<<

Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

138 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

Coleoptera Collected Using Three Trapping Methods at Grass River Natural Area, Antrim County, Michigan Robert A. Haack1, * and William G. Ruesink2 1 USDA Forest Service, Northern Research Station, 3101 Technology Blvd., Suite F, Lansing, MI 48910 (emeritus) 2 Illinois Natural History Survey, 1816 S Oak St, Champaign, IL 61820 (emeritus) * Corresponding author: (e-mail: [email protected])

Abstract Overall, 409 Coleoptera (369 identified to species, 24 to only, and 16 to subfamily only), representing 275 genera and 58 families, were collected from late May through late September 2017 at the Grass River Natural Area (GRNA), Antrim Coun- ty, Michigan, using baited multi-funnel traps (210 species), pitfall traps (104 species), and sweep nets (168 species). All three collecting methods were used in three distinct habitats: a rich conifer swamp (cedar), near the edge of a red pine plantation (pine), and within a mesic northern hardwood forest (hardwoods). Additional collections were made along two trails and in an open field by sweep netting only. Of the 409 species, 322 were collected in one or more of the cedar, hardwoods, and pine habitats, and 152 were collected along the two trails and the grassland site. Of the 322 species collected in the three main habitats, 40 species (36 genera and 14 families) were collected in all three habitats, 105 species (80 genera and 32 families) were collected in the cedar, 176 (131 genera and 38 families) in the hardwoods, and 199 (158 genera and 47 families) in the pine habitats. With respect to adult seasonal activity, 21% of the 409 species were first collected in May, 40% in June, 23% in July, 10% in August, and 6% in September. Of the 210 species collected in funnel traps, 144, 123, and 114 species were collected, respectively, in traps baited with α-pinene, ethanol, or ipsenol. Diversity indices were calculated for the funnel trap data by site and lure. Overall, 32 of the 409 species were considered exotic to North America, and 18 were considered new state records for Michigan. In addition, 16 species of aquatic (12 genera in 5 families) were identified from GRNA stream samples collected during 2013–2019. Keywords: beetles, diversity index, state records, funnel trap, pitfall trap, sweep net

Interest in and con- on collections in both Michigan’s Lower and servation has grown in recent decades along Upper Peninsulas, including Isle Royale with recognition of the many important eco- in Lake Superior where they recorded 123 system services that provide (Foottit beetle species. Townsend (1889) published and Adler 2017, Samways 2019). Moreover, a list of beetles (161 species) collected in St. recent reports of global insect declines have Joseph County, MI. Later, Adams (1909) heightened public awareness of the threats and Wolcott (1909) added several more spe- faced by many insects (van Klink et al. 2020). cies to the list of beetles known to occur on Just over one million species of insects have Isle Royale, raising the total to 206 species. been described worldwide (Zhang 2011), Andrews (1916) published an extensive including nearly 400,000 species of beetles list of beetle species (623 species) found on (Coleoptera) (Bouchard et al. 2017). In North the Charity Islands in Saginaw Bay, Lake America north of Mexico, over 25,000 beetle Huron. Andrews (1921) published another species have been described (Marske and long list of beetle species (886 species) found Ivie 2003), and in Michigan there are about at Whitefish Point in the Upper Peninsula 4000 beetle species recorded (W. G. Ruesink of Michigan. One additional early list of et al., unpublished data). Michigan beetles (580 species) was published Beetle surveys in various parts of by Hatch (1925) for Charlevoix County, Michigan have been published since the including Beaver, Garden and Hog Islands late 1800s. Schwarz (1876) published one in Lake Michigan. More recently, several of the first lists based on beetles collected authoritative lists of Michigan beetles have mostly in the Detroit area. Two years later, been published for specific families or sub- Hubbard and Schwarz (1878) published a families, such as (Wellso et al. more extensive list (> 2000 species) based 1976), Cerambycidae (Gosling 1973, 1983;

Published by ValpoScholar, 2020 1 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 139

Figure 1. Map of Grass River Natural Area (GRNA), Antrim County, MI, showing all property as of 2020 except for a few parcels at the north end of Lake Bellaire. The green-col- ored parcels are open to hunting, while the orange-colored area is closed to hunting. Labels for the 2017 sampling sites are: C = cedar habitat, G = grassland, H = hardwoods, P = pine, RT = Rail Trail, and SMT = Sedge Meadow Trail. The three creeks where sampling occurred are marked (Cold, Finch, and Shanty Creeks). The red star inside the inset map of Michigan is the approximate location of GRNA.

Gosling and Gosling 1977), (Grey et al. 2017). Using MNFI terminology the and Cognato 2019), (Gosling 1980), nine natural communities (habitats) found at Scolytinae (Cognato et al. 2009), Tenebrion- GRNA, in decreasing order of size as estimat- idae (Spilman 1973), and several families of ed in 2017, were rich conifer swamp (503 ac; aquatic beetles (Bright 2020). 204 ha), mesic northern forest (283 ac; 115 ha), northern fen (185 ac; 75 ha), poor coni- The Grass River Natural Area (GRNA) fer swamp (82 ac; 33 ha), hardwood-conifer is in Antrim County in northwestern Lower swamp (53 ac; 21 ha), northern wet meadow Michigan, where it straddles much of the (49 ac; 20 ha), northern shrub thicket (41 Grass River that connects Lake Bellaire to ac; 17 ha), dry-mesic northern forest (30 Clam Lake (Fig. 1). GRNA began in 1969 ac; 12 ha), and emergent marsh (6 ac; 2 ha) with a single 62 ac (25 ha) parcel and has (Hackett et al. 2017). now expanded to 1492 ac (603 ha), consisting of 73 discrete and mostly contiguous land Beetles can be collected in a variety of parcels (GRNA 2020). The staff at GRNA ways with equipment such as sweep nets, welcomes and supports on-site field research beating sheets, light traps, pitfall traps, (GRNA 2020). baited or unbaited flight-intercept traps, Berlese funnels, and Malaise traps (Dillon The Michigan Natural Features In- and Dillon 1972, White 1983, Evans 2014). ventory (MNFI) has identified 77 natural In the present study, we used sweep nets, community types in Michigan (Kost et al. pitfall traps, and baited multi-funnel traps 2007, Cohen et al. 2015), of which 9 were to collect beetles at GRNA in 2017. In this identified by MNFI staff at GRNA (Hackett paper, we provide a list of the beetle species

https://scholar.valpo.edu/tgle/vol53/iss2/9 2 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

