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Chapter 9 of Ojibway Prairie, a Southern Ontario Tallgrass Prairie

Steve M. Paiero and Stephen A. Marshall Department of Environmental Biology, University of Guelph Guelph, Ontario, Canada Paul D. Pratt Windsor Department of Parks Windsor, Ontario, Canada Matthias Buck Department of Environmental Biology, University of Guelph Guelph, Ontario, Canada

Abstract. This chapter describes the fauna of Ojibway Prairie, a tallgrass prairie complex in southern Ontario, highlighting the tallgrass-dependent and tallgrass-associated species among the over 2,000 insect species found there so far. The presence of tallgrass-dependent and tallgrass-associated species reflects Ojibway Prairie’s status as a fragment of a formerly more continuous grassland and thus supports the prairie peninsula hypothesis. The chapter includes a discussion of insect species associated with other southern Ontario tallgrass prairie sites and compares these species with those found in Ojibway Prairie. Also discussed are rare species found at Ojibway Prairie but not associated specifically with tallgrass habitats. Forty-four insect species new to Canada or new to Ontario (1 Orthoptera, 3 , 10 Coleoptera, 16 Diptera, and 14 ) are recorded from Ojibway Prairie.

Résumé. Ce chapitre décrit l’entomofaune de la prairie Ojibway, un complexe de prairies à herbes hautes du sud de l’Ontario, en portant une attention particulière aux espèces dépendantes des herbes hautes ou associées à ces dernières et qui sont au nombre des quelque 2 000 espèces d’insectes recensées jusqu’ici à cet endroit. La présence d’insectes dépendants des herbes hautes ou associés à ces dernières est un reflet de l’état actuel de la prairie Ojibway, qui n’est plus qu’un fragment d’une prairie autrefois plus continue, et vient appuyer l’hypothèse de la « péninsule de prairie ». Le chapitre traite également des espèces d’insectes associées à d’autres prairies à herbes hautes du sud de l’Ontario et compare ces espèces à celles qui se trouvent dans la prairie Ojibway. Il examine enfin les espèces rares qui vivent dans cette prairie mais qui ne sont pas associées directement aux habitats d’herbes hautes. On a établi la présence dans la prairie Ojibway de 44 espèces nouvelles pour le Canada ou l’Ontario (1 orthoptère, 3 hémiptères, 10 coléoptères, 16 diptères et 14 hyménoptères).

Introduction North American tallgrass prairies form a subclimax habitat dominated by grasses and forbs, which can grow to more than 2 m in height and are maintained by dry microclimates and seasonal fires limiting the growth of woody plants. This habitat occurs largely in transition zones between the Great Plains and the eastern forests, but many small patches of tallgrass prairie are scattered within a triangle bounded by Oklahoma, southern Ontario, and southeastern Manitoba. These eastern grasslands are considered to be remnants of a “prairie peninsula” that developed in the Hypsithermal period (approximately 6,000– 8,000 years ago), when temperatures were approximately 1–2 °C higher than current

Paiero, S. M., S. A. Marshall, P. D. Pratt, and M. Buck. 2010. Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie. In of Canadian Grasslands (Volume 1): Ecology and Interactions in Grassland Habitats. Edited by J. D. Shorthouse and K. D. Floate. Biological Survey of Canada. pp. 199-225. © 2010 Biological Survey of Canada. ISBN 978-0-9689321-4-8 doi:10.3752/9780968932148.ch9 200 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

averages (Transeau 1935; Wright 1968). During this period, eastern forests would have receded as a result of desiccation (from increased annual temperatures, western winds, and decreased soil moisture) and mortality from fire, allowing grasslands to expand into eastern North America (Wright 1968). Following the Hypsithermal period, when average annual temperatures returned to those of modern day, the prairie peninsula was fragmented as woody plants recolonized their previous range (Wright 1968), leaving prairie remnants only in those areas most favourable to grasslands (see Chapter 4). The remaining tallgrass prairie (like the faunistically similar oak savanna) was further fragmented by increased human activity following the immigration of Europeans approximately 150 years ago (Szeicz and MacDonald 1991). Prairie remnants now persist only where well-drained soils, microclimate, and periodic fires (Wright 1968) have maintained these otherwise “forest climax” areas. In Ontario, tallgrass prairie and oak savanna are now estimated to encompass less than 1% of their original range (Bakowsky and Riley 1994). The presence of disjunct populations of tallgrass-dependent organisms, particularly brachypterous or apterous species incapable of sustained flight, throughout these prairie sites is evidence that they were indeed formerly part of a continuous prairie peninsula that was once connected to the Great Plains (Ross 1970; Mlot 1990; Hamilton 1994). The prairie peninsula had allowed prairie organisms otherwise restricted to the Great Plains to migrate along a prairie corridor into northeastern North America. As the peninsula was later fragmented by the encroachment of the eastern forests, disjunct populations of these tallgrass species were left in the remnants, creating communities of diversity not found elsewhere in eastern North America. One such prairie remnant is found within the city limits of Windsor, Ontario. The Ojibway Prairie Remnants Area of Natural and Scientific Interest (ANSI) (42°46′N 83°05’W) is composed of five different areas (Ojibway Prairie Provincial Nature Reserve, Ojibway Park, Tallgrass Prairie Heritage Park, Spring Garden Natural Area, and the Black Oak Heritage Park, collectively referred to as the Ojibway Prairie Complex; Fig. 1). It is the second largest prairie site in Ontario (320 ha, second only to Walpole Island, which has approximately 650 ha of tallgrass prairie and oak savanna habitat). The Ojibway Prairie Complex supports a rich Carolinian flora as well as 116 prairie indicator plant species (Pratt 1989) and is one of the few sites in Ontario that uses fire management to sustain the tallgrass prairie along with its surrounding savanna and open oak woodland. From 1979 to 2007, 21 early spring burns were done at the Ojibway Prairie Complex in an effort to maintain this unique Ontario grassland.

Previous Taxonomic Surveys of Northeastern Tallgrass Prairies John Macoun, the Dominion Botanist of Canada, discovered the “eastern extension of the prairie flora” at Ojibway Prairie when he visited the site in August 1892. His observation was mostly ignored by biologists until the early 1970s, when efforts were made by numerous groups to protect this site. Faber-Langendoen and Maycock (1983) surveyed the floral composition and soil characteristics of several southern Ontario tallgrass prairie remnants and found six types of prairies. Ojibway Prairie was classified as a wet prairie, with soil characteristics similar to those of the western prairies and a flora dominated by forbs with strong prairie affinities. Crins (1997) found over 30 nationally rare vascular plants specific to tallgrass prairies in Ontario. The distribution of these tallgrass prairie indicator species in Ontario has been well documented by the Ontario Natural Heritage Information Centre since. By contrast, relatively little work has been done on communities of Ontario tallgrass. Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 201

Fig. 1. Aerial view of the Ojibway Prairie Remnants Area of Natural and Scientific Interest (Windsor, Ontario, Canada) with its five different areas (green).

