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 Pp10991 UOYIOU MoU 9 ounf | Toye) SUaTDAUT DyAasTNyD c Ringrose et al. p1o0de1 ulayyiou Mou = ¢ |: Any 0} Z| ouNe S1-¢ Arne br 6¢ (AayeM) SuBgunjsad vIpyjayjinboyD p1osa1 WOYyIIOU Mou ¢] Ajng ‘g ung ¢ Ayn G € sesleA lajiva sajaydoup p1o0da1 WdyyIoU Mou QZ ounr I YIOIIYIOA, jadwas Sapay [[yur des L Ajng—g ounr = ¢ Aynf—9 oune GG ik (Aqiry) 4ojound sapap QOUTAOIA 10} PIODAI SIV, Q] oun I (nayinbod) smpjjnd sapay p10901 U1d]sed puke UIOYIOU Mou g oun I (1dyJeA\) SUDIOAOAd SAapay [[yur des pl Ajng—g oung =—s_/, Ayn¢—Z oune TE 1G JeXq sdiuoid sapapy pi0se1 WIdYyINOS MOU L Ajng I (Qpajsione7) sadisiu sapay p10901 UJOYIOU MoU 97-8 ounr 61 IeAq suapnajul sapay p1o0da1 WdYyIOU MoU Ol Ajng ‘g ounr LI-z oune G G YIOIDYIOA, sNIDIIdu1 Sapapy [[yur des wiajsamyyou Z ounr 8] (19y[eM) 4ad1dui sapay [[yur des €[-g oung Z|] Ayn¢—Z oune II vd IeAq snjuopoxay sapap p1o0de1 WoYyyIoU Mou SZ ‘€Z oune ZI Arne Z Z (Jaye) SUDIINAIXA SAapay pioda1 UayyIoU Mou Q|[ sung b (UdSIBJ\\) SI]DSAOp Sapapy [[yur ded usajsamyyou QZ ‘S[ ounr IL ‘O] ounce G Il (499) aq]) SlUNWWUOD Sapay p10dei WoYyOU Mou €Z ounr I UII SNaaUld sapay pi0de1 UJOYyIOU Mou QZ ‘SZ oune G (p[eqooy |) sisuappubs sapay [[yur des L Ajnp—g oungc 1 Ajng—z] sung 67 | (SunoX pure ya4) smpssasqv sopay ratiys I107 TlOT =—s EL OZ OLILJUG 10J 9duvYd UOHNQLYSIG poainjdes (s)ajVq Avak Jad ye) saiadg ‘Aueq|y OY JO UP} OS] UIYIM Z[OT UI pur ‘oye Apueg pure oye] NOLL Sig Jo WY OS] UTYIM | [OZ UI payoaT[oo saroads aeprolnD “| ATAVL 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). 18 16 14 12 10 8 z ——+—+SH1 100 200 300 km expected number of species Oo NY fF OD 0 50 100 150 200 catch size FIGURE 1. Rarefaction analysis of mosquito catches (means and SDs) within 150 km of 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 July 10 —July 16 | 82° 3'24" — 51° 21'22" lke {sasha ae July 10—June 16 [83° 22'44"_52° 18'23" July 3 = July 9 a OZone July 3 — July 9 S2ogtoneals pens" a0! june 26 = Ily Oe wa 82240 een soy S| June 26 —July 2 [83° 17'24"_51° 29" 53" | [xx KO Junes19= June 255/820 4119s spe Sas! June 19 — June 25__|81° 39' 23" 51° 58' 8" Tuned 2 = unew SaaS ONS 7 oles ORSu June 12—June 18 | 80° 23' 11" 51° 26' 40" June 5—June 11 __ {82° 39' 13" _51° 55" 53" June §—June 11 _[81°57'48"_52° 461 35" & a July 6—July 13 193° 32' 10" _53° 36'9" | ft aaa uly6— July 13) | 91e Aton 5220 7m37% |. ae June 28 —July 5 __| 94° 13'38" 52° 49' 28" ns June 28—July5 193° 2'33"__ 53°27" 40" June 16 — June 23 [88° 33' 33" 54° 28' 19" | = June 16—June 23 [oo°2r'3er savor | [| — 15 92° 1'44" 54° 9' 30" al 15 ggosa'si" 53245135" |_| 4 le! ial — S le le 2°o 5» |5 QO |o WW |00 |00 Sampling dates 0001 0 — — — June 7 89° 40' 43" 54° 25' 50" May 31 — June 7 S926 25 SSE ZNSE 3) (3) Ex se |8 = (5) a) > = 5 5) VS BS bt —) Aedes abserratus 002 Aedes canadensis 0004 Aedes communis 0006 Aedes excrcians 0005 Aedes dorsalis 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 suaypdul DjJasijny supganjsad pipijayjinboy lajAba Sajaydoup yaduas Sapay 4ojound sapay snyoyjnd sapay supJ0ao0id sapay sdiuoid sapay sad1isiu sapay SUIPNAJUI Sapapy snjooydui sapay Jasidul sapay SNJUOPOXIY Sapapy 2 5) 5 June 25 sayep suljdures 10 — June 16 Jul June 26 — July 2 June 26 — Jul June 19 June 19 — June 25 June 12 —June 18 June 12 — June 18 June 5 — June 11 June 5 — June 11 July 6 — Jul July 6 — Jul June 28 — Jul June 28 — Jul o|June 16 — June 23 June 16 — June 23 May 31 — June 7 May 31 — June 7 N b ae | >) N _— ~ psnuynos.