140 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

collected with details on their habitat asso- pitfall traps, and sweep netting. Sampling ciations, seasonality, methods by which they with funnel traps occurred from 19 May to were collected, and status as being native 24 September 2017, and similarly from 19 or exotic to the United States as well as a May to 22 September 2017 for pitfall traps, new state record for Michigan. In addition, and from 23 May to 20 September 2017 for diversity indices were calculated for the sweep netting. Starting in June or July, re- funnel trap data. spectively, the pitfall traps and funnel traps were “closed” for two weeks each month, Methods and Materials usually the first two weeks of each month. Sweep netting occurred at all sites, starting Habitats sampled. Trapping and in May or June, and occurred usually two to sweep netting was conducted in three GRNA three times per month, including the trails habitats, including rich conifer swamp and grassland site. (which we refer to as “cedar”), mesic north- ern forest (hardwoods), and along the edge Funnel traps. At each of the three of a mature, red pine (Pinus resinosa Sol. main collecting sites, three 12-unit multi-fun- ex Aiton) plantation (pine) (Fig. 1). The rich nel traps (Contech Enterprises Inc., Victoria, conifer swamp was dominated by northern British Columbia, Canada) were deployed. white cedar (Thuja occidentalis L.), with oth- The color of the funnels was green as shown er occasional tree species such as tamarack in Petrice and Haack (2015). The individual [Larix laricina (Du Roi) K. Koch], balsam fir funnels in all traps were coated with Fluon [Abies balsamea (L.) Mill], red maple (Acer (Northern Products Inc., Woonsocket, RI), a rubrum L.), yellow birch (Betula alleghanien- slippery substance that improves trapping sis Britton), and black ash (Fraxinus nigra efficiency (Graham et al. 2010). The traps Marshall). The mesic northern forest was were suspended from lower branches of trees dominated by hardwood trees such as sugar so that the bottom of the collection cup was maple (Acer saccharum Marshall) and Amer- about 1 m above groundline. To make sure ican beech (Fagus grandifolia Ehrh.), with the traps were clearly visible to flying in- occasional yellow birch, northern red oak sects, any interfering branches were pruned (Quercus rubra L.), eastern hemlock [Tsuga away. The distance between traps at each canadensis (L.) Carrière], and white pine site ranged from 4–25 m, depending on the (Pinus strobus L.). Along the edge of the red location of suitable trees. The collection cup pine plantation were occasional red maple, at the bottom of the funnel trap was fitted black cherry (Prunus serotina Ehrh.), and with a small screen to allow rainwater to white pine. A detailed floristic description of drain. Inside each collection cup, a circular each GRNA habitat is given in Hackett et al. piece of window screening was fitted near (2017). The Latitude-Longitude coordinates the bottom that suspended captured insects of the three main trapping sites were: N above any moisture that accumulated at 44.9135° Lat and W 85.2186° Long for the the base of the cup. A few pieces of No-Pest cedar site, N 44.9128° Lat and W 85.2246° Strip (Spectrum Group, St. Louis, MO), were Long for the hardwood site, and N 44.9050° placed inside each collection cup on top the Lat and W 85.2223° Long for the pine site. screen to quickly kill trapped insects. The Sweep netting was also conducted active ingredient in No-Pest Strips is dichlor- along two GRNA trails (Rail Trail and Sedge vos, an organophosphate insecticide. Three Meadow Trail) and in an open field (grass- different lures were used at each site, using land; N 44.9100° Lat and W 85.2333° Long) one lure per trap, including an α-pinene UHR (Fig. 1). The Rail Trail occurs along the for- (ultra-high release) pouch with a release mer Pere-Marquette Railway track bed and rate of 2.3 g per day at 26°C (Alpha Scents, typically grades on each side from grasses, Inc., West Linn, OR), an ethanol UHR pouch forbs and shrubs to trees that reflect the with a release rate of about 300 mg per day adjacent habitats. The Sedge Meadow Trail at 20°C (Contech Enterprises Inc.), and a traverses three habitats: rich conifer swamp, racemic ipsenol bubble-cap with a release hardwood-conifer swamp, and northern wet rate of about 0.1–0.2 mg/d at 25°C (Contech meadow. The hardwood-conifer swamp was Enterprises Inc.). These lures are common dominated by northern white cedar, yellow attractants to a wide range of bark- and birch, balsam poplar (Populus balsamifera wood-infesting insects and their associates L.), and bigtooth aspen (Populus grandiden- (Miller et al. 2015, Millar and Hanks 2017, tata Michaux), whereas the northern wet Rabaglia et al. 2019). The lures and No-Pest meadow was dominated by sedges, grasses, Strips pieces were changed at approximately and occasional small shrubs (Hackett et al. 6-week intervals. At the end of a sampling 2017). period, all insects were removed and placed Sampling methods and frequen- in a labeled zip-lock plastic bag and frozen cy. The three main methods of collecting until sorted. The collection cup was then beetles at GRNA involved funnel traps, cleaned, the screen and No-Pest Strips re-