In the most significant studies of Ontario tallgrass arthropods to date, Hamilton (1994, 1995) recorded numerous prairie-dependent (Hemiptera: Cicadellidae) and (Hemiptera: Fulgoroidea) species from tallgrass prairie sites throughout Ontario and the northeastern United States, supporting the theory that some of these sites are remnants of the prairie peninsula. Bouchard et al. (2001) surveyed insects in southern Ontario alvars, which share some floral components with prairies (Catling and Brownell 1995), and found 18 tallgrass-associated Auchenorrhyncha of potential use as indicators of alvar quality. Skevington et al. (2000) surveyed the insects of Lambton County, including several oak savanna and grassland sites, and recorded several tallgrass-associated species of Cicadellidae, Cercopidae, and Caliscelidae. Sugar et al. (1998) conducted a Malaise trap survey of aculeate Hymenoptera of an Ontario oak savanna and found one putative prairie species, Dufourea marginata (Cresson) (Halictidae). Several insect surveys have been conducted in tallgrass prairies in the United States. Panzer (1988) found numerous prairie-dependent Lepidoptera, Hemiptera, and Orthoptera species in tallgrass prairie remnants in Illinois and Indiana, and discussed how to best manage these remnants to conserve prairie species. Larsen and Work (2003) listed several grassland-associated species of Carabidae from restored and original tallgrass prairie in Iowa. Comer et al. (1995) collected 201 insect species from Lakeplain Prairie in Michigan, including numerous Auchenorrhyncha that they considered to be “potentially host specific” on prairie vegetation. Wilson et al. (1993) surveyed the Fulgoroidea of an 202 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

Illinois tallgrass prairie and documented their plant hosts, showing that several species were restricted to prairie vegetation. Panzer et al. (1995) sampled selected groups (mostly Lepidoptera, Orthoptera, and Hemiptera) in Illinois prairie remnants and found 1,100 species of insects, of which 256 were considered prairie “remnant-dependent.” This ranking was based on comparisons between natural areas and degraded, or disturbed, sites to determine if species were capable of surviving in disturbed habitats. Reed (1996) provided a literature review of prairie species with a list of 728 putative tallgrass associates, though a large number of the species listed are common in non-prairie sites and are not treated here as tallgrass associates.

Materials and Methods The insects of Ojibway Prairie have been sampled by University of Guelph personnel on an intermittent basis over the past 30 years, resulting in a steady accumulation of significant new records of rare and habitat-restricted arthropod species in the University of Guelph Insect Collection. Recognition of the importance of the many new records among the relatively small collections accumulated on this casual basis led to the initiation of a focused and intensive sampling program (as the senior author’s M.Sc. thesis) in the spring of 2001. Sampling continued, using pan traps and other techniques, through the summer of 2002, and a few additional records have been added since. The majority of collections took place in the Ojibway Prairie Provincial Nature Reserve, but occasional collections were made in other parts of the Ojibway Prairie Complex. Yellow pan traps were used because pan traps sample both flying and crawling insects, and yellow attracts a wide range of taxa (Kirk 1984; Marshall et al. 1994). Twenty- four pan traps were used continuously from May to October of 2001, with six pans in each of four sites in the nature reserve, including two sites that had been burned in May 2001. Each pan was flush with the soil surface and partially filled with salt-saturated water along with a small quantity of non-scented liquid dish soap to decrease the surface tension. Pan traps were emptied every three to four days and replaced when there was an observable decline in the colour intensity or if they were damaged. A single Malaise trap was also set on a private tract of land with grassland affinities near Ojibway Prairie (the probability of damage to a conspicuous trap in a public park precluded use of the trap in the prairie complex proper). Sweep netting and hand sampling was done opportunistically but yielded taxa that might otherwise have been missed by the traps. Sampling was continued from 2002 to 2007 by hand collecting, sweep netting, and short-term (48 h or less) placements of large numbers (50–200) of pan traps. Specimens were removed from the pan trap residues, rinsed in clean alcohol, and stored in 70% ethanol until they could be properly sorted and prepared. Generally, all specimens that were in identifiable condition were retained unless there were large series of single species. Malaise trap residues were subsampled for taxa of interest, with the remainder of the sample retained in 70% ethanol for future inspection. Hard-bodied insects (e.g., Coleoptera, large Hymenoptera) were air-dried and mounted, whereas small Diptera, Hemiptera, and other small, soft-bodied insects were dried by using a critical point dryer. Larger soft-bodied specimens were pinned directly and then dried by immersion in three 24-h baths of ethyl acetate. Those specimens identified to at least the generic level were databased by using Biota (Colwell 1997) and assigned individual specimen codes. Personnel at the University of Guelph Insect Collection identified most specimens, but specimens of some taxa were sent to specialists for identification or confirmation. All of the specimens collected for this Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 203

study were deposited at the University of Guelph Insect Collection, with the exception of material that specialists retained for further scientific study or voucher material. Ojibway Prairie specimens collected prior to 2001 and deposited in the University of Guelph Insect Collection were also identified and entered into the database. The database was further supplemented with lists generated by previous collections and counts taken at Ojibway Prairie. Organized butterfly counts sponsored by the North American Butterfly Association have been conducted annually at Ojibway since 1994, and 90 species of butterflies have so far been recorded. Other Lepidoptera records came from the moth collection of John E. Pilkington (collected at Ojibway between 1974 and 1985, now deposited in the Canadian National Collection in Ottawa). The Odonata of the Ojibway Prairie Complex have been surveyed by P.D. Pratt annually since 1985.

Results and Discussion

Specimens and Species Totals A total of 15,723 specimens were mounted and labelled, including specimens taken by hand and aerial sweep netting (4,175), Malaise traps (2,226), pan traps (9,322), and light traps (136). Over 12,500 of these specimens were identified to the genus or species level. This material was augmented with records of butterflies, moths, dragonflies, and damselflies obtained through amateur collecting done since 1974 and through several published records (Hamilton 1994, 1995). Over 2,200 taxa are now recorded from Ojibway Prairie (58 Odonata, 2 Mantodea, 1 Phasmatodea, 1 Dermaptera, 45 Orthoptera, 10 Psocoptera, 322 Hemiptera, 11 Neuroptera, 493 Coleoptera, 5 Mecoptera, 500 Diptera, 435 Hymenoptera, and 334 Lepidoptera). The Lepidoptera were not a focus in the M.Sc. study, and moths and butterflies recorded here are from butterfly counts and the Pilkington moth collection. Additionally, the Parasitica (Hymenoptera), most families of nematoceran Diptera, Anthomyiidae (Diptera), Muscidae (Diptera), and a few groups of Coleoptera collected and prepared during this project (approximately 1,000 to 1,200 specimens) remain largely unidentified beyond the family level. Further sampling and study of these groups will undoubtedly add several hundred species to the current list. Updated lists of Ojibway Prairie species are available online at the University of Guelph Insect Collection and Ojibway Prairie websites (http://www.uoguelph.ca/debu/Insects_Of_Ojibway_Species_List_2008. pdf and www.ojibway.ca/insect.htm).

What Are Tallgrass-Dependent and Tallgrass-Associated Insects? The term “tallgrass-dependent” is here used to refer only to species that require specific prairie plant hosts. Tallgrass-dependent host-specific phytophagous Hemiptera in the family Cicadellidae () were a focal group for this study. The phrase “tallgrass associate” is used here to refer to species more closely associated with tallgrass prairies than with other habitats but not necessarily dependent on prairie plant hosts. Tallgrass- associated species include species apparently restricted to tallgrass prairie in Ontario but found in a wider range of habitats in the eastern United States, as well as non-phytophagous species (predaceous, parasitic, or saprophagous species) known only from tallgrass sites but not specifically dependent on particular tallgrass plants. Panzer et al. (1997) provided a discussion of these and similar terms, along with discrepancies in the use of these terms by different authors. Hamilton (1994, 1995), Reed (1996), and Mawdsley (2003) provide 204 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

further remarks on tallgrass associates and tallgrass-dependent insect species. Species found in both natural and anthropogenic grasslands are difficult to place in either category because the disturbance of eastern forests for agriculture, forestry, and urban development has created meadows and fields (the cultural “steppe” of Lindroth 1957), some with grassland flora, that are readily colonized by vagile grassland species. Although some of these species might have originally been tallgrass-dependent or tallgrass-associated species, these now widespread synanthropic species are not treated here as tallgrass dependent or tallgrass associated.