¢ ATaVL Ringrose et al. JESO Volume 144, 2013 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 number of species/class —-NT OO — veil line|S 2 ages 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. 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YSIoIpusg L86l LII N 9L Syeyqey sno[oorpew wWoynos = =NOTPYsIeYY 2 Wepsuls L861 LIT a 9£ oeployd eigjdiq 1sunj SutAvoap ul Ys] UopuoT ‘ydjanyD NO [[eysIeI WUMOIG SR6I SII a “J wnuojynf It snovsdig — avaovavsdiq SUIISIA S]OASUI “OD pueUIpreH NO pprr p86l FII = T< eINolg aoutAold 10} Mou wiayjnos = NO ulWOL ZR6l ZII cS) tC oepiquie’) viajdoa]o) Spfey W109 LAO Ne) STA PAIAL O86l OIT s urpuioy Oo OF weoy ainjsed ‘pray us0o 4st] uopuo7 ‘ydjany = NO 3 wsqpeolg OR6I OII aS If oepisAder ein|diq soutAold JOJ MOU = WIDJSaMyINOS «=NOQ AseN AWUNWOL 6/61 6OI n ge L souorido SI] “OD puewipreH NO pprf 8261 801 = L epodos] 4ST] (OD/puewaipren NO ppnf LL61 LOI S o 19 eraydopiday ist] Aponng IN SLOW 961 901 ao) =| 9 ero} do[AIpD, SUOTIIPpe ISI] NO ey x AIDADIA CL6I SOI = saisadg ‘dds # snua5 Aque JIPIO sp.10m Aay AWBIOT vary (s)doyjny «vad "[OA S) 140d 5 E ““panuiquos 7 qTaVL P3} JESO Volume 144, 2013 Huber cri Aes vypjoundiq 8LT LSL Sol (uaseH) ajiaro (‘q) ppsadiuid II € cIv 8L saisadg ‘dds # oepiqeie) vaplody sajpdo.iay oepridwiog ‘|s ‘aeproayds aeploAquieia) pyegynoy “eyAyduiAs DUBD]JOUY Iepluoliseuso0y) SNIMMOT aeUuA]OIS snuas5 Arey eiajideryiyd “Weoy piaydoajopd oepisjnuide) Jo sauisesed sIBATe via}doyuG suONNqiysIp/sp10de1 MoU eiojdouawAY eiaj}douswiAy elajdiwioH eiajdouswiAy viaydoajoa ero diwiay viaj}doajoa arourly eiajdorsmaydq elojdouswiAY R1eUOPO e}eUuOpO, eiajdoajog elaydos] Rjoquie|[0D eiajdooay Fey ere) epeues Joy Mou ul uonedinxa AuNOS JO} MoU 1S] JSA10} [RII0q sIBAle ul eyoudyiousyony Sp1O901 Mou 1SI| ISI] yeuurars Yeo 1ST] uorsuedxa pre Maypiou s1ojepaid/sproyisesed Sp10901 MOU Isl] uoHNgINsIp sp.0m Ady UIO}]SOM ye aisadsen UOY NOS ‘o- uo}dwie 7] ua}seo “‘woyynos Pole BIRSPIN A}IRIOT aw NO “urd “urd NO ‘urd NO ere) NO NO 30 ued NO NO NO NO NO WV “ur NO Baty AeMoyey ‘ye 19 preyonog ye 19 [jeysieyy yong 2 olaied a0sponH Je 19 O1aIed yong auiednq 2 uinbeg ‘Je Jo preyonog ayeponbioysjy asesayq] 2 uinbeg ydjopuey 7 AUayROoW ye Jo JOBNS jaeyorues 2 UOISULADYS Ssurped 1Yysug SaJA SIOWPHIS Joyony 2 Jodiey (s).1oyjny L00T 9007 9007 POT pO0T FOOT POT 1007 LO00C L00C 000T 8661 8661 Lo6l 9661 9661 S661 S661 b661 ABad 149d Lel 9ET cel vel vel rel vel cel cel cel Ie] 6cl 6cl 8cl Lél EG 9TI 9TI STl “TOA “panunuos 7 q1aVL 24 SO Volume 144, 2013 4 JI Taxonomy and faunistics in Ontario, 1952—2012 oye] nsjeunseg smpiydiiny SNUSOIOAAG, oepidiuAd vlajdouswAY SOUIAOId JOJ MOU dy) UO BILBLIN NO pesy w49qnH ZIOT EFI SHO 8 (dquqny) snaudus sduydoyjo) aepluavoAT viajdopida7] uoNNqiysip woynos NO UMOIg-sUIWIOUDq ZIOZ EFI pS ETE TTANY ZUMPLIOWY Snpuansuljsip snquog oepidy viajdouswAYH POLOWy YLION 10} Mou Imy MV ww PleyJous 1107 Trl (yoy) sniuvdii SNISIUO]AF] — VeplostuoyotL epodos] SOUIAOIG OJ MOU "OD MAIJUDY ~=—-—- NO Uaplo 3% ould[yoW [10Z Trl ([e1S) sdyoy pydiowojpAyT —- epi wui0jyeUdg viojdiwiay Axjunoo Joy Mau ue) = RAD w ulesoy [LOT Trl Ad Ad br }SI] ‘S]OOSUI Jeog Jejog NO Plojsaiog [107 Trl 91 seplayeyq vlajdoajoa sp10901 MoU ‘WY 'N sejsnoq [107 Trl Janajaday winiojaaisa aplyovsayy oepidy eio}douawA HY dOUIAOId JOJ MoU souLeyYeD IS NO ye 19 playseus OLOT I+rl 97 