Published by ValpoScholar, 2020 3 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 141

positioned in the cup, and then the cup was part of the MiCorps Volunteer Stream Mon- reattached to the trap. itoring Program, sponsored by Michigan’s Pitfall traps. Two pitfall traps were Department of Environment, Great Lakes, installed at the three main collection sites. and Energy (https://micorps.net/). Beetles Each trap consisted of an 18 oz (0.5 l) plastic were removed from the stored vials, pinned, cup with a ca.10-cm diameter opening that labelled, and later identified. was sunk into the ground so that the top was Specimen identification and loca- flush with the soil surface. Two 1-m-long bar- tion. Nearly all beetles were identified by riers, made from black plastic lawn edging, WGR using an AmScope 7X-90X binocular were partially buried on opposite sides of zoom microscope (AmScope, Irvine, CA, USA) the cup and positioned flush with the cup’s and the keys in Downie and Arnett (1996). rim at the hardwoods and pine sites, but The identifications were checked against not the cedar site because the sphagnum information and photographs on BugGuide surface there was too irregular for the edging (https://bugguide.net/). In cases where some to make an effective barrier. Such barriers doubt existed regarding the identification, direct ground invertebrates towards the cup literature suggested on BugGuide as well and have been shown to increase trap catch as the keys in Arnett and Thomas (2001) (Durkis and Reeves 1982, Hansen and New and Arnett et. al. (2002) were consulted. In 2005, Skvarla et al. 2014). At the start of a addition, for some beetle groups, taxonomic sampling period each cup was cleaned and advice and assistance were provided by then filled to a depth of ca. 5 cm with 70% national experts (see Acknowledgments). ethanol. To reduce dilution from rain, a 25- The specimens discussed in this paper are cm diameter plastic plate was placed ca. 5 cm currently in the authors’ private collections above each collection cup, supported by large but later a few will be retained at GRNA nails. At the end of each sampling period, all with the vast majority being deposited in beetles were removed and placed in labeled the Michigan State University, Department vials with fresh 70% ethanol and then stored of Entomology, Albert J. Cook until mounted for identification. Research Collection in East Lansing, MI. Sweep netting. A standard 15-inch New Michigan state records. The (38 cm) diameter heavy duty sweep net with status of each beetle species as being a a sailcloth bag was used. Typically, 10-30 potential new state record for Michigan min was spent sweeping each site when was based on data presented in the sever- visited. Most sweeping was conducted on al authoritative lists of Michigan beetles non-woody vegetation, but in each habitat mentioned in the introduction plus over 350 about ten percent of all sweeps consisted other publications that have been reviewed of strongly beating the net against woody in preparation of a formal list of Michigan shrubs and lower tree limbs. The number of Coleoptera by W. G. Ruesink et al. Many of sweeps taken was not predetermined and these papers mention only one or a few spe- varied with each sampling date, depending cies, but others cover entire major families on conditions and productivity. The number for all of North America north of Mexico. of sweeps varied among habitats. For exam- Examples of the latter include Beal (2003) ple, in the hardwoods habitat only 40–50 [Dermestidae], Bousquet (2012) [Carabidae], sweeps would be taken because there was Bousquet et al. (2018) [Tenebrionidae], Her- little appropriate vegetation given that most man (2001) [Staphylinidae, in part], O’Brien tree branches were too high to reach with the and Wibmer (1982) [], Peck net and relatively little ground vegetation and Newton (2017) [], Pelletier grew in areas of deep shade. By contrast, as and Hébert (2014) [Cantharidae], and Ri- many as 500 sweeps were made in the grass- ley et al. (2003) [Chrysomelidae, excluding land site where there was about 1 ha of open Bruchinae]. In addition, the online SCAN herbaceous vegetation interspersed with a database (https://scan-bugs.org/ ) of over 100 few shrubs available for sweeping. The bee- North American arthropod collections was tles collected during sweeping were placed consulted to determine if they had Michigan specimens for any of the potential new state in labeled vials with fresh 70% ethanol and records, as well as photographs of identified stored until mounted for identification. specimens submitted to BugGuide (https:// Aquatic beetles. In addition to the bugguide.net/) and iNaturalist (https://www. beetles collected by the methods described inaturalist.org/) of beetles collected in Mich- above, the authors also examined vials of igan. Gary Parsons, Collection Manager of stored aquatic macroinvertebrates collect- the A. J. Cook Arthropod Research Collection ed by netting in three GRNA creeks (Cold (ARC) at Michigan State University (MSU), Creek, Finch Creek, and Shanty Creek; was also consulted to check if specimens of Fig. 1) during the spring and fall sampling any of the potential new state records had efforts during 2013-2019. These collections recently been deposited in the ARC-MSU were made by GRNA staff and volunteers as collection.

https://scholar.valpo.edu/tgle/vol53/iss2/9 4 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

142 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

3 2017 VI VII Months when collected in V VI VII VIII IX VII VI VII VIII IX V VI VII VIII IX V VI VII VIII IX VI VIII IX VI V VI VII VIII VIII IX VI VI VII VIII IX VI VI VII IX V V VI V VI VII VIII IX V VI VII VIII IX V VI VII VIII V VI VII VIII IX VIII IX V VI V VI VII VIII IX VI VI VII VIII IX VI VII

2

S F F F F F F P F F F P P FS FS FS FS FS FS FS FP FP FPS FPS FPS FPS FPS Methods by which collected

1

P P P C G H H PS CP CP CP HP HP HP GPS GPS CHP CHP HPO CHPS where CGHPO CGHPSO CGHPSO CGHPSO CGHPSO CGHPSO CGHPSO Habitats collected