Tallgrass-Dependent and Tallgrass-Associated Leafhoppers (Hemiptera: Cicadellidae) Several tallgrass-dependent and tallgrass-associated leafhopper species have been recorded from the northeastern prairie peninsula (Hamilton 1994, 1995, 2000, 2005; Comer et al. 1995; Panzer et al. 1995; Bouchard et al. 2001; Sauer 2002; Paiero et al. 2004) and have been critical in supporting the prairie peninsula hypothesis because of the host specificity and limited vagility of many species (Hamilton 1994). Some species of Cicadellidae are flightless, or at least have flightless females unable to traverse large distances between suitable host plants, and are thus unlikely to disperse to disjunct tallgrass sites in the absence of tallgrass corridors. Hamilton (1994, 1995) sampled numerous northeastern grasslands and found tallgrass-dependent leafhoppers as far east as southern Ontario. Ojibway Prairie was one of the sites studied by Hamilton (1994, 1995), and five of the nine “prairie species” he recorded from the Ojibway complex (Dorydiella kansana Beamer, Chlorotettix spatulatus Osborn and Ball, Laevicephalus minimus (Osborn and Ball), L. unicoloratus (Gillete and Baker), and Graminella oquaka DeLong) were also collected during our study. One of these species, L. unicoloratus, is here considered to be a tallgrass associate. The remaining four species, and another five species recorded from Ojibway for the first time in our study (Cuerna fenestella Hamilton, Destria fumida (Sanders and DeLong), Erythroneura carbonata McAtee, Polyamia caperata (Ball), and Neohecalus magnificusHamilton), are here considered to be tallgrass dependent. The host plants and distributions of these species are as follows: Dorydiella kansana is restricted to only a few tallgrass prairies and alvars in Ontario where it is recorded on Eleocharis elliptica Kunth and Scleria spp. (Bouchard et al. 2001); Chlorotettix spatulatus is recorded on Schizachyrium scoparium (Michx.) Nash (formerly Andropogon scoparius; Panzer et al. 1995; Hamilton 2005); Laevicephalus minimus feeds on Bouteloua curtipendula (Michx.) Torr. (Panzer et al. 1995; Hamilton 2005); Graminella oquaka is recorded on Panicum virgatum L. (Panzer et al. 1995; Hamilton 2005); Destria fumida (newly recorded here from Ontario) is recorded on Spartina gracilis Trin. (Hamilton 2005) and is recorded from Manitoba, Saskatchewan, and several midwestern states (Kramer 1976; Maw et al. 2000); Erythroneura carbonata occurs in prairies in Alberta, Saskatchewan, and Manitoba and in Ojibway Prairie and feeds on Lysimachia (Sauer 2002); and Polyamia caperata, a common species in Ojibway Prairie, feeds on Schizachyrium scoparium (Bouchard et al. 2001). Neohecalus magnificusis recorded on several Spartina species and occurs only in tallgrass prairies (Hamilton 2000). The host of Cuerna fenestella is unknown, but this species is otherwise known only from Manitoba tallgrass (Hamilton 1970). The remaining four prairie species collected by Hamilton (1994, 1995) from Ojibway Prairie (Flexamia prairiana DeLong, Graminella pallidula (Osborn), Limotettix elegans Hamilton, and L. urnura Hamilton) were not found during this study, but are nonetheless considered here to be tallgrass dependent. Flexamia prairiana is recorded on Bothriochloa spp., Andropogon gerardii Vitman, and Schizachyrium scoparium (Whitcomb and Hicks Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 205

1988), and Graminella pallidula occurs on “prairie grasses” (DeLong 1948). Both Limotettix elegans and L. urnura feed on Eleocharis elliptica (Bouchard et al. 2001). We consider the following species to be tallgrass associated: Balclutha abdominalis (Van Duzee), Chlorotettix fallax Sanders and DeLong, Gypona melanota Spangberg, Hecalus major (Osborn), Laevicephalus unicoloratus, Memnonia flavida (Signoret), Mesamia nigridorsum (Ball), Stroggylocephalus mixtus (Say), and Xerophloea major Baker. These species are often found in tallgrass prairies, but occur in a variety of other habitats throughout eastern North America and are recorded from the following tallgrass- associated flora:C. fallax on Panicum virgatum (Hamilton 2005); H. major on Calamagrostis canadensis (Michx.) P. Beauv., Hordeum jubatum L., and Poa compressa L. (Hamilton 2000); L. unicoloratus on Andropogon gerardii and Schizachyrium scoparium (Hamilton 2005); Memnonia flavida on Andropogon and Bouteloua curtipendula (Hamilton 2000); Mesamia nigridorsum on Helianthus species (DeLong 1948); and X. major on Monarda and other forbs (Bouchard et al. 2001). No hosts are known for Balclutha abdominalis or Stroggylocephalus mixtus. The only non-cicadellid considered here to be tallgrass dependent is Delphacodes parvula Ball, which has a range consistent with tallgrass prairie and hosts that are usually found in tallgrass habitat. Hamilton (2005) recorded D. parvula feeding on Andropogon gerardii and Schizachyrium scoparium. The differences between the study by Hamilton (1994, 1995) and our study for both tallgrass and non-tallgrass cicadellids suggest that the total leafhopper fauna of Ojibway Prairie is significantly larger than that recorded by either study. Hamilton (1995) suggested that the southern portion of the Ojibway Park (specifically the Ojibway Prairie Provincial Nature Reserve), which is maintained by spring burns, is depauperate (virtually no leafhopper fauna remaining) compared with the Tallgrass Prairie Heritage Park, which is not burned. Our collections, based almost exclusively in the provincial nature reserve, included many of the species collected by Hamilton (1994, 1995) at the Heritage Park along with others not found during his study. Our data thus do not support the suggestion that spring burns have a significant impact on the cicadellid fauna as suggested by Hamilton. If the sites were to be rated according to Hamilton (1995: Tables 2 and 3) by using the new species list, the provincial nature reserve would rate as an “above average” site (previously ranked “poor”).

Tallgrass Associates Many of the tallgrass-associated species discussed in the following subsections are known in Ontario only from tallgrass prairie sites, where they often occur in large numbers. Elsewhere in eastern North America, however, these species occur in oak savannas, meadows, and other grassland habitats.

Orthoptera At least three species of Orthoptera are closely associated with Ontario tallgrass prairies. Dichromorpha viridis (Scudder) occurs in rich pastures and meadows in the eastern United States (Vickery and Kevan 1985), but in Ontario it is restricted to a few sites with tallgrass affinities (Marshall et al. 2006). Melanoplus scudderi (Uhler) is recorded from only a few tallgrass prairies and oak savannas in Ontario (Marshall et al. 2006) but is found in open meadows and pastures in the eastern United States. Anaxipha exigua (Say) is commonly found in several mesic tallgrass prairies and is known in southern Ontario from only a few Carolinian sites (Marshall et al. 2006). Although Vickery and Kevan (1985) indicate that A. 206 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

exigua is associated with margins of lakes and rivers, we found it in several mesic Ontario tallgrass prairies distant from any permanent open water.

Heteroptera (Hemiptera) Numerous species of Heteroptera (Hemiptera) are associated with Ontario tallgrass. Chariesterus antennator (Fabricius) (Coreidae) is restricted to tallgrass prairie and oak savanna in Ontario, where it is normally associated with Euphorbia, though it is known to feed on a variety of plants. All known Ontario sites for this species are significant tallgrass or oak savanna sites (Ojibway Prairie, Manestar Nature Reserve, and Walpole Island). Slater and Baranowski (1978) stated that C. antennator was distributed throughout the eastern and central United States but “scarce and scattered north of the middle Atlantic states.” Three species of tallgrass-associated Pentatomidae occur at Ojibway Prairie. Acantholomidea denticulata is a rare species in Ontario and is apparently “highly remnant-dependent” (Panzer et al. 1995). It feeds on Ceanothus herbaceus Raf. (Slater and Baranowski 1978) and C. herbaceous (Panzer et al. 1995), and it may also feed on C. americanus L., a species found in several Ontario tallgrass prairies. Another pentatomid, Coenus delius (Say), is uncommon in the northeast where it feeds on a variety of grasses (Slater and Baranowski 1978). In Ontario, it is abundant in several tallgrass sites and it is here considered a tallgrass associate. Trichopepla semivittata (Say) feeds on Apiaceae (Slater and Baranowski 1978) in rich meadows or mesic prairies but is uncommon in Ontario except at Ojibway Prairie. Several species of seed bugs (Lygaeoidea) are considered to be tallgrass associates. Ischnodemus falicus (Say) (Blissidae) is an abundant species in Ojibway Prairie where it feeds on Spartina, and it is known in Ontario only from several sites with tallgrass or oak savanna. Panzer et al. (1995) recorded this species as being moderately dependent on prairie remnants. Another species, Cryphula trimaculata (Distant), is rare in Canada (Maw et al. 2000) but widespread in the United States. It apparently feeds on grasses (Slater and Baranowski 1978) but no specific host is recorded. Two other seed bugs, Oedancala dorsalis (usually on Carex) and Slaterobius insignis (Uhler) (on Andropogon), are often found in large numbers in tallgrass sites but often occur in other habitats. Two other Hemiptera, Barce fraterna (Say) (Reduviidae) and Protenor belfragei Haglund (Alydidae), are characteristic bugs of eastern grasslands. Barce fraterna, a widespread but uncommon eastern North American species that commonly occurs within bunches of grasses such as Andropogon gerardii, is the only predator treated here as a tallgrass associate. Protenor belfragei is a sedge and grass feeder (Slater and Baranowski 1978) abundant in stands of Schizachyrium scoparium in tallgrass remnants in southern Ontario. Protenor belfragei also occurs in sedge meadows and similar habitats.

