnH ployiseied oepuewAy| eiajdouswiAy (ueyed) avipuajpo saydoup jeyuat[AD snjnaiwd snsoydouayds
(aepruolnoiny :e1ajdoajoD)
[uapuayiyD saplojopAzau
snsoydouayds=] (snioiiqe 4)
(9007) 20qnH plowseied aepliewAy viajdouswAY (ueyen) apipuajpo saydoup saplojopazau snsoydouayds
(aepiuornoin, :e1ajdoajo5)
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(9007) 49qnH ployseied aepuewAy] viajdouswiAY (ueyen) apipuajoo saydpup uapuaynlyD sipiow snsoydouayds
(sepruolnoing
elajdoajog) uapuantyo
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(aepluornoing :esajdoajoa)
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(sepruorlpnoiny
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= [neyinbo5
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| (oepruegey :eiajdiq) xajdwios
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5 (ZS6|) UosuIgoy Jowepoid oepuosoyUy viojdiway (Aes) snsorpisul sniic,
a (ZS6|) UOsUIgoY Joyepoid oepidosAiy) viaidoinan, ‘dds pdosday)
= (Qepr[[aul990_9 :e19}d09]0))
aoe (ZTS6|) UosuIgoy Jowepoid aeploooyjUy elo}diwiay (Aes) snjnosnut siio20yjup (a93U0997) wnjound snsoyjalg
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Ontario records of Sperchopsis tessellata JESO Volume 144, 2013
ONTARIO RECORDS OF SPERCHOPSIS TESSELLATA
(ZIEGLER), ARARELY COLLECTED LOTIC WATER
SCAVENGER BEETLE (COLEOPTERA, HYDROPHILIDAE)
S.A. MARSHALL
School of Environmental Sciences, University of Guelph
Guelph, ON NIG 2W1
email: samarsha@uoguelph.ca
Scientific Note J. ent. Soc. Ont. 144: 113-114
Sperchopsis LeConte is a distinctive monotypic eastern North American genus
of water scavenger beetles including only S. tessellata (Zeigler), a rarely collected species
restricted to the margins of cold, clear, rapidly flowing streams where it prefers undercut
sandy or gravelly banks. Spangler (1961) reviewed the distribution and biology of S.
tessellata, recording it from the Canadian provinces of Nova Scotia and Quebec as well
as from localities throughout eastern United States. Smetana (1988), in his important
review of the Hydrophilidae of Canada, provides only three Canadian collection records
for S. tessellata: one from Nova Scotia, one from Quebec and one from New Brunswick.
Roughley (1991) used Smetana’s review as the basis for a checklist of Hydrophilidae of
Canada, but gave the known Canadian distribution of S. tessellata as Nova Scotia, Quebec
and Ontario instead of Nova Scotia, New Brunswick, and Quebec. There are no Ontario
specimens of S. fessellata in the Wallis-Roughley Museum of Entomology (University
of Manitoba) so Roughley’s listing of Sperchopsis from Ontario was probably a /apsus.
The relatively large (about 7 mm long), strikingly convex, pitted adults of this species are
easily distinguished from other water scavenger beetles, so it is unlikely that it would be
overlooked in collections or samples from aquatic insect surveys. The new Ontario records
of Sperchopsis given below are based on the only known Ontario collections of the genus.
I first collected and identified this species from Ontario in 1976, from the Credit
River near Belfountain in Wellington County. Despite subsequent searching in apparently
suitable parts of other streams and rivers in southern Ontario (including the Eramosa, Saugeen,
Speed, Grand, Sauble, Rankin and Crane Rivers), no further Sperchopsis specimens were
found until 2007 and 2008, when one beetle was found on rotting wood embedded in an
undercut sandy bank in the Credit River near Erin, and another was collected in the Credit
River very close to where the first Ontario specimen had been collected 31 years earlier.
Data for the Ontario specimens of Sperchopsis, all deposited in the University of
Guelph Insect Collection, are as follows (latitude and longitude are not on the original label):
Ontario, Wellington County, Belfountain, Credit River, 43°48'6.56"N 79°59'47.10"W, April
3, 1976, S. A. Marshall; Belfountain at the fork of the Credit River, May 5, 2007, Adam
Brunke; Ontario, Wellington County, Credit River at the crossing of highway 124 near Erin,
43°50'6.01"N 80° 1'20.04”,W, May 1, 2008, S. A. Marshall.
Published December 2013
103
Marshall JESO Volume 144, 2013
FIGURE 1. Adult Sperchopsis tessellata from the Credit River, Ontario. Body length
approximately 7 mm.
In view of the general rarity, taxonomic distinctness, and restricted habitat of this
beetle, it is of potential importance as a species of conservation concern in Ontario.
Acknowledgements
Thanks to Miles Zhang for confirming that there are no Canadian Sperchopsis specimens in
the J. B. Wallis museum.
References
Roughley, R. 1991. Hydrophilidae. Pp. 130-135 in Bousquet (ed), Checklist of beetles of
Canada and Alaska. Agriculture Canada publication 1861/E.
Smetana, A. 1988. Review of the family Hydrophilidae of Canada and Alaska (Coleoptera).
Memoirs of the Entomological Society of Canada 142: 1-316.
Spangler, P. 1961. Notes on the biology and distribution of Sperchopsis tessellatus (Ziegler)
(Coleoptera: Hydrophilidae). The Coleopterists Bulletin 15: 105-12.
114
I. clavata, new generic placement for a misclassified species JESO Volume 144, 2013
IDIOTYPA CLAVATA (PROVANCHER, 1888) (HYMENOPTERA:
DIAPRIIDAE), NEW GENERIC PLACEMENT FORA
MISCLASSIFIED SPECIES
J. T. HUBER
Natural Resources Canada, c/o Canadian National Collection of Insects, AAFC,
960 Carling Avenue, Ottawa, ON, Canada, K1A 0C6
email: john.huber@agr.gc.ca
Scientific Note J ent. Soc. Ont. 144: 115-117
In 1888, Provancher (1889) described Camptoptera (as “Camptotera”’) clavata
from a male and a female collected at Ste. Gertrude, Quebec, though he did not note
the actual number of specimens examined. Presumably it was only two. He also did not
designate a primary type. Girault (1911) borrowed what was thought to be the unique
specimen of C. clavata, labelled “Camptotera clavata Prov. 1598”, but it arrived badly
damaged so he could only state that it definitely did not belong to Mymaridae. Girault
remounted the fragments remaining—initially stated to be “a single fore wing and several
tarsi’; later in the same description corrected to “these notes are based on a fore wing and
tibiae and tarsi of two legs’—in Canada balsam on a slide and described the fore wing
venation and leg remnants before returning the specimen to the sender, Abbé V.A. Huard,
Musée de |’ Instruction Publique, Quebec [City]. Girault’s designation must be construed as
a lectotype designation according to ICZN Article 73.1.3 and Recommendation 73F:
“Where no holotype or syntype was fixed for a nominal species-group taxon established
before 2000, and when it is possible that the nominal species-group taxon was based on
more than one specimen, an author should proceed as though syntypes may exist and,
where appropriate, should designate a lectotype rather than assume a holotype.” Gahan and
Rohwer (1917) correctly treated Girault’s “type” as a lectotype.