1 1 2 1 1 4 13 No. exotic species collected

1 1 2 1 1 4 1 7 1 6 2 2 1 2 2 1 4 1 3 2 1 15 11 11 21 30 38 No. GRNA genera

1 1 3 1 1 5 1 1 7 2 2 1 2 2 1 7 1 3 2 1 20 17 18 21 41 40 47 No. GRNA species

48 88 77 57 84 61 45 82 11 85 56 52 in 229 151 788 473 958 243 481 145 117 965 435 258 No. 2439 1869 2919 North species America

Antlike flower beetles Common name Antlike leaf beetles Fungus Bostrichid beetles Straight-snouted beetles Jewel beetles Moss beetles Soldier beetles Ground beetles Loghorned beetles Leaf beetles Minute tree-fungus beetles Checkered beetles Lady beetles Minute hooded beetles Silken fungus beetles Weevils Skin beetles Click beetles Handsome fungus beetles Pleasing fungus beetles Plate-thigh beetles False click beetles Earth-boring scarab beetles Clown beetles Water scavenger beetles Lined flat bark beetles

nthicidae Table 1. Summary data by beetle family for Coleoptera collected at Grass River Natural Area (GRNA) in 2017, including common name of the family and number of species in North America north Mexico from Evans (2014), genera collected at GRNA, number of species collected and considered exotic to the United States, habitats where specimens were collected, methods by which beetles were collected, and months when collected. Family Aderidae A Anthribidae Bostrichidae Brentidae Buprestidae Cantharidae Carabidae Cerambycidae Chrysomelidae Ciidae Cleridae Curculionidae Dermestidae Elateridae Endomychidae Eucinetidae Eucnemidae Laemophloeidae

Published by ValpoScholar, 2020 5 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 143 V V VI VII VIII IX V VI VIII IX VII VIII IX VI VII VIII IX VII VI VII VIII V VI VII VIII IX VI VII VIII IX IX VI VII V VI VII VIII IX V VI VII VI VII VIII IX V VI VII VIII IX VI VI VII V VI VII VIII IX V VI VIII IX V VI VI V VI VII VIII IX V VI VII VIII IX VI VII VI V VI VII VIII IX V VI VII VIII IX VII VIII IX V VI VIII VI VII V VI

S S F F P F F F F F F F F F F F F FS FS FS FS FS FS FP FP FP FPS FPS FPS FPS FPS

P C O H H H CS CP CP CP PO HP HP HP HP HP CHP GPO CHP CHP CHP CHP CHPO GHSO CGHPS CGHPS CGHPO CGHPSO CGHPSO CGHPSO CGHPSO

4 5

1 6 3 2 1 2 4 1 3 1 3 1 4 1 3 1 1 3 1 5 2 3 2 1 2 2 2 2 8 14 20

1 7 5 3 1 3 4 1 4 1 9 1 6 1 4 1 1 3 1 8 3 4 2 1 2 3 2 2 20 43 13 r imarily the Rail Trail, P = pine, and S Sedge Meadow Trail. See text for details. f amily status, such as Cimberidinae being elevated to Cimberidae. ptember, X = October.

9 15 25 76 50 56 26 87 19 50 50 46 30 32 26 20 59 126 140 381 424 520 189 173 122 117 471 109 1700 4360 1184

Pine flower snout beetles Fireflies Minute brown scavenger beetles Round fungus beetles Stag beetles Net-winged beetles False darkling beetles Blister beetles Soft-winged flower beetles Root-eating beetles Tumbling flower beetles Hairy fungus beetles Sap beetles False blister beetles Shining flower beetles Featherwing beetles Ptilodactylid beetles Deathwatch & spider beetles Fire-colored beetles Scarab beetles Marsh beetles False flower beetles Carrion beetles Silvanid flat bark beetles Cryptic slime mold beetles Rove beetles Darkling beetles Polypore fungus beetles Throscid beetles Bark-gnawing beetles Zopherid beetles

4

Habitats: C = cedar, G grassland, H hardwoods, O other, but p Methods: F = funnel traps, P pitfall and S sweeping. Months: V = May, VI June, VII July, VIII August, and IX Se Some sources have elevated certain subfamilies of to Lampyridae Leiodidae Lucanidae Melandryidae Meloidae Monotomidae Mordellidae Mycetophagidae Nemonychidae Nitidulidae Oedemeridae Phalacridae Ptiliidae Pyrochroidae Scarabaeidae Silphidae Silvanidae Sphindidae Staphylinidae Tenebrionidae Tetratomidae Trogossitidae Zopheridae 1 2 3 4

https://scholar.valpo.edu/tgle/vol53/iss2/9 6 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

144 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

Status as native or exotic. As above, notes were recorded for each species as to whether they were native or exotic to the United States. The main sources of informa- tion were again Downie and Arnett (1996) and the internet site BugGuide. The native range of the exotic species was described as Asian, Eurasian, European, or Palearctic (northern portion of Eurasia). Diversity indices. Several diversity indices have been used in ecological studies to characterize community diversity, with some, such as the Shannon index, combining species richness (i.e., the total number of different species) and the proportion of each species into a single value (Washington 1984, Chao et al. 2014). Using EstimateS (Colwell 2013), we calculated the Shannon index (H) and the effective (or equivalent) number of species [ENS = exp(H)] values for the funnel trap data given that the total number of each beetle species collected was recorded from all traps. We computed separate values for each of the three main habitats (cedar, hard- woods, and pine) by combining all funnel trap data (i.e., from the three different lures) within each habitat, and also for each of the three lures (α-pinene, ethanol, or ipsenol) by combining the funnel trap data by lure type across the three habitats. The ENS value is calculated as the exponential of the Shan- non index, exp(H), and is equivalent to the number of equally common species required to produce the same diversity index value (Jost 2006). For example, Shannon index values of 2, 3, 4, and 5 would be equivalent to communities with 7, 20, 55, and 148 equally common species, respectively.