Auchenorrhyncha (Hemiptera) In addition to the tallgrass-dependent and tallgrass-associated Auchenorrhyncha species listed earlier, several other tallgrass-associated Auchenorrhyncha occur in the Ojibway Prairie Complex. Two Phylloscelis species (Fulgoridae), collected on mint species in Ojibway Prairie, are known in Ontario only from tallgrass prairie but are widely distributed over most of the eastern United States (Wilson and McPherson 1980). Phylloscelis pallescens Germar is monophagous on Pycnanthemum tenuifolium Schrad. (Wilson et al. 1993). Two of the fulgoroids that we collected during this study, Megamelus metzaria Crawford () and Aphelonema simplex Uhler (Caliscelidae), were considered Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 207

by Hamilton (2005) as characteristic of eastern prairies. These two species are occasionally found in other grassland types and have distributions not entirely consistent with prairie (Wilson and McPherson 1980). Both species are here considered to be tallgrass associated. Hamilton (2005) recorded M. metzaria on Spartina gracilis and A. simplex on S. gracilis and other grasses. Wilson et al. (1993) recorded A. simplex on S. pectinata Bosc ex Link. One membracid species, Publilia reticulata Van Duzee, is an uncommon northeastern species currently known in Canada only from Ojibway Prairie (Paiero et al. 2004). The distribution of this species is similar to that of the eastern tallgrass prairies (Kopp and Yonke 1973a), and it is reported by Kopp and Yonke (1970) to be a tallgrass species. The wide range of host plants (Kopp and Yonke 1973b), however, indicates that P. reticulata likely occurs in habitats other than tallgrass prairies. An additional seven species (Campylenchia latipes (Say), Acutalis tartarea (Say), Micrutalis calva (Say), Entylia carinata (as E. bactriana Germar), Publilia concava (Say), Stictocephala bubalus (Fabricius), and Ceresa diceros (Say)) were putatively recorded by Kopp and Yonke (1970) as “prairie inhabitants,” but actually are widespread species (Kopp and Yonke 1973a, 1973b, 1973c, 1974) and are not considered here as tallgrass associates.

Coleoptera Of the Coleoptera collected during this study, relatively few species, mostly in the Buprestidae and Carabidae, are here considered to be tallgrass associated. The uncommon buprestid species Pachyschelus laevigatus (Say) and P. purpureus (Say) are recorded on Desmodium and Lespedeza, respectively. Both of these plant genera have species associated with tallgrass prairies and both beetle species have been found only in a few localities in southern Ontario (Bright 1987). Acmaeodera pulchella (Herbst), which develops on herbaceous plants such as Ceanothus and Asclepias (Bright 1987), is known in Canada only from the prairie provinces and a few tallgrass sites in Ontario. Larsen et al. (2003) treated five of the carabid species (Calleida punctata LeConte, Elaphropus granarius (Dejean), Panagaeus fasciatus Say, Stenolophus ochropezus (Say), and Acupalpus carus (LeConte)) that we record from Ojibway Prairie as “prairie-dependent” and three species (Agonum cupripenne (Say), Harpalus erraticus Say, and Stenolophus conjunctus (Say)) as “associated with grasslands.” The latter three species occur in a wide variety of habitats in Ontario and are not considered further here. The former five species are here treated as tallgrass associated rather than as tallgrass dependent. Larsen et al. (2003) do not make a strong case for high habitat specificity in these groups, and other authors (e.g., Lindroth 1961, 1963, 1966, 1968, 1969; Downie and Arnett 1996a) record these species from related habitats. Mawdsley (2003) recorded 18 species of prairie-associated Cleridae, of which two (Enoclerus rosmarus (Say), Placopterus thoracicus (Olivier)) occur in Ojibway Prairie. Both are common and widespread species known from a wide variety of habitats in Ontario and are here considered neither tallgrass dependent nor tallgrass associated. Diabrotica cristata (Harris) (Chrysomelidae) is another species that has been considered as a tallgrass species by some authors (e.g., Wheeler 1988). Campbell and Meinke (2006) further suggested that prairie fragmentation represents a major loss of habitat for D. cristata, but its widespread distribution in the eastern United States (Downie and Arnett 1996b) indicates that it is at most a prairie associate.

Diptera True have been generally neglected in studies of grassland insects, presumably because of their overwhelming abundance and the significant taxonomic impediment to 208 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

the study of many taxa. Some fly groups, however, are well-known and include host- specific grassland species. Several species of here recorded from Ojibway Prairie, including six newly recorded from Canada and two newly recorded from Ontario, represent disjunct populations of species previously known only from grasslands in the United States. Xanthomyia platyptera (Loew) is newly recorded from Canada. The host of this species is currently unknown, but its distribution (Foote et al. 1993) indicates a tallgrass association. Eurosta comma Wiedemann is common in Ojibway Prairie and is newly recorded in Canada. It is relatively common east of the Rockies in the United States, where it develops on Solidago species (Foote et al. 1993). Xanthaciura tetraspina (Phillips), here newly recorded from Canada, develops on Eupatorium and Ageratum (Foote et al. 1993) but is uncommon throughout its range. Tomoplagia obliqua (Say) develops on Aster and Vernonia (Foote et al. 1993) and is a new Canadian record. palposa (Loew), a grassland species that develops on Cirsium (Foote et al. 1993), was previously reported only from western Canada and is here newly recorded from Ontario. Neaspilota reticulata Norrbom and Foote develops on Coreopsis tripteris L. (Norrbom and Foote 2000) and is here newly recorded from Canada. Neaspilota albidipennis (Loew) and N. vernoniae (Loew), also new Canadian records, develop on Vernonia species (Foote et al. 1993). Although these species develop on tallgrass prairie flora, two of the tephritids mentioned (E. comma, X. platyptera) have been collected in non-tallgrass sites in Ontario and have ranges not entirely consistent with eastern tallgrass prairies. The other species have distributions fairly consistent with the tallgrass habitat but also occur in other regions, possibly in artificial grasslands or similar synanthropic habitats. Several other dipteran families, such as the Otitidae, Platystomatidae, and Chloropidae, have some tallgrass-associated species. Ontario’s grassland Otitidae include two species of Tritoxa, one of which (Tritoxa incurva Loew) is here newly recorded from Canada. Tritoxa species are specialists on Allium (Allen and Foote 1975), including the wild onions A. cernuum Roth and A. stellatum Fraser ex Ker Gawl. found in tallgrass prairies. Tritoxa incurva is known from throughout the eastern United States. Several species of Platystomatidae are known from Ontario grasslands, but only two (Rivellia socialis Namba and R. conjuncta Loew) appear to be tallgrass associates. Other Rivellia species are widespread in the east and occur in a variety of grassland habitats. Only one species of tallgrass-associated Chloropidae, Opetiophora straminea Loew, is currently recognized from Ojibway Prairie. This uncommon grassland species was previously recorded from the west-central United States south into Mexico (Sabrosky 1987) and is here newly recorded from Canada. The biology of this species is not known, but its distribution indicates an association with prairies. Chloropidae and other families of Diptera, especially largely phytophagous and frequently host-specific groups such as Cecidomyiidae and Agromyzidae, undoubtedly include further tallgrass-associated species, but work on these groups remains incomplete and many of our specimens are identified only to genus. Some grassland-associated Agromyzidae have been recorded from Ontario alvars (Crecco and Wheeler 2002), and some of the same species probably occur in tallgrass prairies.