At my request, J. Perron, retired curator of the Provancher Collection, searched for
the Girault slide and noted that it had been lost. So nothing at all remains of the lectotype.
However, he found another specimen labeled in Provancher’s hand as C. clavata (Fig. 1)
and sent it to me for study. It is unquestionably one of the syntypes because its label number
#616 (Fig. 1) corresponds to catalogue number 1598 of Provancher’s personal collection,
which is the number of the lost type seen by Girault (J.-M. Perron, personal communication).
It is a species of Trichopria (Diapriidae) (Figs. 1-5), similar to 7. virginica (Ashmead) (L.
Masner, personal communication).
Peck (1963) catalogued the literature on C. c/avata. Both he and Burks (1979) had
treated the species as unplaced within Chalcidoidea, even though Girault (1911) had stated
“The fore wing... has the venation of a Pteromalid”. The question is whether Girault’s brief
description of those remnants actually fits that of a North American species of Pteromalidae.
If the Code is scrupulously followed, only Girault’s redescription of the lectotype can
be used to determine the correct identity of Camptoptera clavata. Provancher’s original
description must be disregarded, because it did not explicitly include a type designation,
Published December 2013
LES
Huber JESO Volume 144, 2013
ip : Ths amit Rat g : mi ai
FIGURES 1-5. Camptoptera clavata, paralectotype female. 1, habitus, lateral; insert:
labels. 2, head, anteroventral. 3, antenna, dorsal. 4, head and thorax, dorsal. Scale for Fig.
1=1 mm.
and the second Provancher syntypical specimen of C. c/avata is not a name-bearing type.
Yet, based on Provancher’s original description and the only remaining specimen of the
syntype series (the paralectotype) C. clavata could also be a species of Diapriidae.
Girault described the fore wing venation as “the costal cell is well developed,
the submarginal vein long and slender, eight or more times longer than the short, straight,
broad marginal vein, which is twice the length of the stigmal vein, which is distinct but
116
I. clavata, new generic placement for a misclassified species JESO Volume 144, 2013
without a neck; postmarginal vein somewhat shorter than the stigmal and short and broad,
subconic. Apex of the submarginal vein just before it joins the submarginal is colorless.”
No Nearctic member of Pteromalidae remotely fits Girault’s description of the venation.
The only Nearctic pteromalid that has a relatively short, straight and broad marginal vein is
Pachyneuron mucronatum Girault but in this genus the submarginal vein is much less than
eight times the length of the marginal which, in turn, is about as long as the stigmal vein
and the postmarginal vein is longer than the stigmal vein. As for the tibia and 5-segmented
tarsi, no species of Pteromalidae exactly fits Girault’s description: “The tarsi are 5-jointed,
with the spur forked and the strigil well-developed on the cephalic legs. The proximal tarsal
joint is long. The tibiae are curved and enlarged distad, almost club-shaped. The proximal
tarsal joint of the cephalic legs is curved at the base.” In contrast, Girault’s description fits
almost perfectly species of /diotypa (Diapriidae) (Masner, personal communication and
my own examination of Nearctic specimens). Thus, both the lectotype and paralectotype
are shown to belong to the same family, i.e., Diapriidae, though unfortunately not to the
same genus. Camptoptera clavata is therefore removed from being an unplaced genus
within Chalcidoidea, as catalogued by Peck (1963) and Burks (1979), and is here placed in
Idoiotypa (Diapariidae) as /. clavata (Provancher), comb. n.
Acknowledgements
I thank J. M. Perron for finding and sending me the only remaining specimen of
Camptoptera clavata, informing me about the correspondence of catalogue numbers for
this species. My retired colleague, L. Masner (Canadian National Collection of Insects,
Ottawa), kindly checked the generic placement of the paralectotype and suggested its likely
relationships. J. Read is thanked for taking the photographs and preparing the plate.
References
Burks, B. D. 1979. Family Mymaridae. Pp. 1022—1033 in Krombein, K. V., Hurd, P. D. Jr.,
Smith, D. R., and Burks, B.D. (eds), Catalog of Hymenoptera in America North of
Mexico 1. Washington, DC: Smithsonian Institution Press. 1198 pp.
Gahan, A. B. and Rohwer, S. A. 1917. Lectotypes of the species of Hymenoptera (except
Apoidea) described by Abbé Provancher. The Canadian Entomologist 49: 298—
308, 331-336, 391400.
Girault, A. A. 1911. Notes on the Hymenoptera Chalcidoidea, with descriptions of several
new genera and species. Journal of the New York Entomological Society 19: 175—
189.
Peck, O. 1963. A catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). The
Canadian Entomologist, Supplement 30. 1092 pp.
Provancher, L. 1989. Additions et Corrections au Volume II de la Faune Entomologique
du Canada Traitant des Hyménoptéres. Québec: C. Darveau. 475 pp + Errata
(1 p).
7
JESO Volume 144, 2013
118
New Ontario records for NV. m. marmoratus JESO Volume 144, 2013
NEW ONTARIO RECORDS FOR NANOPHYES M. MARMORATUS
(GOEZE, 1777) (COLEOPTERA: BRENTIDAE), INTRODUCED
INTO NORTH AMERICA FOR CLASSICAL BIOLOGICAL
CONTROL OF PURPLE LOOSESTRIFE
E. ST. LOUIS
Faculty of Science, University of Ottawa,
Ottawa, ON, Canada KIN 6N5
email: estlol03@uottawa.ca
Scientific Note J. ent. Soc. Ont. 144: 119-123
In 1992, Nanophyes marmoratus marmoratus (Goeze) (Coleoptera: Brentidae),
the Purple Loosestrife Flower Weevil, native to Eurasia (Thompson et al. 1987), was
identified as a candidate for biological control of Purple Loosestrife, Lythrum salicaria (L.)