aepruolpnoing viajdoajod S]soJOJ poompiey =-|eyUD ‘yINOS ~=NO ye 19 19}901g OLOT Irl 8I snquog oepidy piaydousw Ay ASojousyd NO ysourng 97 B09 O07 Itl avourly ‘e1ajdoa}o9 68 esojdoyuO sieaye NO ‘yeqo supe O07 IPI 8 DUIJDAID oepidy via}douswAY saruous yeinjeu woyinos NO yes yonIyIIA OLOT Ir Ags[oonH © oeprynoundig viaidiq Sp10991 soy Mou ZV W UOUIADIS 600T OFI (MOLY) DauDIsDd DAapv]vDy —-aeplavqereog viajdoajop douTAoId JO} Mou SN —_ SIBOY 2 IBIND 6007 OFI 8 SvouRTy’ SUOTIPPE 90 uinbed 3007 6€1 ‘ye 10 Il deploAquiviad vlajdoajoa SSO] ‘auT]Dap sar1sads NO ajeponbioj9W LOOT BEI SI snquog oepidy piajdouswiA} spjoy ejours aw ‘yey yooun, LOOT LEI saisadg “dds 4 snuo5 AyWune J 1IpsO sp.10Mm oy AWYBIOT vay (s)doyjny «Ava "[OA ‘14%d je Ponuynves'¢ Aalelv.L 25 Huber JESO Volume 144, 2013 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), 31 Mason JESO Volume 144, 2013 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 32 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 33 Mason JESO Volume 144, 2013 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 34 Biological control in Ontario, 1952—2012 JESO Volume 144, 2013 (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. 35 Mason JESO Volume 144, 2013 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 36 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). 337) Mason JESO Volume 144, 2013 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 38 Biological control in Ontario, 1952-2012 JESO Volume 144, 2013 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: 39 Mason JESO Volume 144, 2013 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 40 Biological control in Ontario, 1952-2012 JESO Volume 144, 2013 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 41 Mason JESO Volume 144, 2013 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 42 Biological control in Ontario, 1952—2012 JESO Volume 144, 2013 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 43 Mason JESO Volume 144, 2013 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 44 Biological control in Ontario, 1952-2012 JESO Volume 144, 2013 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 45 Mason JESO Volume 144, 2013 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 46 Biological control in Ontario, 1952—2012 JESO Volume 144, 2013 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). 47 Mason JESO Volume 144, 2013 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 48 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. 49 Mason JESO Volume 144, 2013 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 50 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 Silk 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. 52 Biological control in Ontario, 1952—2012 JESO Volume 144, 2013 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. 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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. 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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. 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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). 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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. 62 Biological control in Ontario, 1952-2012 JESO Volume 144, 2013 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. 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