Hymenoptera Although and bees abound in tallgrass prairies, few of them appear to be tallgrass associates. Reed (1996) listed several Hymenoptera as “potential prairie specialists,” including five bee species (Colletes compactus Cresson, Melissodes agilis Cresson, M. rustica (Say), Bombus fervidus (Fabricius), and Perdita halictoides Smith)) that occur in the Ontario grasslands that we studied. Most of these species occur throughout eastern North Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 209

America in a variety of habitats, and we consider only P. halictoides as a potential prairie specialist. It is widely distributed in eastern North America, but in Ontario this species is known only from grassland and oak savanna sites. Two ant species (Acanthomyops claviger Roger and Monorium minimum (Buckley)) collected during this study were also regarded by Reed (1996) as potential prairie specialists, but these, too, are widespread species that occur in habitats other than grasslands.

Lepidoptera Only a few hundred species of Lepidoptera, certainly a small fraction of the total fauna present, have been recorded at Ojibway Prairie. Tallgrass associates recorded to date include butterflies such asErynnis baptisiae (Forbes), Achalarus lyciades Geyer (Cossidae), Satyrium edwardsii Grote and Robinson, and S. acadica Edwards (Lycaenidae). Tallgrass associates also include noctuid moths such as Papaipema baptisiae (Bird), which develops in Baptisia; P. cerussata (Grote), which develops in Vernonia; P. pterisii Bird, which develops in Pteridium; P. sciata Bird, which develops in Veronicastrum; and Faronta rubripennis Hodges, which develops in Panicum virgatum. Further sampling of the Lepidoptera is warranted at Ojibway Prairie and additional tallgrass-associated and tallgrass-dependent species will likely be found. Unfortunately, light competition with the adjacent racetrack lights may interfere with sampling (light trapping) of nocturnal species, and light pollution from the same source may negatively affect lepidopteran populations in the area.

Tallgrass Associates Found in Other Ontario Tallgrass Prairies Although Ojibway Prairie has probably now been more thoroughly surveyed than any single grassland site in Canada, the more than 2,000 species listed here do not constitute a comprehensive survey. Several obstacles prevent a complete arthropod inventory of a complex habitat, especially the increasing sampling and processing cost of adding more taxa as an asymptote is approached, as well as the taxonomic impediment that blocks the identification of taxa for which there are neither taxonomic resources nor systematic specialists. Furthermore, insect faunas do not remain the same from year to year and decades may be required to fully inventory even a few selected taxa for only a relatively simple habitat such as an English garden (Owen 1991). In the absence of a complete inventory, it is difficult to distinguish between species that do not occur at the study site and species that are resident but have been overlooked. Nonetheless, some species that occur at other Ontario grasslands appear to be absent at Ojibway Prairie. One provincially rare species of Orthoptera, Pseudopomala brachyptera (Scudder), is not present in Ojibway Prairie though it has been found in several other sites in Ontario. This brachypterous species feeds on Agropyron smithii (Rydb.) A. Löve and Schizachyrium scoparium (Vickery and Kevan 1985), which occur at Ojibway Prairie, and is restricted to only a few tallgrass remnants in southern Ontario, including Pinery Provincial Park and the Simcoe-Delhi railway prairie. Cantrall (1943) reported that tallgrass prairie was P. brachyptera’s major original habitat, but it has expanded its geographical and habitat range to include a variety of grassland habitats across the continent (Vickery and Kevan 1985). Panzer et al. (1995) reported this species as being moderately remnant dependent. Prairie and grassland Auchenorrhyncha (Hemiptera) species occur in other southwestern Ontario grasslands but have not been recorded from Ojibway Prairie. These species include tallgrass-dependent and tallgrass-associated Cercopidae, Caliscelidae, and Cicadellidae (Hamilton 1994, 1995, 2000; Bouchard et al. 2001). 210 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

At least two species of grassland-associated Diptera occur in Ontario at sites other than Ojibway Prairie. One asilid species, Proctacanthella cacopiloga (Hine), is a western species that has disjunct populations in southern Ontario tallgrass and oak savanna habitat (Skevington 1999). This species is common and conspicuous where it occurs. Its absence from Ojibway Prairie probably reflects a lack of exposed sandy soils. One otitid species, Tritoxa flexa (Wiedemann), is known in Ontario from the Stone Road Alvar (Pelee Island), a grassland site with strong faunal similarity to Ojibway Prairie. Tritoxa flexa develops on Allium (Manis 1941) and is found from Manitoba to Connecticut south into Georgia. The bees of Ontario are well-known in comparison to other Hymenoptera. Although the majority of species in Ontario are relatively widespread and not restricted to any specific habitat, one halictid, Dufourea marginata (Cresson), was recently recorded from oak savanna in Ontario by Sugar et al. (1998), who considered it a “prairie specialist.” Its known distribution and recorded pollen sources, Helianthus and Bidens (Mitchell 1961), are consistent with tallgrass dependency. Another Ontario bee species, Bombus fraternus (Smith), was recorded by Panzer et al. (1995) as a tallgrass-dependent species, but the known distribution of this species, Michigan to Florida (Mitchell 1962), and polylectic habits suggest that it is, at most, a tallgrass associate.

New Distributional Records This study has yielded national or provincial records in several orders, some of which appear in recent papers (see Table 1). A further 44 new Canadian and provincial insect records are recorded here for the first time (Table 1), along with notes on known biology and distribution.

New Species In addition to the new distributional records, one new species of fly,Loxocera ojibwayensis Buck (Diptera: Psilidae), was described from Ojibway Prairie (Buck and Marshall 2006), the only known locality for this species despite intensive sampling in southern Ontario and a review of available material from North American collections. No host plant is known for L. ojibwayensis, but its sister species, L. cylindrica Say, develops on Carex (Buck and Marshall 2006), and it is possible that this new species develops on a prairie-associated Carex species. Several other specimens of Diptera and Hymenoptera appear to be undescribed species and will have to be dealt with in the context of future revisions of the taxa involved.

Rarer Species In addition to the tallgrass species, newly described species and new distributional records mentioned earlier, many nationally and provincially rare and restricted species also occur at Ojibway Prairie. We discuss the more noteworthy records here. Four provincially rare Carolinian Orthoptera species occur at Ojibway Prairie. Microcentrum rhombifolium (Saussure) was previously known only in Point Pelee National Park (Vickery and Kevan 1985). Melanoplus walshii Scudder and Neoxabea bipunctata (DeGeer) have both only recently been recorded in Canada from Ojibway Prairie, whereas Oecanthus exclamationis Davis was previously recorded in Canada only from Point Pelee Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 211