(Lythraceae), an alien weed invasive in North America. Although N. m. marmoratus was
assessed as having a lower potential impact than the other biological control candidates,
Neogalerucella calmariensis L., N. pusilla (Duftschmid) (Chrysomelidae) and Hylobius
transversovittatus (Goeze) (Curculionidae), it was still released due to its high likelihood
of establishment (Blossey and Schroeder 1995). From 1994-2005, N. m. marmoratus was
released in several US states (Skinner 1996; Blossey 2001). In Canada, 720 adults were
released in 1997 in southern Manitoba in three marshy sites (Lindgren et al. 2002). In
Ontario there is no record of this species ever having been released (D. Mackenzie, personal
communication). The present study is the first to monitor the spread of N. m. marmoratus
in Canada since its release in North America, thus addressing Corrigan et al.’s (2013)
recommendation to assess its establishment in Canada.
Nanophyes m. marmoratus adults are small (1.4-2.1 mm long), with light
yellowish-brown elytral markings and a long rostrum. Females (Fig. 1a) are slightly larger
and with more yellow on the elytra than males (Fig. 1b). The life cycle (egg to adult
emergence) is about four weeks. The weevils first appear in mid-spring, when they mate
on the flowering inflorescences before laying eggs singly in flower buds (Batra et al. 1986;
Blossey and Schroeder 1995). The young whitish larvae feed on the stamens and ovary of
unopened flower (Wilson et. al. 2004). The adult beetles emerge, feed on foliage and mate
in late August before overwintering in leaf litter (Lindgren et al. 2002).
In 2012 and 2013 several Purple Loosestrife populations in eastern Ontario were
surveyed. Only the well-established, leaf-feeding Neogalerucella beetles were expected
to be found but Nanophyes m. marmoratus was repeatedly discovered as well, on Purple
Loosestrife flowers from mid-June to late August. One of these specimens, from Pakenham,
ON, 45.3333°N 76.2833°W, 10 September 2012 (1 female, CNC), was reported by Douglas
et al. (2013) along with specimens collected in Quebec in 2011 from the area between the
Ottawa and St. Lawrence Rivers (Fig. 2). Nanophyes m. marmoratus has now been identified at
eighteen sites in eastern Ontario (Fig. 2) and quantified in fourteen of them. The average density
was 0.78 (+ 0.55) weevils per stem (n=513 stems), with a range of 0-14 weevils per stem.
Published December 2013
119
St. Louis JESO Volume 144, 2013
FIGURE la. (left) Female and 1b. (right) male NV. m. marmoratus showing elytral patterns.
Attacked buds do not flower; they become blackened, filled with frass and usually
fall off the plant (Wilson et al. 2004). In populations where N. m. marmoratus is common,
Purple Loosestrife fruit densities are noticeably lower, with higher proportions of aborted
seedpods (Fig. 3). Flower bud abortion was often seen on the lower half of the inflorescence
with many to all of the buds along this section missing. Affected buds that remain on plants
each have a single small exit hole—additional subtle evidence of the weevils’ presence.
The damaged bud can look very similar to fully developed fruits (seed capsules). Weevil
damage is easily spotted and distinguishable from that caused by Neogalerucella beetles.
The weevils’ presence reduces overall seed production. At high weevil densities, larval
feeding can reduce fruit output of a Purple Loosestrife plant by up to 70% (Van Dreische
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FIGURE 2. Records for Nanophyes m. marmoratus in eastern Ontario and southwestern
Quebec. Light grey points are new reports, dark grey points are those reported in Douglas
(2013).
120
New Ontario records for N. m. marmoratus JESO Volume 144, 2013
FIGURE 3. Abscission of flower buds caused by MN. m. marmoratus.
et.al. 2002; personal observation). In the fourteen sites with N. m. marmoratus, reductions in
reproductive output were measured as the portion (%) of the total lengths of inflorescences
bearing aborted flower buds. These reductions averaged 12.6% (+ 10.6%) with a range of
0—34.5% over all sites. When considering only the damaged plants (n=85) within the sites,
total reproductive output reductions averaged 53% (+ 26.2%) with a range of 0-100%.
Early in the biocontrol programme, it was thought likely that combining
biological control agents would significantly decrease Purple Loosestrife density (Malecki
et al. 1993; Blossey and Schroeder 1995; Wilson et al. 2004; Skinner 2006) but Coombs
(2004) suggested that V. m. marmoratus would thrive when Neogalerucella was low or
absent. At one Ontario site, anecdotal evidence suggested that obvious niche partitioning
occurred; N. m. marmoratus was found primarily on host plants situated in a field, whereas
Neogalerucella beetles were more concentrated on plants in the adjacent, recently-mowed,
roadside ditch. The mowed plants had an increased production of tender, young shoots
preferred by Neoalerucella, whereas unmowed plants in the field retained their flowers,
preferred by Nanophyes m. marmoratus.
Nanophyes m. marmoratus is now present in southwestern Quebec and Ontario
(Douglas et al. 2013). In Ontario, over 403 individuals were counted at 18 sites (50 vouchers,
University of Ottawa). Populations of N. m. marmoratus are present through extensive
areas of eastern Ontario, having been found along the Ottawa River watershed from above
Petawawa to the National Capital Region and throughout the Rideau waterway from Kingston
to Ottawa. Over the last 19 years tens of thousands of these weevils have been released and
redistributed in several northeastern states (Blossey, personal communication). The origins
of the Ontario populations are likely the closest US release sites, i.e., Buffalo (42.8553°N
121
St. Louis JESO Volume 144, 2013
78.8552°W) and Laurel Marsh, NY (42.8709°N 77.2424°W), about 300 km from the most
southerly Ontario site at Queen’s Biological Field Station (44.5681°N 76.3201°W). Though
N. m. marmoratus dispersal has not been described, Ferrarese and Garono (2010) noted that
dispersals across large expanses of open water have occurred in Oregon. They suggested
that adults are capable of dispersing 100-300 km/year. Blossey (personal communication),
also indicated that NV. m. marmoratus adults are strong dispersers and have generally spread
in a northeastern pattern, helped by the prevailing winds. It would be useful to survey
additional areas for the presence of NV. m. marmoratus, especially those between the original
release sites in the USA and Manitoba and the sites mapped here.