Results For all three collecting methods com- bined, 409 beetle species were identified from the 2017 GRNA samples, including 369 taxa identified to species, 24 to genus only, and 16 to subfamily only (Appendix 1). These 409 species represented 58 families, with the five most speciose being Curculionidae (47 species), Staphylinidae (43), Carabidae (41), Chrysomelidae (40), and Elateridae Figure 2. Venn diagrams indicating the number (21) (Table 1). By contrast, there were 20 of beetle species and corresponding percent of the families represented by only a single species, total for A: the 322 species that were collected in and another 22 families represented by only the cedar, hardwoods, and pine habitats (the label 2-5 species each (Table 1). There was a sig- of one specimen was lost so the numbers add to nificant positive linear correlation between 321); B: the 409 species that were collected in the number of species collected per beetle funnel traps, pitfall traps, and by sweep netting; family at GRNA and the corresponding total and C: the 210 species that were collected in fun- number of North American species (north of nel traps baited with either α-pinene, ethanol, or Mexico) recognized in those same families, ipsenol (values add to 209 due to one lost label). using the values presented in Table 1 and Evans (2014) (r = 0.91, R2 = 0.83, N = 58, P < 0.0001). The 40 taxa that were not identi- fied to species represented 16 beetle families, 22 of which were Staphylinidae, and 16 of

Published by ValpoScholar, 2020 7 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 145

Table 2. Number of families, genera, and species of beetles collected at Grass River Natural Area in 2017 in the cedar, hardwoods, and pine habitats by trapping method (funnel traps, pitfall traps, and sweep net- ting); see text for details. No. families, genera, species by trapping method Habitat Funnel traps Pitfall traps Sweep net Cedar 30, 63, 78 10, 23, 34 11, 18, 24 Hardwoods 34, 98, 116 11, 38, 57 10, 27, 38 Pine 43, 118, 140 14, 41, 53 19, 47, 53

the 22 staphylinids were only identified to Lures. Of the 210 species collected the subfamily (Appendix 1). in funnel traps, 144 species (119 genera in Habitats. Of the 409 species collected 39 families) were collected in funnel traps at GRNA, 322 were collected at the three baited with α-pinene, 123 species (97 genera main sampled habitats (cedar, hardwoods, in 36 families) in ethanol-baited traps, and and pine) using all three trapping methods, 114 species (97 genera in 38 families) in compared with 152 species being collected ipsenol-baited traps (Fig. 2c, Appendix 1). Of by sweeping along the two GRNA trails and these 210 species, 45 species were collected only with α-pinene, 25 species only with eth- the grassland site (Appendix 1). Of the 322 anol, and 23 species only with ipsenol (Fig. species, 105 species (80 genera in 32 fami- 2c). By contrast, 56 species were collected lies) were collected in the cedar habitat, 176 with all three lures (Fig. 2c). The label was species (131 genera in 38 families) in the lost on one specimen and thus the values in hardwoods habitat, and 199 species (158 Fig. 2c only add to 209, not 210. Considering genera in 47 families) in the pine habitat all collections made from the nine funnel (Appendix 1). Similarly, of these 322 spe- traps used in 2017, 707 individual beetles cies and the three main habitats sampled, were collected in the three funnel traps 27 species were collected only in the cedar baited with α-pinene lures, 513 individuals habitat, 76 only in the hardwoods habitat, in the three traps with ethanol lures, and and 99 only in the pine habitat, whereas 40 553 individuals in the three traps with species were collected in all three habitats ipsenol lures (Appendix 1). When combining (Fig. 2a). For the three GRNA locations the funnel trap data at each site, 79 beetle where only sweep netting occurred, 57 beetle species were collected at the cedar site, 116 species (47 genera in 18 families, including at the hardwoods site, and 143 at the pine 16 unique species) were collected along the site. Several beetle species showed a strong Sedge Meadow Trail, 69 species (56 genera preference for one of the three lures tested. in 18 families, including 18 unique species) For example, α-pinene was most attractive along the Rail Trail, and 83 species (67 to the curculionid Pissodes affinis Randall genera in 19 families, including 36 unique (capturing 96% of 23 individuals) and the species) in the grassland site (Appendix 1). histerid Paromalus bistriatus Erichson Trapping methods. Of the 409 (95% of 20), ethanol was most attractive to species collected in this study, 210 species the curculionid Anisandrus sayi Hopkins (165 genera in 51 families) were collected in (97% of 72) and the nitidulid funnel traps, 104 species (66 genera in 20 sanguinolentus (Olivier) (93% of 27), and families) in pitfall traps, and 168 species (124 ipsenol was most attractive to the ceram- genera in 26 families) with sweep nets (Fig. bycid Monochamus scutellatus (Say) (61% of 2b, Appendix 1). Similarly, the greatest num- 66) and the clerid Madoniella dislocata (Say) ber of species were collected in funnel traps (73% of 108) (Appendix 1). when considering the cedar, hardwoods, and Diversity indices. The Shannon pine habitats separately (Table 2). Of the index values H, based on funnel trap data 409 species, 145 species were collected only only, were 3.297 for the cedar habitat, 3.971 with funnel traps, 86 species only with pitfall for the hardwoods habitat, and 4.075 for the traps, and 108 species only with sweep nets pine habitat. The corresponding effective (Fig. 2b). By contrast, only three species were number of equally common species (ENS) collected with all three methods (Fig. 2b), for these Shannon index values are 27 for including the carabid gregarious cedar, 53 for hardwoods, and 59 for pine. (Say), the coccinellid Brachiacantha decem- Similarly, considering the three lures tested, pustulata (Melsheimer), and a ptilodactylid the Shannon Index values H were 4.314 for Ptilodactyla sp. (Appendix 1). the α-pinene, 4.005 for ethanol, and 3.764 for

https://scholar.valpo.edu/tgle/vol53/iss2/9 8 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