National Park and Pelee Island (Marshall et al. 2006). All of these species feed on a variety of shrubs and trees and they occur throughout the eastern United States. Three provincially rare Carolinian Hemiptera have been recorded from Ojibway Prairie. Arilus cristatus (Linnaeus) and Lygaeus turcicus (Fabricius) are both uncommon species in Ontario, where they are known from only a few localities, including Ojibway Prairie. Lygaeus turcicus is recorded feeding on a Helianthus at Ojibway. Arilus cristatus is a well-known predator that is often abundant in the southern United States but rare along the northern fringe of its distribution. Neokolla lugubris (Signoret) is a meadow species that is known in the northeastern United States but is currently known in Ontario only from Ojibway Prairie (Paiero et al. 2004). One rarely collected Mecoptera species, Merope tuber Newman, occurs at Ojibway Prairie. This insect is widely distributed across North America but is rarely collected except in Malaise traps. Thornhill and Johnson (1974) indicate that adults are likely nocturnal and possibly phytophagous but larval natural history is unknown. Several rarely collected species of Neuroptera occur at Ojibway Prairie. Psectra diptera Burmeister is a Holarctic species that is rare throughout its range (Throne 1971) and occurs as either four- or two-winged morphs (Carpenter 1940). Two mantispid species, Leptomantispa pulchella (Banks) and Climaciella brunnea (Say), occur at Ojibway. Leptomantispa pulchella is a species that is widespread in the United States but only ranges into Canada in the southern parts of British Columbia and Ontario (Cannings and Cannings 2006). Climaciella brunnea is widespread in Ontario, but it is not often collected. Provincially rare Lepidoptera include butterflies such as Papilio cresphontes (Cramer), Euphyes dukesi (Lindsey), Poanes massasoit (Scudder), and Asterocampa celtis (Boisduval and Le Conte). The underwing moth Catocala connubialis Guenée, which develops on oaks, has been recorded at only one other location in Ontario. Several provincially rare species of Asilidae (Diptera) were collected. Three Carolinian species, Holcocephala calva (Loew), Cerotainia albipilosa Curran, and C. macrocera (Say), are known from only a few localities in southwestern Ontario (Skevington 1999). Two further species, Promachus vertebratus (Say) and Asilus sericeus Say, are uncommon Carolinian species here recorded from Ojibway Prairie. Four other fly species, Amphicnephes pullus (Wiedemann) (Platystomatidae), Tabanus trimaculatus Palisot de Beauvois, Chrysops brunneus Hine (Tabanidae), and Heteromeringia nitida Johnson (), are widespread in the eastern United States but are known from only a few Carolinian sites, including Ojibway Prairie in Ontario. Heteromeringea nitida was only recently recorded from Canada (Lonsdale and Marshall 2007). A significant number of rare species of Hymenoptera were collected during this study. Olixon banksii (Brues) (Rhopalosomatidae) had not yet been officially recorded from Ontario, though Brothers and Finnamore (1993) make mention of one species of Rhopalosomatidae in “extreme southern Ontario.” This species, though rare in collections, seems to be associated with oak savanna, and Krauth (2000) also recorded it from a tallgrass–oak savanna habitat. It is parasitic on Blattodea (Krombein 1979c). Sierolomorpha canadensis (Provancher) (Sierolomorphidae) is a widely distributed species in North America, but it is rarely collected and little is known about its biology. Another species, Paratiphia texana Cameron (Tiphiidae), is an uncommon Carolinian species in Ontario that is known from only a few other sites in southern Ontario (Allen 1966). A total of 115 species of spheciform wasps are recorded from the Ojibway Prairie Complex, more than the total currently recorded from any other site in Ontario. Buck (2004) reviews the spheciform wasps of Ontario, of which three species, Tachysphex apicalis Fox, 212 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

Recorded from several tallgrass several from Recorded sites in southwestern Ontario Introduced species. Collected from several other Carolinian Ontario sites Recorded from several Carolinian Ontario sites Known from one other site in southern Ontario. New subfamily record for eastern Canada Range has been slowly expanding since the mid- 1900s. Known in Canada only from Ojibway Prairie Recorded from several Carolinian Ontario sites tallgrass several from Recorded sites in southwestern Ontario tallgrass several from Recorded sites in southwestern Ontario Now known from several other Carolinian Ontario sites

Wheeler 1984 Paiero et al. 2004 Paiero et al. 2004 Marshall et al. 2006 Marshall et al. 2006 Marshall et al. 2006 Marshall et al. 2006 Pratt and Paiero 2004 References Notes Northeastern USA Hoebeke and Eastern and central USA Eastern USA Eastern USA Eastern USA Eastern USA Eastern USA south into Eastern USA Mexico Eastern USA Previously Previously Documented American North Distribution CAN CAN CAN CAN CAN CAN CAN CAN Distribution Distribution Record CAN Associated with Euphorbia Associated with Fagus and Quercus Found in colonies of various ant species

Aethus nigritus (Fabricius)* Chariesterus antennator (Fabricius) Neuroctenus simplex Neuroctenus (Uhler) Dichromorpha Dichromorpha viridis (Scudder) Melanoplus scudderi scudderi (Uhler) Melanoplus walshi (Scudder) Neoxabea bipunctata (DeGeer) Myrmecophila Bruner* pergandei Archilestes grandis Archilestes Rambur Cynidae Coreidae Aradidae Gryllidae

Hemiptera (Heteroptera) Orthoptera Acrididae Order FamilyOdonata Species Lestidae Biology A list of new or recent Canadian and Ontario records based all or in part on specimens collected at Ojibway Prairie, Windsor, Ontario, Canada, between 2001 and 2007. Windsor, list of new or recent Canadian and Ontario records based all in part on specimens collected at Ojibway Prairie, 1. A Table Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 213 Recorded from several southwestern Ontario sites Recorded from several southwestern Ontario sites Recorded in Canada only from Ojibway Prairie Recorded from several Ontario sites Recorded in Canada only from Ojibway Prairie Ojibway Prairie Recorded in Canada only from Ojibway Prairie Recorded in Ontario only from Ojibway Prairie Known in Ontario only from Ojibway Prairie Recorded from several Carolinian Ontario sites Recorded from several Carolinian Ontario sites Recorded from several southwestern Ontario sites

Paiero et al. 2004 Paiero et al. 2004 Paiero et al. 2004 Paiero et al. 2004 Paiero et al. 2004 Paiero et al. 2004 Sauer 2002 Recorded in Ontario only from Paiero et al. 2004 Paiero et al. 2004 Paiero et al. 2004 Paiero et al. 2004 Eastern USA Eastern USA Northeastern USA SK to NS, south to WY WY SK to NS, south and MO CT Northeastern USA AB to MB into eastern USA Eastern USA Eastern USA Quebec; widespread in USA CAN CAN CAN ON CAN CAN ON E CAN MB to KS, east NY E CANWI MB, SK, and CAN CAN CAN On various woody plants On various woody plants On various deciduous trees On Spartina Tallgrass Tallgrass associate; recorded on Steironema, Glaux, or Lysimachia Tallgrass Tallgrass associate Carolinian species Carolinian species

Ormenoides venusta (Melichar) Anormenis septentrionalis (Spinola) Anotia westwoodi Fitch Megamelus metzaria Crawford Neokolla lugubris (Signoret) Delphacodes waldeni Metcalf Destria fumida (Sanders and DeLong)* Erythroneura carbonata McAtee Cuerna fenestella Hamilton Dendrocoris Dendrocoris humeralis (Uhler) Hoplistoscelis Reuter sordidus Pangaeus bilineatus (Say) Flatidae Derbidae Delphacidae Cicadellidae Pentatomidae Nabidae

Hemiptera ("Homoptera")

214 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck Known in Canada only from Ojibway Prairie Known in Canada only from Ojibway Prairie This species has been collected at one other site in “Windsor.” Known in Ontario only from Ojibway Prairie Ojibway Prairie Recorded in Ontario only from Ojibway Prairie Recorded in Canada only from Ojibway Prairie This species has been collected at one other southwestern Ontario site. This species has been collected at several other Carolinian Ontario sites. Recorded in Canada only from Ojibway Prairie

Downie and Arnett 1996 b Downie and Arnett 1996 a Downie and Arnett 1996 b Downie and Arnett 1996 b Cannings and Cannings 2006 Downie and Arnett 1996 a ; 2005 Westcott Downie and Arnett 1996 a as Molumba lunata ; Downie and Arnett 1996 b Kozár 2004 Known in Canada only from Paiero et al. 2004 References Notes MI and NY Eastern and central USA throughout and Manitoba the eastern USA Eastern USA from NY from NY Eastern USA to FL in USA into South in USA America Eastern USA AR, IL, MA, FL, TX and IN, FL, NY, CT, and MO Eastern USA, AZ, and Eastern USA, UT Eastern USA Previously Previously Documented American North Distribution CAN CAN ON CAN E CANCAN BC, ON, widespread CAN CAN CAN CAN Distribution Distribution Record Quercus On Quercus coccinea On several species of Asteraceae and Rosaceae On various herbaceous plants