Acknowledgements
I would like to acknowledge P. Bouchard who confirmed the identity of collected specimens
and J. Corrigan, D. McKenzie, B. Hall and M. Stastney for assistance, guidance and support.
Funding was provided by an Ontario Ministry of Research and Innovation Early Researcher
Award to Risa D. Sargent.
References
Batra, S. W., Schroeder, T. D., Boldt, P. E. and Mendl, W. 1986. Insects Associated with
Purple loosestrife (Lythrum salicaria) in Europe. Proceedings of the Entomological
Society of Washington 88: 748-759.
Blossey, B. 2001. Impact and management of purple loosestrife (Lythrum salicaria) in
North America. Biodiversity and Conservation 10: 1787-1807.
Blossey, B. and Schroeder, D. 1995. Host specificity of three potential biological weed-
control agents attacking flowers and seeds of Lythrum salicaria (Purple Loosestrife).
Biological Control 5: 47-53.
Coombs, E. M., Clark, J. K., Piper, G. L. and Cofrancesco A. F. Jr. 2004. Biological control
of invasive plants in the United States. Western Society of Weed Science, Oregon
State University Press, Corvallis, Oregon. 457 pp.
Corrigan, J., Gillespie, D. R., De Clerck-Floate, R. and Mason, P. G. 2013. Lythrum salicar-
ia L., Purple Loosestrife (Lythraceae) Pp. 363-366 in Mason, P. G. and Gillespie,
D. R. (eds) Biological Control Programmes in Canada 2001-2012. CABI Publish-
ing, Wallingford, Oxfordshire, UK. 518 pp.
Douglas, H., Bouchard, P., Anderson, R. S., de Tonnancour, P., Vigneault, R. and Webster,
R. P. 2013. New Curculionoidea (Coleoptera) records for Canada. Zookeys 309:
13-48.
Ferrarese, E. and Garono, R. J. 2010. Dispersal of Galerucella pusilla and Galerucella
calmeriensis via passive water transport in the Columbia River Estuary. Biological
Control 52: 115-122.
Lindgren, C. J., Corrigan, J. and De Clerk-Floate, R. A. 2002. Lythrum salicaria L., Purple
Loosestrife (Lythraceae). Pp. 383-390 in Mason, P. and Huber, J. T. (eds), Biological
122
New Ontario records for N. m. marmoratus JESO Volume !44, 2013
Control Programmes in Canada, 1981-2000. CABI Publishing, Wallingford, UK.
583 pp.
Malecki R. A., Blossey, B., Hight, S. D., Schroeder, D., Kok L. T. and Coulson, J. R., 1993.
Biological Control of Purple loosestrife. Bioscience 43: 680-686.
Skinner, L. 1996. Integrated control of Purple loosestrife in Minnesota. Minnesota
Department of Natural Resources. Fifteenth North American Prairie Conference. Pp.
219-222.
Thompson, D. Q., Stuckey, R. L. and Thompson, E. B. 1987. Spread, impact and control
of purple loosestrife (Lythrum salicaria) in North American Wetlands. United States
Fish and Wildlife Service, Fish Wildlife Research 2: \—55.
Wilson, L. M., Schwarzlaender, M., Blossey, B. and Randall, C.B. 2004. Biology and
biological control of purple loosestrife. USDA Forest Service, FHTET-04-12. 78
Pp.
Van Driesche, R., Blossey, B., Hoddle, M., Lyon, S. and Reardon, R. 2002. Biological
control of invasive plants in the Eastern United States. USDA Forest
Service FHTET-2002-04. 413 pp.
123
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First records of Z. indianus from Ontario and Quebec JESO Volume 144, 2013
FIRST RECORDS OF ZAPRIONUS INDIANUS GUPTA (DIPTERA:
DROSOPHILIDAE) FROM COMMERCIAL FRUIT FIELDS IN
ONTARIO AND QUEBEC, CANADA
J. M. RENKEMA", M. MILLER!, H. FRASER?, J-PH. LEGARE’, AND
RB HALLETT
'School of Environmental Sciences, University of Guelph,
50 Stone Road West, Guelph, ON, Canada NIG 2W1
email: renkemaj@uoguelph.ca
Scientific Note J. ent. Soc. Ont. 144: 125—130
Zaprionus indianus Gupta (Diptera: Drosophilidae) was described in India (Gupta
1970) (Fig. 1) but is suspected to be native to the Afrotropical Region (Chassagnard and
Kraaijveld 1996). In the New World, it was first found in late 1998 in Sao Paulo, Brazil
and has since spread rapidly throughout South and Central America (Vilela 1999; Goni
et al. 2001; Tidon et al. 2003). Zaprionus indianus was first detected in North America
in July 2005 in Florida (Steck 2005) and is now reported from many eastern, central and
southwestern states (van der Linde et al. 2006; van der Linde 2013). This species is now
globally widespread and considered cosmopolitan, present in temperate and tropical regions
(Tidon et al. 2003; Commar et al. 2012).
Zaprionus indianus is a generalist, with the ripe fruits of at least 74 plant species
in 31 families in Africa recorded as breeding sites (Lachaise and Tsacas 1983). It has a
similarly wide host range in South and North America and has become a significant pest
of figs (Ficus carica L.) in Brazil (Santos et al. 2003; Stein et al. 2003; van der Linde et al.
2006). While Z. indianus is often associated with damaged or fallen rotting fruit, larvae
are able to invade the soft tissue of figs before harvest and have been reared in Florida from
tree-ripened Malphigia emarginata (Barbados cherry), Punica granatum (pomegranate),
Eriobotrya japonica (loquat) and Dimocarpus longan (longan) (van der Linde et al. 2006;
Pasini et al. 2011). In northeastern USA, Z. indianus has been reported in high numbers
in net collected samples in a vineyard and was found in apple cider vinegar traps in cherry,
raspberry and blackberry fields (Biddinger et al. 2012).
Here we report the first records of Z. indianus in Canada, with all specimens found
in southern Ontario and Quebec. Specimens of Z. indianus were found during surveys for
Drosophila suzukii (Matsumura) in apple cider vinegar traps in pre- and post-harvest fields of
Published December 2013
“Author to whom all correspondence should be addressed.