146 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

Figure 3. Percentage of beetle species collected by three sampling methods at Grass River Natural Area in 2017 by month of first collection and habitat for the beetle species collected in the cedar (105 species), hardwoods (176 species), and pine (199 species) habitats. Values above each column are percentage values within each habitat.

ipsenol, which correspond to ENS values of New Michigan state records. Of the 75, 55, and 43, respectively. 369 species that were identified to the species Seasonality. Of the 409 beetle species level, we found published, in print, collection collected at GRNA in 2017, 21% (85 species) records from Michigan for 336 species. For were first collected in May, 40% (162) in the remaining 33 species, 14 were listed June, 23% (95) in July, 10% (41) in August, online in SCAN as having been collected in and 6% (26) in September (Appendix 1). A Michigan and deposited in the Albert J. Cook broadly similar pattern emerged for each Arthropod Research Collection at Michigan of the three main habitats sampled, with State University (1 of the 14 was also depos- most species being first collected in May or ited in the C.A. Triplehorn Insect Collection June (Fig. 3). Moreover, when considering at The Ohio State University), 1 was listed on just those beetle species collected in funnel BugGuide as having been collected in Mich- traps, the resulting season-long species igan, and the remaining 18 were considered accumulation curves for each of the three new state records for Michigan (Appendix main habitats were similar in shape with 1). These 18 new state records are members more species consistently being collected at of 12 beetle families, including Brentidae (1 the pine site compared with the hardwoods species), Cantharidae (2), Chrysomelidae and cedar sites (Fig. 4). At the family level, (4), Cryptophagidae (1), Curculionidae (3), 48% (28 families) of the 58 beetle families Hydrophilidae (1), Laemophloeidae (1), recorded were first collected in May, 38% Melyridae (1), Silvanidae (1), Staphylinidae (22) in June, 9% (5) in July, 3% (2) in August, (1), Tenebrionidae (1), and Throscidae (1) and 2% (1) in September (Table 1). The two (Appendix 1). Four of these 18 species are beetle families first collected in August were exotic to the United States, including the Ciidae and Endomychidae, with members of brentid Perapion curtirostre (Germar), the Ptiliidae being first collected in September chrysomelids Chrysolina hyperici (Forster) (Table 1). Considering the number of beetle and Mantura chrysanthemi (Koch), and the species collected each month for all 409 spe- staphylinid Tasgius melanarius (Heer). cies, 21% (85 species) were collected in May, Status as native or exotic. Overall, 59% (243) in June, 46% (187) in July, 33% 32 of the beetle species collected at GRNA (133) in August, and 27% (111) in September were exotic to the United States (Appendix (Appendix 1). 1). These 32 species represent 9 beetle

Published by ValpoScholar, 2020 9 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 147

Figure 4. Cumulative number of species collected in funnel traps (all lures combined within sites) by month in three habitats (cedar, hardwoods, and pine) sampled during May to September 2017 at Grass River Natural Area.

families, including Brentidae (1 species), Muona (1999), who used large numbers of Buprestidae (1), Carabidae (1), Chrysomel­ pitfall traps and flight-intercept traps in idae (4), Coccinellidae (2), Curculionidae three separate studies at Oulanka National (13), Dermestidae (1), Scarabaeidae (4), and Park in Finland, where the beetle fauna is Staphylinidae (5). The natural range of these very well known, but reported collecting 32 exotics was Asia for 4 species, Europe for only 54-61% of the known terrestrial beetle 12, Eurasia for 5, and the Palearctic for 11 species present in each study. If these same (Appendix 1). ratios are applied to our GRNA data (409 Aquatic beetles. Examination of the species), then the total number of terrestrial invertebrates from the stored GRNA stream beetle species at GRNA would be estimated samples from 2013–2019 revealed 16 species at 670–757 species. Still we would postulate of adult aquatic beetles, representing 12 that there would be even more beetle species genera in 5 families (Table 3). Of these 16 present at GRNA than these estimates given species, 11 were identified to the species level that sampling intensity in Muona’s (1999) and 5 to genus only (Table 3). Overall, there studies involved 100–240 pitfall traps and were 4 species of , 1 Elmidae, 32–120 flight-intercept traps, depending 3 Gyrinidae, 2 Haliplidae, and 6 Hydro- on the study. Additionally, sampling over philidae. All taxa identified to species are successive years typically results in more considered native to the United States and species being discovered at any site. To none were new state records for Michigan. demonstrate this point, consider the 10-year Aquatic beetles were collected from each of study by Martikainen and Kaila (2004) in the three creeks sampled at GRNA, including Finland that focused on saproxylic beetles Cold Creek, Finch Creek, and Shanty Creek (those species dependent on dead or decaying (Table 3, Fig. 1). wood; Ulyshen and Šobotník 2018) at two forest sites. They reported that only 50% of all 258 saproxylic species collected over the Discussion entire 10-year study were collected every Although over 400 beetle species were single year, that many species showed great collected at GRNA in 2017, this number un- year-to-year variation in population size (as doubtedly represents only a fraction of the reflected in trap catch numbers), and that total number of beetle species present. Insect new species were collected in every year of diversity tends to increase with sampling the study. intensity in insect surveys (Samways et al. Our single-year trapping effort fo- 2010). As an example, consider the work of cused on just three habitats, using three

https://scholar.valpo.edu/tgle/vol53/iss2/9 10 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