(Banks) Tritoma angulata Tritoma Say* Photinus pyralis Linnaeus* Epicauta atrata (Fabricius)* Leptomantispa pulchella Mastogenius Knull crenulatus Chauliognathus marginatus (Fabricius)* Clypestraea lunata (LeConte)* Cryptorrhynchus tristis LeConte* Orthezia solidaginis Sanders* Publilia reticulata Publilia reticulata VanDuzee Erotylidae Lampyridae Meloidae Cantharidae Corylophidae Curculionidae Ortheziidae Membracidae

Neuroptera Mantispidae Coleoptera Buprestidae

Order Family Species Biology Table 1 (continued) Table Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 215 This species has been collected at one other southwestern Ontario site. Recorded in Canada only from Ojibway Prairie This species has been collected at one other southern Ontario site. This species has been collected at one other Carolinian Ontario site. Known in Canada only from Ojibway Prairie This species has been collected at one other southwestern Ontario site. Known in Canada only from Ojibway Prairie This species has been collected at several other Carolinian Ontario sites. This species has been collected at several other Carolinian Ontario sites. Greathead 1999; Kits et al. 2008 Evenhuis and Greathead 1999; Kits et al. 2008 Evenhuis and Greathead 1999; Kits et al. 2008 Arnett 1996 b Downie and Arnett 1996 a Arnett 1996 b Downie and Arnett 1996 b Baker and Fischer 1975 Eastern USA into TX into Eastern USA Evenhuis and Midwestern and eastern USA Midwestern and eastern USA MI, LA, and FL MI, LA, and FL Eastern and central Downie and USA AL and ME, NY, Downie and NJ, IN, OH, IN NY, TX and south into FL and west to ND MO where it is noted NY as a “recent colonizer” Wheeler (McCabe and 1994) from the south and is recorded from MI where it prefers “moist luxuriant meadows” (Baker and Fischer 1975) Central and eastern USA CAN CAN CAN CAN CAN CAN CAN CAN CAN Parasitic on solitary Hymenoptera Parasitic on Noctuidae Parasitic on Apoidea

(Say)*

Macquart Toxophora aphitea Toxophora Walker Poecilanthrax bicellata Tmesiophorus carinatus (Say)* Rhipiphorus luteipennis (LeConte)* Platydema elliptica (Fabricius)* Ommatius tibialis (Say)* Taractius octopunctatus Anthrax pluto Wiedemann Temnopsophus Temnopsophus bimaculata Horn* Pselaphidae Rhipiphoridae Tenebrionidae Bombyliidae Melyridae

Diptera Asilidae

216 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck Known in Canada only from Ojibway Prairie Known in Canada only from Ojibway Prairie Known in Canada only from Ojibway Prairie This species has been collected at several other Carolinian Ontario sites. This species, known in Canada only from Ojibway Prairie, was previously thought to be cavernicolous but probably occurs in mammal burrows as well as in caves. This species has been collected Ontario. in sites other several at Ojibway Prairie Ojibway Prairie This species has been collected at several other Carolinian sites in southwestern Ontario. This species has been collected at at collected been has species This one other Carolinian Ontario site.

Foote et al. 1993; Norrbom and Foote 2000 Norrbom and Foote 2000 Foote et al. 1993 Foote et al. 1993 Reeves 2003 2001 Rohácek et al. Namba 1956 Known in Canada only from Namba 1956 Known in Canada only from Stone et al. 1965 Stone et al. 1965 References Notes Eastern USA MI and OH Eastern USA Eastern USA MD, GA, and SC Marshall and NB, MA, and TN NB, MA, and CT, KS, MA, NY, NC, KS, MA, NY, CT, OK, TX and VA, IA and IL IA Eastern USA UT to MN and MI UT south to Mexico Previously Previously Documented American North Distribution CAN CAN CAN CAN CAN CAN CAN CAN CAN Distribution Distribution Record CAN On Vernonia Coreiopsis On Coreiopsis tripteris On Vernonia On Solidago A cavernicolous A species No host plant known No host plant known On Allium

Neaspilota vernoniae Neaspilota (Loew)* Neaspilota reticulata Neaspilota reticulata Norrbom and Foote* Neaspilota albidipennis (Loew)* Eurosta comma Eurosta Wiedemann* Spelobia pickeringi Marshall* Bromeloecia seltzeri Bromeloecia Marshall* Rivellia conjuncta Loew* Rivellia socialis Namba* Tritoxa incurva Tritoxa Loew* Opetiophora straminea Loew* Tephritidae Sphaeroceridae Platystomatidae Otitidae Chloropidae

Order Family Species Biology Table 1 (continued) Table Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 217 Introduced species. Collected from one other southwestern Ontario site Now known in Canada from several other Carolinian Ontario sites Known in Ontario only from Ojibway Prairie This species has been collected at one other Carolinian Ontario site. This species has been collected at several other Ontario sites. This species has been collected at one other Ontario locality (in central Ontario). additional Ontario sites additional Ontario sites one other Carolinian Ontario site one other Carolinian Ontario site additional Ontario sites additional Ontario sites Kimsey 1982 Paiero and Buck 2004 Stone et al. 1965 Foote et al. 1993 Foote et al. 1993 Foote et al. 1993 Buck 2004 Recorded from several Buck 2004Buck 2004 Recorded from several Now known in Canada from Buck 2004 Now known in Canada from Buck 2004 Recorded from several Widespread in USAWidespread Bohart and Eastern USA BC to MB, north Baffin Island, through Atlantic provinces, the Also and into NY. occurs in Europe Eastern USA Eastern and central USA Eastern USA BC, AB, and BC, widespread in USA AB, and BC, transcontinental USA USA east of Rocky USA Mountains in USAWidespread Buck 2004USA Recorded from several CAN CAN ON CAN CAN CAN ON ON E CANCAN SK and throughout ON E CAN MB, SK, and eastern Parasite of Sceliphron (Sphecidae) Occurs in alkaline areas and has been found in salt marshes and in beach grasses No host plant known On Eupatorium and Ageratum On Aster and Vernonia

(Loew)* Chrysis fuscipennis Brullé* Perdita bequaerti Perdita bequaerti Viereck Xanthomyia platyptera Pelomyiella mallochi (Sturtevant)* Xanthaciura tetraspina (Phillips)* Cerceris crucis Cerceris and Viereck Cockerell Cerceris echo Cerceris Michel finitima Cerceris Cresson insolita Cerceris Cresson kennicottii Cerceris Cresson snowii Fox Crabro Tomoplagia obliqua Tomoplagia (Say)* 1 Chrysididae Andrenidae Tethinidae

Hymenoptera

218 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck additional Ontario sites additional Ontario sites Ojibway Prairie additional Ontario sites additional Ontario sites additional Ontario sites additional Ontario sites additional Ontario sites Known in Canada only from Ojibway Prairie Ojibway Prairie additional Ontario sites additional Ontario sites Introduced species. Recorded in Canada only from Ojibway Prairie Buck 2004Buck 2004 Recorded from several Buck 2004 Recorded in Canada only from Recorded from several Buck 2004Buck 2004 Recorded from several Buck 2004 Recorded from several Recorded from several Buck 2004Buck 2004 Recorded in Ontario only from Buck 2004 Recorded from several Romankova Recorded from several 2004 Buck 2004 Recorded from several References Notes Eastern USA Eastern USA Eastern USA NB and eastern USA Buck 2004Eastern USA Recorded from several Central and eastern USA Eastern USA Eastern USA throughout USA Eastern USA Eastern USA Northeastern USA and ON Rocky Mountains Previously Previously Documented American North Distribution CAN CAN CAN CAN CAN CAN CAN CAN E CANCAN AB, BC, and CAN CAN Distribution Distribution Record E CAN east of AB and USA

Fox (Fox) Ectemnius scaber (Lepeletier and Brullé) Entomognathus lenapeorum Viereck Entomognathus memorialis Banks Mimumesa longicornis (Fox) Nysson simplicicornis Oxybelus cressonii (Robertson) Oxybelus decorosus (Mickel) Pseneo kohlii apicalis Tachysphex Fox crassus Tachytes Patton intermedius Tachytes (Viereck) Anthidium oblongatum (Illiger) Ectemnius dilectus (Cresson) Megachilidae