Ontario Ministry of Agriculture and Food and Ministry of Rural Affairs, Box 8000, 4890
Victoria Avenue North, Vineland ON, Canada LOR 2E0
*Ministéres de l’Agriculture, des Pécheries et de |’Alimentation du Québec, Complexe
scientifique, 2700 rue Einstein, local C. RC 105, Québec, QC, Canada GIP 3W8
125
Renkema et al. JESO Volume 144, 2013
Se, We oe
FIGURE 1. Zaprionus indianus fro
A. Marshall).
m Africa dorsolateral habitus (Photograph by Stephen
peach, blueberry, raspberry, strawberry, cherry and plums. Many of the Ontario specimens
were collected by the Ontario Ministry of Agriculture and Food and Ministry of Rural A ffairs
(OMAF/MRA); all were identified by M. Miller and S. A. Marshall and deposited in the
University of Guelph Insect Collection, Guelph ON (DEBU). The specimens collected in
Quebec were identified at the Laboratoire de diagnostique en phytoprotection of Ministere
de l’Agriculture, des Pécheries et de |’ Alimentation du Québec (MAPAQ), confirmed by M.
Miller and S. A. Marshall, and deposited in the Collection d’insectes du Québec, Québec
QC (CIQ). Voucher specimens from both provinces are deposited in the Canadian National
Collection, Ottawa, ON.
Material examined. ONTARIO. Essex. Near Harrow, blueberries, 10.1x.2013, OMAF/
MRA (1912, DEBU). Near Harrow, blueberries, 3.ix.2013, OMAF/MRA (192, DEBU).
Near Harrow, peaches, 10.ix.2013, OMAF/MRA (219264, DEBU). Near Ruthven, peaches,
12.ix.2013, OMAF/MRA (1934, DEBU). Chatham-Kent. Near Blenheim, raspberries,
10.ix.2013, J. Renkema (19, DEBU). Near Blenheim, raspberries, 10.ix.2013, OMAF/
MRA, (1214, DEBU). Near Blenheim, 2.ix.2013, J. Renkema (12, DEBU). Niagara.
Near Beamsville, plums, 12.ix.2013, OMAF/MRA, (1914, DEBU). Near Niagara-on-the-
Lake, cherries, 11.ix.2013, OMAF/MRA, (14, DEBU). Simcoe. Near Barrie, peaches,
10.viii.2013, H. Fraser (19, DEBU). QUEBEC. Chateau-Richer. La Cote-de-Beaupre,
strawberries, 2.x.2013, MAPAQ (1%, CIQ). Compton. Coaticook, raspberries, 7.x.2013,
MAPAQ (14, CIQ). Laval. Laval, strawberries, 7.x.2013, MAPAQ (19, CIQ). Laval,
strawberries, 15.x.2013, MAPAQ (19, CIQ). Pierreville. Nicolet-Yamaska, strawberries,
8.vii.2013, MAPAQ (19, CIQ). Nicolet-Yamaska, raspberries, 19.vi1i.2013, MAPAQ
(12, CIQ). Sainte-Pétronille. L’ile-d’ Orléans, raspberries, 3.x.2013, MAPAQ (19,
126
First records of Z. indianus from Ontario and Quebec JESO Volume 144, 2013
FIGURE 2. Zaprionus indianus from Ontario, Canada, male, lateral habitus.
CIQ). Sainte-Sabine. Brome-Missisquoi, strawberries, 6.ix.2013, MAPAQ (19, CIQ).
Brome-Missisquoi, strawberries, 10.x.2013, MAPAQ (CIQ). Salaberry-de-Valleyfield.
Beauharnois-Salaberry, raspberries, 17.ix.2013, MAPAQ (12, CIQ). Yamaska. Pierre-De
Saurel, blueberries, 21.viii.2013, MAPAQ (1, CIQ).
Zaprionus indianus 1s the only member of Zaprionus Coquillett present in Canada
to date. It is distinguished from all other Canadian Drosophilidae by its reddish-brown
head and thorax with unique silvery stripes that extend dorsally from the antennae to the
tip of the scutellum (Fig. 3) and laterally from the leading edge of the thorax to the base of
each wing (Fig. 2) (Gupta 1970; Steck 2005; van der Linde et al. 2006; Yassin and David
2010). Because future invasion by other Zaprionus species is possible (van der Linde
2010), including the invasive Z. tuberculatus Malloch, currently established in Egypt and
Israel, and the potentially invasive Z. ghesquieri Collart, introduced to Hawaii and Cyprus,
but without established populations (Patlar et al. 2012; Yassin A, 2013, pers. comm.) we
provide additional features that would confirm that specimens are Z. indianus.
The keys to African (Yassin and David 2010) and European (Bachli et al. 2004)
Zaprionus species, the description in van der Linde (2010), and the original species
description by Gupta (1970) were used to identify our specimens of Z. indianus. Zaprionus
indianus specimens have 4—6 composite spines with second short branches arising directly
from the fore femur (a character of all 15 members of the vittiger species group) (Fig. 4);
the silver stripes with black borders are narrow and the black borders do not widen at the
scutellum; the scutellum lacks a white tip; the abdomen is light yellow; and the subapical
setae on the fourth and fifth abdominal tergite arise from dark spots. In males, the aedeagal
flap is smooth apically and serrated basally (distinguishing it from Z. africanus Yassin and
David with a deeply serrated apical margin and Z. gabonicus Yassin and David with a
127
Renkema et al. JESO Volume 144, 2013
FIGURE 3. Zaprionus indianus, head and FIGURE 4. Zaprionus indianus, fore
thorax, dorsal view. femur, lateral view, showing the composite
spines.
complete lack of serration apically and basally). In females, the oviscape has six peg-like
ovisensilla (Z. africanus with 7 or 8 ovisensilla), and the spermatheca length to width ratio
is 0.95—1.16 (Gupta 1970; Steck 2005; van der Linde et al. 2006; Yassin and David 2010).
Zaprionus indianus is unlikely to become an established pest of fruit in Ontario and
Quebec. The small numbers of flies we report from Ontario and Quebec suggest Z. indianus
may have moved in from the United States in late summer and autumn 2013. However,
it can adapt to a wide range of climates (Karan et al. 2000), and if it can successfully
overwinter it may also spread rapidly in Canada, as evidenced by its rapid expansion in
the USA since its first discovery there (van der Linde 2013). Large populations are often
observed the year after its initial detection, particularly in urban environments (Ferreira and
Tidon 2005). Unlike D. suzukii, Z. indianus is not known to infest ripe, undamaged fruit, but
if it can use ripening fruit already attacked by D. suzukii, there is the potential for increased
damage to harvested fruit. Therefore, future monitoring for D. suzukii should include Z.
indianus. Further study on the biology and ecology of this fly is warranted, if population
levels in Canada are found to increase.