148 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

Table 3. Species of aquatic beetles collected during 2013-2019 in three creeks that cross Grass River Natural Area, including Cold Creek, Finch Creek, and Shanty Creek (see text for details and Fig. 1 for creek locations). FAMILY Subfamily Species Creek name (number collected, date) DYTISCIDAE (Predaceous diving beetles) Agabinae Ilybiosoma seriatum (Say) Shanty (3, 16 V 2015), Shanty (1, 3 X 2015); sp. Finch (1, 28 IX 2019) Hydroporinae Liodessus crotchi Nilsson Finch (1, 27 IX 2014), Cold (1, 17 X 2017) Neoporus sp. Cold (2, 16 V 2015), (Shanty (1, 16 V 2015) ELMIDAE (Riffle beetles) Elminae Dubiraphia bivittata (LeConte) Shanty (1, 16 V 2015) GYRINIDAE (Whirligig beetles) Gyrininae Gyrinus maculiventris LeConte Cold (1, 27 IX 2014) Gyrinus lecontei Fall Finch (3, 28 IX 2019) Gyrinus sp. Cold (1, 6 V 2017), Shanty (2, 21 V 2014) HALIPLIDAE (Crawling water beetles) Haliplus canadensis (Wallis) Cold (1, 16 V 2015) Haliplus immaculicollis Harris Cold (1, 16 V 2015) HYDROPHILIDAE (Water scavenger beetles) lutescens (Stephens) Cold (2, 16 V 2015) Enochrus ochraceus (Melsheimer) Shanty (2, 16 V 2015) Hydrobius fuscipes (Linnaeus) Shanty (1, 16 V 2015) Hydrobius melaenus (Germar) Shanty (2, 3 X 2015) Paracymus sp. Cold (1, 12 X 2013) Tropisternus sp. Finch (6, 12 X 2013)

sampling methods and deploying only three Of the three collection methods used funnel traps and two pitfall traps per site. in the present study, baited funnel traps We recognize that many more beetle species caught the greatest number of beetle species, would have been collected at GRNA if we had whereas pitfall traps caught the fewest (Fig. sampled in multiple years, started sampling 2b). Multiple collection methods have been earlier and ended later in the year, sampled compared in many other studies. At forest- more habitats, deployed more traps per site, ed sites in Finland, Hyvärinen et al. (2006) used a greater variety of baits in the funnel found that two types of flight-intercept traps traps, sampled at different heights within caught a greater number of beetle species the canopy, and sampled over multiple than did pitfall traps. Similarly, in a forest- years. In addition, many more beetle species ed area of Arkansas, Skvarla and Dowling would have been collected if we had used (2017) compared 12 trapping methods for additional methods, such as flight-intercept collecting four groups of beetles (Carabidae, window-pane traps, light traps, Malaise Buprestidae, Cerambycidae, and Curculion- traps, and Berlese funnels. Nevertheless, oidea excluding Scolytinae) and found that our modest trapping effort did document the best trapping method varied by beetle about 10% of the beetle species known to family. For example, the greatest diversity occur in Michigan, given that Michigan has of buprestids was collected in Malaise traps around 4000 recognized beetle species (W. G. and unbaited green funnel traps; whereas Ruesink et al., unpublished data). Malaise traps, canopy SLAM traps, and

Published by ValpoScholar, 2020 11 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 149

funnel traps were best for cerambycids; Coleoptera. For example, in New Zealand, pitfall traps and funnel traps were best using Malaise traps, Hutcheson (1990) noted for non-scolytine curculionoids; and pitfall that both insect species richness and abun- traps were best for carabids. In the present dance increased from spring to summer and study, 33 (80%) of the 41 carabid species then declined into fall. In Arkansas, using collected at GRNA were collected in pitfall multiple trapping methods, Skvarla and traps, compared with only 4 carabid species Dowling (2017) noted different adult activity in funnel traps and 6 species in sweep-net patterns among different groups of beetles: samples (Appendix 1). Carabidae were active from spring to fall The three lures used in the present with a slight peak in June, Buprestidae and study are commonly used in surveys for Cerambycidae were most active in June and July, and Curculionoidea (excluding Scolyti- bark- and wood-infesting beetles, especially nae) were most active in May and June. In Cerambycidae and Scolytinae (Brockerhoff et the present study, we had similar results for al. 2006, Millar and Hanks 2017, Rabaglia members of these same beetle groups with et al. 2019, Rassati et al. 2019). In fact, most species initiating adult activity in June since 2007 the USDA Forest Service has (Appendix 1). implemented a nationwide survey for exotic bark and ambrosia beetles, using funnel Shannon index (H) values have been traps baited with α-pinene, ethanol and a reported in dozens of Coleoptera survey stud- three-component pheromone lure containing ies, but usually with a focus on just one or a ipsdienol, cis-verbenol, and methyl-butenol few beetle families. However, all Coleoptera (Rabaglia et al. 2019). As noted in the pres- families and species were included in the ent study, Miller et al. (2015) and Rabaglia analyses of a few studies and therefore can et al. (2019) also reported that many ceram- be compared to our results for the cedar (H bycid and scolytine species showed strong = 3.297; ENS = 27), hardwoods (H = 3.971, preferences for individual lures similar to ENS = 53), and pine sites (H = 4.075, ENS = the lures we tested, either individually or 59). For example, Coulson et al. (1971), work- in various combinations. ing in North Carolina, reported diversity values for the canopy Coleoptera collected in Volatile compounds such as α-pinene a monoculture stand of white pine (seasonal and ethanol are often produced at increased range of H = 0.969–2.205, ENS = 3–9) and a levels in stressed trees, and thus many bark- mostly oak-hickory (Quercus-Carya) forest and wood-infesting insects have evolved to (seasonal range of H = 1.931–2.941, ENS = use these compounds to locate stressed host 7–19). Trieff (2002), working at multiple sites plants (Mattson and Haack 1987, Millar in Tennessee, collected canopy Coleoptera by and Hanks 2017). Besides cerambycids and fogging northern red oak trees and reported scolytines, many other beetles use α-pinene a range in season-long H values of 3.04 to and ethanol, as well as bark beetle pher- 3.70 (ENS = 21–40) for the various sites. In a omones, as kairomones to locate stressed third study, working in a mixed-conifer forest plants (e.g., some Bostrichidae, Buprestidae, in California, Apigian et al. (2006) reported and Niti­dulidae) or potential prey (e.g., some an overall H value of 3.73 (ENS = 42) for all Cleridae, Histeridae, and Tenebrionidae) Coleoptera collected in pitfall traps collected (Chénier and Philogène 1989, Schroeder over a 3-year period. and Lindelöw 1989, Erbilgin and Raffa 2002, Bouget et al. 2009, Millar and Hanks 2017). Faunistic surveys provide basic knowl- edge on the occurrence, abundance and Variation in seasonal adult activity distribution of selected species within a among insects reflects their overwintering given area. To that end, the present study life stage, overwintering site, and voltinism provides information on over 400 beetle spe- pattern (Wolda 1988). In temperate areas, cies found at the Grass River Natural Area many insects overwinter as larvae (Danks in Antrim County, Michigan, and will serve 1978), and thus must still complete meta- as a baseline for future surveys of Coleoptera morphosis before adult activity can begin. in Michigan. Many others overwinter as adults and are thus most abundant in late spring or in late Acknowledgments summer and autumn. Adults of temperate insects tend to have one of three seasonal The authors thank Anthony Cognato patterns: a single peak, multiple peaks (for (Michigan State University) and Thomas multivoltine species), or one broad maximum McElrath (Illinois Natural History Survey) (Wolda 1988). Although we collected new for identification assistance on Scolytinae beetle species at GRNA in all months from and Latridiidae, respectively; John Epler May to September, June had the greatest (independent researcher, Crawfordville, number (162 of 409) of new species collect- Florida), Frank Etzler (Clemson Univer- ed for any single month. Similar seasonal sity), Margaret Thayer (Field Museum, patterns have been reported by others for Chicago), and Daniel Young (University of