Order Family Species Biology Table 1 (continued) Table Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 219 Known in Canada only from Ojibway Prairie This species has been collected at two other Carolinian Ontario sites. Recorded from several other sites in Ontario collected at one other Carolinian Ontario site. This species has been collected at one other Carolinian Ontario site. This species has been collected at several other Carolinian Ontario sites. Recorded in Canada from only one other southwestern Ontario site This species has been collected at one other Carolinian Ontario site. This species has been collected at two other Carolinian Ontario sites. This species has been collected at several other Carolinian Ontario sites. This species has been collected at one other Carolinian Ontario site. Recorded from several other sites in Ontario Introduced species. Recorded from several other sites in Ontario Krombein 1979 c Evans 1997 Krombein 1979 b Krombein 1979 b Krombein 1979 b Evans 1950 This species has been Krombein 1979 a Krombein 1979 b Krombein 1979 b 1979 b Krombein 1979 b Paiero and Buck Buck and Paiero 2004 (as S. costalis Cresson) Paiero and Buck 2004 NC, SC, TX, KS, and IL NC, SC, CO, ID, and MI 1986; Wasbauer Eastern USA MS to NC; west to IL MS to NC; west IL and IN to FL; west KS NY TX and MI, NY; west to TX west to MI, NY; and CO MA to FL, west NE MA TX and and west to GA NY to CO east of Rocky USA Mountains (transcontinental)USA Krombein Northeastern USA (including MI and NY) Eastern USA Eastern USA CAN CAN CAN CAN CAN CAN CAN CAN CAN CAN CAN CAN CAN Carolinian species Carolinian species Carolinian species Cleptoparasite on megachilid bees

Priocnemis kevini Wasbauer* Priocnemis minuscula (Banks)* Dipogon calipterus (Say)* Entypus unifasciatus (Say)* Minagenia osoria (Banks)* Pseudomethoca sanbornii (Blake)* Ageniella cupida (Cresson)* Ageniella fulgifrons (Cresson)* Allaporus pulchellus (Banks)* Anoplius hispidulus Dreisbach* Anoplius percitus Evans* Stelis louisae Cockerell Megachile sculpturalis Smith Mutillidae Pompilidae

220 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck Amaurochroa Amaurochroa This species has been collected at two other Carolinian Ontario sites. This species has been collected at several other Carolinian Ontario sites. This species has been collected at several other Ontario sites. This species has been collected at several other Ontario sites. This species has been collected at several other Carolinian Ontario sites. This species has been collected at several other Ontario sites. This species has been collected at several other Ontario sites. et et et et

Krombein Buck 1979 c ; al. 2008 ; Buck 1979 c ; al. 2008 Buck 1979 c ; al. 2008 Krombein Buck 1979 c ; al. 2008 2008 Parker 1966; Krombein 1979 c (as L. ornatus Saussure); Buck et al. 1979 c 1979 a References Notes . Amaurochroa cinctipes Amaurochroa Eastern USA QC and eastern USA Krombein MB and eastern USA Krombein Eastern USA MA to FL, NE, CO, MA TX KS, and Northeastern USA and DC NY, Krombein VA, Krombein Previously Previously Documented American North Distribution CAN CAN CAN CAN CAN CAN CAN Distribution Distribution Record Carolinian species Carolinian species Carolinian species Carolinian species (2004). These specimens were in fact et al. (2004). Parazumia symmorpha (Saussure) Stenodynerus ammonia paraensis (Saussure) Stenodynerus blepharus Bohart Parancistrocerus Parancistrocerus (Saussure) perennis Methocha impolita Krombein* Leptochilus republicanus Torre) (Dalla Tachypompilus Tachypompilus ferrugineus (Banks)* nigrescens

2 Tiphiidae Vespidae The species is here recorded from Ontario or Canada for the first time. Distributional record abbreviations: (CAN), eastern (E CAN), Ontario (ON). The species is here recorded from Ontario

Order Family Species Biology ovalis was erroneously recorded from Canada in Paiero An additional two Sphecidae and 25 Crabronidae species were also recorded as new to either Canada or Ontario by Buck (2004) from numerous localities, including Ojibway species were also recorded as new to either Canada or Ontario by Buck (2004) from numerous localities, including An additional two Sphecidae and 25 Crabronidae Prairie, but are not included here as they widespread in Ontario. For further information, see Buck (2004). in part on material from Ojibway Prairie (specific localities not given). Note: and Canada by Buck et al. (2008) based Recorded for the first time from Ontario * Table 1 (continued) Table 1 2 Insects of Ojibway Prairie, a Southern Ontario Tallgrass Prairie 221

Entomognathus lenapeorum Viereck, and Cerceris insolita Cresson, are currently known in Ontario (or Canada) only from Ojibway Prairie (the latter was subsequently also discovered on Walpole Island). Since 2004, another species, Pseneo kohlii (Fox), was added to this list. Other rare species that occur in few Ontario localities besides Ojibway Prairie are Mimumesa longicornis (Fox), Tachytes crassus Patton, T. intermedius (Viereck), Oxybelus cressonii (Robertson), O. decorosus (Mickel), Entomognathus memorialis Banks, Crabro snowii Fox, Ectemnius dilectus (Cresson), E. scaber (Lepeletier and Brullé), Nysson simplicicornis Fox, Cerceris crucis Viereck and Cockerell, C. echo Michel, C. finitima Cresson, and C. kennicottii Cresson. The high number of provincially or nationally rare species is remarkable and unparalleled at any other site in Ontario. Structurally rich oak savannas and savanna-like habitats, especially if interspersed with small, sparsely vegetated areas or sandy patches, represent the habitat type that supports the greatest diversity of species of spheciform wasps in Ontario.

Conclusions Tallgrass prairie sites in southern Ontario support a significant number of highly habitat- restricted species, many of which are rare on a provincial, national, or even global level. The many rare insect species we here record from Ojibway Prairie, several of which are newly recorded for Canada or for Ontario, suggest that this site is one of the most important biological hotspots in Canada and possibly the most significant site for the preservation of provincially rare species in Ontario. Furthermore, the significant numbers of prairie-dependent and prairie-associated species we here document from Ojibway provide unequivocal evidence for the prairie peninsula. Further data from other tallgrass prairie habitats in southern Ontario are needed to fully document the distribution of these habitat-restricted insects. Most of the taxa discussed earlier would have been difficult to recognize or assess outside of the context of a thorough insect survey. Although the 2,217 taxa now identified from Ojibway Prairie represent a more comprehensive inventory than has been developed for any comparable site in Canada, the total arthropod fauna of the Ojibway Prairie Complex probably exceeds 3,000 species. Further sampling is needed by using other techniques (light traps, soil extraction, bait traps, and emergence traps), combined with resolution of taxonomic groups not dealt with in this study (such as most sternorrhynchan Hemiptera, nematoceran Diptera, Hymenoptera Parasitica, most microlepidopterans, and Collembola, along with spiders and mites), to test this prediction and to document the presence of further grassland species in Ontario.

Acknowledgements We thank the World Wildlife Fund and the Ontario Natural Heritage Information Centre for supporting this project and providing site information about grasslands in Ontario. We thank the staff at the Ojibway Nature Centre, Windsor Parks Department, for their assistance in maintaining and servicing the traps and S. Hayes for preparing the map. We relied on collaborating systematists for the identification of many of our specimens, and we gratefully acknowledge the following collaborators for their contributions: K.G.A. Hamilton (Auchenorrhyncha), H. Goulet (Symphyta and Carabidae), L. Masner (Proctotrupoidea), O. Lonsdale (Diptera and Coleoptera), R.A. Anderson (Curculionidae), T. Romanokova (apiform Apoidea), J. Skevington (Pipunculidae), M. Parchami-Araghi (Oestroidea), J. Llyod (Lampyridae), and J. Kits (Bombyliidae). 222 S. M. Paiero, S. A. Marshall, P. D. Pratt, and M. Buck

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