Acknowledgements
We thank participating farmers for allowing access to fields for D. suzukii surveys,
Anne Horst for coordinating the surveys in Ontario, collaborators at Ontario Ministry of
Agriculture and Food and the Ministry of Rural Affairs and the Ministere de I’ Agriculture,
des Pécheries et de |’ Alimentation du Québec and agronomic adviser groups of Quebec for
collecting samples. Stephen Marshall (University of Guelph) and Amir Yassin (Université
Paris Diderot) provided helpful feedback on an earlier draft of the manuscript and the former
kindly allowed us to use a photograph of Z. indianus.
128
First records of Z. indianus from Ontario and Quebec JESO Volume 144, 2013
References
Biachli, G., Vilela, C. R., Escher, S. A. and Saura, A. 2004. The Drosophilidae (Diptera) of
Fennoscandia and Denmark. Fauna Entomologica Scandinavinca 38. 362 pp.
Biddinger, D., Joshi, N. and Demchak, K. 2012. African fig fly: another invasive drosophilid
fly discovered in PA. Plant and Pest Advisory, Rutgers New Jersey Agricultural
Experiment Station, Fruit Edition 17: \—2.
Chassagnard, M. T. and Kraaijveld, A. R. 1996. The occurrence of Zaprionus sensu
stricto in the Palearctic region (Diptera: Drosophilidae). Annales de la Société
entomologique de France (N.S.) 27: 495-496.
Commar, L. S. Galego, L. G. D., Ceron, C. R. and Carareto, C. M. A. 2012. Taxonomic and
evolutionary analysis of Zaprionus indianus and its colonization of Palearctic and
Neotropical regions. Genetics and Molecular Biology 35: 395-406.
Ferreira, L. B. and Tidon, R. 2005. Colonizing potential of Drosophilidae (Insecta, Diptera) in
environments with different grades of urbanization. Biodiversity and Conservation
14: 1809-1821.
Goni, B., Fresia, P., Calvino, M., Ferreiro, M. J., Valente, V. L. and da Silva L. B. 2001. First
record of Zaprionus indianus Gupta 1970 (Diptera: Drosophilidae) in southern
localities of Uruguay. Drosophila Information Service 84: 61-65.
Gupta, J. P. 1970. Description of a new species of Phorticella Zaprionus (Drosophilidae)
from India. Proceedings of the National Academy of Sciences, India 36: 62.
Karan, D., Dubey, S., Moreteau, B., Parkash, R. and David, J. R. 2000. Geographical clines
for quantitative traits in natural populations of a tropical drosophilid: Zaprionus
indianus. Genetica 108: 91—100.
Lachaise, D. and Tsacas, L. 1983. Breeding sites in tropical African drosophilids. Pp. 221—
332 in Ashburner, M., Carson, H.L. and J.N. Thompson, J.N. (eds), The Genetics
and Biology of Drosophila. London, Academic Press. 382 pp.
Pasini, M. P. B., Link, D. and Schaich, G. 2011. Attractive solutions efficiency in capturing
Zaprionus indianus Gupta, 1970 (Diptera: Drosophilidae) in Ficus carica L.
(Moraceae) orchard in Santa Maria, Rio Grande do Sul, Brazil. Entomotropica
26: 107-116.
Patlar, B., Koc, B., Yilmaz, M. and Ozsov, E. D. 2012. First records of Zaprionus tuberculatus
(Diptera: Drosophilidae) from the Mediterranean Region, Turkey. Drosophila
Information Service 95: 94-96.
Santos, J. F., Rieger, T. T., Campos, S. R. C., Nascimento, A. C. C., Félix, P. T., Silva, S. V.
O., and Freitas, F. M. R. 2003. Colonization of Northeast Region of Brazil by the
drosophild flies Drosophila malerkotliana and Zaprionus indianus, a new potential
insect pest of Brazilian fruticulture. Drosophila Information Service 86: 92-95.
Steck, G. J. 2005. Zaprionus indianus Gupta (Diptera: Drosophilidae), a genus and species
new to Florida and North America. Pest Alert, Florida Department of Agriculture
and Consumer Services, Division of Plant Industry. DACS-P-01677.
Stein, C. P., Teixeira, E. P., and Novo, J. P. S. 2003. Aspectos biologicos da mosca do
figo, Zaprionus indianus Gupta 1970 (Diptera: Drosophilidae). Entomotropica 18:
219-221.
129
Renkema et al. JESO Volume 144, 2013
Tidon, R., Leite, D. F. and Leao, B. F. D. 2003. Impact of the colonization of Zapronus
indianus (Diptera, Drosophilidae) in different ecosystems of the Neotropical
region: 2 years after the invasion. Biological Conservation 112: 299-305.
van der Linde, K. 2010. Zaprionus indianus: species identification and taxonomic position.
Drosophila Information Service 93: 95—98.
van der Linde, K. 2013. Zaprionus indianus distribution in the United States. http://www.
kimvdlinde.com/professional/Zaprionus%20distribution%20US.php (accessed 10
October 2013).
van der Linde, K., Steck, G. J., Hibbard, K., Birdsley, J. S., Alonso, L. M. and Houle, D.
2006. First records of Zaprionus indianus (Diptera; Drosophilidae), a pest species
on commercial fruits from Panama and the United States of America. Florida
Entomologist 89: 402-404.
Vilela, C. R. 1999. Is Zaprionus indianus Gupta, 1970 (Diptera, Drosophilidae) currently
colonizing the Neotropical region? Drosophila Information Service 82: 37-39.
Yassin, A. and David J. R. 2010. Revision of the Afrotropical species of Zaprionus (Diptera:
Drosophilidae), with descriptions of two new species and notes on internal
reproductive structures and immature stages. Zookeys 51: 33-72.
130
First record of L. /i/ii in a wild population of S. amplexifolius —§ JESO Volume 144, 2013
FIRST RECORD OF LILIOCERIS LILH (COLEOPTERA:
CHRYSOMELIDAE) EGGS IN A WILD POPULATION OF
STREPTOPUS AMPLEXIFOLIUS (LILIACEAE)
C. D. KEALEY*, N. CAPPUCCINO’, P. G. MASON?
Institute of Environmental Science, Carleton University
Ottawa, ON K1S 5B6
email: ckealeyO8@gmail.com
Scientific Note J. ent. Soc. Ont. 144: 131-134
Lilioceris lilii (Scopoli) (Coleoptera: Chrysomelidae), the Lily Leaf Beetle, is an
invasive European species first found at Montreal, Canada, in the 1940s (Gold et al. 2001).
It is a serious pest of cultivated Lilium spp. and Fritillaria spp. (Liliaceae) and has spread
across southern Canada and northeastern United States (LeSage 1983; Gold et al. 2001).
The beetle also poses a threat to native lilies in Ontario and Quebec, including Canada
Lily, Lilium canadense L., and Wood Lily, Lilium philadelphicum L. (Ernst et al. 2007;
Bouchard et al. 2008). In fact, in Ontario and Quebec eight out of 20 wild populations of L.
canadense were infested with L. /i/ii (Bouchard et al. 2008). There are also records of L. /ilii
adults feeding on plants in other liliaceous genera, e.g., Polygonatum (Temperé 1926; Fox
Wilson 1942), Streptopus (Ernst 2007), as well as genera in other families, e.g., Solanum
(Solanaceae) (Temperé 1926).
Kealey (2013) investigated Claspleaf Twistedstalk, Streptopus amplexifolius
(L.) DC. (Liliaceae), as a potential novel host of L. /ilii. Streptopus amplexifolius occurs
in rich moist coniferous and deciduous woods in all provinces and territories in Canada
and all adjacent states of the USA (Anonymous 2013). This native plant flowers from late
spring until mid-summer. Streptopus amplexifolius leaves were offered to L. /ilii larvae to
determine survivorship and development time. Leaves of S. amplexifolius were collected
from a wild population growing in Gatineau Park, Quebec, Canada (45.491°N 75.863°W).
Infestations of L. /ilii were recently reported in urban areas south of Gatineau Park, but no
known L. /ilii populations are established within the Park nor on any wild populations of S.
amplexifolius (Cappuccino 2013).
During a routine collection of S. amplexifolius plants for laboratory tests on June
25, 2013, a row of three L. /ilii eggs (Fig. 1) was discovered on the underside of a wild S.
amplexifolius leaf. Surrounding plants were searched for more eggs, though none were
discovered, nor was any obvious feeding damage by adults or larvae observed. The S.
amplexifolius leaf with the L. /ilii egg mass was carefully removed from the stem and
transported to the laboratory. The eggs were left undisturbed, and the leaf was placed on
Published December 2013
* Author to whom all correspondence should be addressed.
'Department of Biology, Carleton University, Ottawa, ON
*Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Ottawa,
ON
131
Kealey et al. JESO Volume 144, 2013
moist filter paper in a 60 x 15 mm Petri dish maintained at 23°C, 70% relative humidity and
16:8 L:D, and monitored for larval hatch. The three eggs hatched on June 30" or July 1%.
Two first instar larvae were still alive on July 2" and feeding damage was observed on the
leaf whereas the third larva was dead and had not fed. Subsequently both surviving larvae
died. The cause of death is unclear.
The eggs (Fig. 1), and hatched larvae (Fig. 2), found in Gatineau Park shared all of
the characteristics of L. /ilii. Lilioceris lilii egg masses are distinct: they are laid parallel to
leaf veins, in a linear arrangement of 2-16 eggs on the underside of host leaves (Salisbury
2008); eggs are bright red or orange in color, though darken when near hatching, and are
covered in a sticky orange layer; individual eggs are 1.0 x 0.5 mm, and masses are laid
from March—September (Haye and Kenis 2004); and Lilioceris lilli larvae are dirty-orange
in color, with a dark head and legs. First instar larvae (Fig. 2) have head capsule widths
between 0.36—0.55 mm, and a distinct egg bursting spine is located on the first abdominal
segment (Livingston 1996; Cox 1994). Larvae also carry a viscous fecal shield of their own
excrement on their backs.
Eggs of other genera within the Criocerinae subfamily may be confused with L.
lilii eggs. Hosts of Lema spp. belong to the distantly related plant families Solanaceae
and Asteraceae, and Oulema spp. are on species of Asteraceae, Commelinaceae and
Poaceae. The only species of Neolema that occurs in Canada, N. cordata White, occurs on
Commelinaceae spp. Two species of Crioceris closely resemble L. /i/ii in the larval stage;
however, both are closely associated with Asparagales and have distinctly different egg
placement and color (White 1993).
This observation marks the first record of L. /ilii ovipositing on S. amplexifolius
in nature and this is the first plant species outside the genera Lilium, Fritillaria, and
Cardocrinium (the known host genera for this beetle) where both oviposition in nature
and successful larval development in the lab have been observed (see Salisbury 2008).
Although Ernst et al. (2007) found that larval performance was poor on S. amplexifolius
leaves in laboratory tests, Kealey (2013) confirmed that almost half (42%) of L. /ilii larvae
can successfully develop to adults on S. amplexifolius. This record is also only the second
oviposition record for L. /i/ii in North America on a host plant in nature outside of urban
areas where development might also be occasionally possible. The observation reported here
is likely the result of an adult that emigrated from an urban area. However, it is unknown
what the potential is for colonization by L. /ilii of novel host plants, such as S. amplexifolius,
in non-urban areas. Among the factors that might encourage a more permanent move to S.
amplexifolius by L. lilii is the enemy-free-space hypothesis in which the herbivore escapes
its specialized parasitoid by feeding on a novel host plant (Brown et al. 1995; Rossbach et
al. 2006). Further study would help to establish if such events are rare or the first step in
adaptation by an invasive alien species to a novel host.
132
First record of L. /i/ii in a wild population of S. amplexifolius | JESO Volume 144, 2013
'
eal hidctcal ae 2 "-44 ee
FIGURE 1. Lilioceris lilii eggs on Streptopus amplexifolius \eaf from Gatineau Park,
Quebec.
FIGURE 2. Lilioceris lilii first instar larva on a Streptopus amplexifolius leaf collected from
Gatineau Park, Quebec. This picture was taken soon after larval death and shows A) feeding
damage B) egg bursters and C) fecal shield characteristic of the species.
133
Kealey et al. JESO Volume 144, 2013
References
Anonymous. 2013. 2. Streptopus amplexifolius (Linnaeus) de Candolle, Fl. Frang. ed. 3.
3: 174. 1805. Flora of North America 26: 145—147. Available from http://www.
efloras.org/florataxon.aspx?flora_id=1&taxon id=242101972 [Accessed 22
November 2013].
Bouchard, A., McNeil, J. N. and Brodeur, J. 2008. Invasion of American native lily
population by an alien beetle. Biological Invasions 10: 1365—1372.
Brown, J. M., Abrahamson, W. G., Packer, R. A. and Way, P. A. 1995. The role of natural-
enemy escape in a gallmaker host-plant shift. Oecologia 104: 52-60.
Cappuccino, N. 2013. Lily leaf beetle tracker. Available at: http://lilybeetletracker.weebly.
com/ [Accessed 8 November 2013].
Cox, M. L. 1994. Egg bursters in the Chrysomelidae, with a review of their occurrence
in the Chrysomeloidea (Coleoptera). Pp. 75—100 in Jolivet, P. H., Cox, M. L.
and Petitpierre, E. (eds), Novel aspects of the biology of Chrysomelidae. Kluwer
Academic Publishers, Dordrecht, The Netherlands. 582 pp.
Ernst, C., Cappuccino, N. and Arnason, J. T. 2007. Potential novel hosts for the lily leaf
beetle Lilioceris lilii Scopoli (Coleoptera: Chrysomelidae) in eastern North
America. Ecological Entomology 32: 45—52.
Fox Wilson, G. 1942. The lily beetle, Crioceris lilii Scopoli: its distribution in Britain
(Coleoptera). Proceedings of the Royal Entomological Society of London (A) 18:
85-86.
Gold, M. S., Casagrande, R. A., Tewksbury, L. A. and Livingston, S. B. 2001. European
parasitoids of Lilioceris lilii (Coleoptera: Chrysomelidae). Canadian Entomologist
133: 671-674.
Haye, T. and Kenis, M. 2004. Biology of Lilioceris spp. (Coleoptera: Chrysomelidae) and
their parasitoids in Europe. Biological Control 29: 399-408.
Kealey, C. D. 2013. Potential host plants of the invasive beetle Lilioceris lilii (Coleoptera:
Chrysomelidae): implications for the biological control agent Tetrastichus setifer
Thomson Hymenoptera: Eulophidae). B.Sc. Honours thesis. Carleton University,
Ottawa, ON. 19 pp.
LeSage, L. 1983. Note sur la distribution présente et future du criocere du lys, Lilioceris
lilii Scopoli (Coleoptera: Chrysomelidae), dans l’est du Canada. Le Naturaliste
Canadien 110: 95-97.
Livingston, S. S. 1996. Biological control and host range of Lilioceris lilii: Anew ornamental
pest in the USA. MLS. Thesis, University of Rhode Island, Kingston, USA. 78 pp.
Rossbach, A., Lohr, B. and Vidal, S. 2006. Parasitism of Plutella xylostella L. feeding on a
new host plant. Environmental Entomology 35: 1350-1357.
Salisbury, A. 2008. The biology of the lily beetle, Lilioceris lilii (Scopoli) (Coleoptera:
Chrysomelidae). Ph. D. Thesis. Imperial College, London, UK. 157 pp.
Temperé, G. 1927. Régime alimentaire anormal de Crioceris lilii(Coleoptera Chrysomelidae).
Proceés Verbaux de la Société Linnéenne de Bordeaux (1926) 78: 131-133.
White, R. E. 1993. A revision of the subfamily Criocerinae (Chrysomelidae) of North
America north of Mexico. United States Department of Agriculture Technical
Bulletin No 1805. v + 158 pp.
134
JESO Volume 144, 2013
THE ENTOMOLOGICAL SOCIETY OF ONTARIO
OFFICERS AND GOVERNORS
2013-2014
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Received on: 01-27-14
Journal of the AMNH LIBRARY
scety of Ont UIA
, 100213326
CONTENTS
I. FROM THE EDITOR oc.ccccccscsccsecsessescocscccesccccocsnscessossacecssessesnccasstaneccartassessvusteussusvel seater ae tenennn 1
Il. ARTICLES
J.L.RINGROSE, K. F. ABRAHAM andD. V. BERESFORD —Newrange records of mosquitoes
(Diptera: Culicidae) from northern Ontarid........0:<cccccssossrsscscsssccsevscsessceseuccessceevescssaueetattanene 3-14
J. T. HUBER — Taxonomy and faunistics in Ontario, 1952-2012: Publications in the “Journal
of the Entomological Society of Ontario”. <......0...c:<seccoossesceassecssessesesevsareuessessassvenressitenteaeenein 15-26
P. MASON — Biological control in Ontario 1952-2012: A summary of publications in the
“Journal of the Entomological Society of Omtario”..............scsssccscsseescssscsssssssssesssessesseecees 27-111
Ill. NOTES
S. A. MARSHALL — Ontario records of Sperchopsis tessellata (Ziegler), a rarely collected
lotic water scavenger beetle (Coleoptera: Hydrophalidae)...............ssssssssssssssesserssesseses 113-114
J.T. HUBER — Idiotypa clavata (Provancher, 1888) (Hymenoptera: Diapriidae), new generic
placement for a misclassified SPeCieS.............sesscessesscescesscsscessesseessesscsscessssssessecsenssessessenses 115-117
E. ST. LOUIS — New Ontario records for Nanophyes m. marmoratus (Goeze, 1777)
(Coleoptera: Brentidae), introduced into North America for classical biocontrol of purple
LOOSCStHIfEC......essssoresssecesossncsesesanceosoocsetesscescsssventenstetdenssaasenssotsesssiasstecestsusencetntacrer tat teennetTeaaat 119-123
J. M. RENKEMA, M. MILLER, H. FRASER, J-PH. LEGARE, and R. H. HALLETT — First
records of Zaprionus indianus Gupta (Diptera: Drosophilidae) from commercial fruit fields in
Ontario and Quebec, Camadar....csccercocsssscsosescsssesocsssosssoccssavecsesesevencsessecsssersvevcesectttscearten 125-130
C.D. KEALEY, N. CAPPUCCINO and P. MASON —First record of Lilioceris lilii (Coleoptera:
Chrysomelidae) eggs in a wild population of Streptopus amplexifolius (Liliaceae)....... 131-134
IV. ESO OFFICERS AND GOVERNORS 2013-2014..........cssscsssscssssssssssssessssessesesessessssssenserenenss 135
V. ESO OFFICERS AND GOVERNORS 2012-2013............sssssesssessesseessessonee inside front cover
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