https://scholar.valpo.edu/tgle/vol53/iss2/9 12 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

150 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

Wisconsin-Madison), for taxonomic advice Insect biodiversity: science and society. 2nd regarding several beetle families; Gary edition. John Wiley & Sons, Hoboken, NJ. Parsons (Michigan State University, MSU) Bouget, C., H. Brustel, A. Brin, and L. Val- for assistance with beetle collection records ladares. 2009. Evaluation of window flight at MSU and the SCAN database; and Rich- traps for effectiveness at monitoring dead ard Hannan (GRNA Board), Toby Petrice wood-associated beetles: the effect of ethanol (USDA Forest Service, Northern Research lure under contrasting environmental condi- Station), Michael Ulyshen (USDA Forest tions. Agricultural and Forest Entomology Service, Southern Research Station), and 11: 143–152. two anonymous reviewers for comments on an earlier draft of this paper. This research Bright, E. 2020. Aquatic insects of Michigan. was supported in part by the USDA Forest Available from: http://www.aquaticinsects. org/ (accessed 22 August 2020) Service. Brockerhoff, E. G., D. C. Jones, M. O. Kim- Literature Cited berley, D. M. Suckling, and T. Donald- son. 2006. Nationwide survey for invasive Adams, C. C. 1909. The Coleoptera of Isle Royale, wood-boring and bark beetles (Coleoptera) Lake Superior, and their relation to the North using traps baited with pheromones and American centers of dispersal, pp. 157-203. In kairomones. Forest Ecology and Management C. C. Adams (ed.), An ecological survey of Isle 228: 234–240. Royale, Lake Superior. Michigan Geological Chao, A., N. J. Gotelli, T. C. Hsieh, E. L. Sand- Survey Annual Report for 1908, Lansing, MI. er, K. H. Ma, R. K. Colwell, and A. M. Andrews, A. W. 1916. Results of the Mershon Ellison. 2014. Rarefaction and extrapolation expedition to the Charity Islands, Lake Hu- with Hill numbers: a framework for sampling ron: Coleoptera, pp. 65-108. In A. G. Ruthven and estimation in species diversity studies. (ed.), Miscellaneous papers on the zoology of Ecological Monographs 84: 45–67. Michigan. Michigan Geological and Biological Chénier, J. V. R. and B. J. R. Philogène. 1989. Publication 20. Field responses of certain forest Coleoptera to Andrews, A. W. 1921. The Coleoptera of the Shi- conifer monoterpenes and ethanol. Journal of ras expedition to Whitefish Point, Chippewa Chemical Ecology 15: 1729–1745. County, Michigan. Papers of the Michigan Cohen, J. G., M. A. Kost, B. S. Slaughter, Academy of Science, Arts, and Letters 1: and D. A. Albert. 2015. A field guide to the 293–390. natural communities of Michigan. Michigan Natural Features Inventory. Michigan State Apigian K. O., D. L. Dahlsten, and S. L. Ste- University Press, East Lansing, MI. 362 pp. phens. 2006. Biodiversity of Coleoptera and the importance of habitat structural features Cognato, A. I., N. Barc, M. Philip, R. Mech, in a Sierra Nevada mixed-conifer forest. En- A. D. Smith, E. Galbraith, A. J. Storer, vironmental Entomology 35: 964–975. and L. R. Kirkendall. 2009. The native and introduced bark and ambrosia beetles Arnett, R. H. Jr. and M. C. Thomas (eds.). of Michigan (Coleoptera: Curculionidae, 2001. American beetles, Volume 1. CRC Scolytinae). The Great Lakes Entomologist Press, Boca Raton, FL. 443 pp. 42: 101–120. Arnett, R. H. Jr., M. C. Thomas, P. E. Skelley, Colwell, R. K. 2013. EstimateS: Statistical and J. H. Frank (eds.). 2002. American estimation of species richness and shared beetles, Volume 2. CRC Press, Boca Raton, species from samples. Version 9. Available FL. 861 pp. from: http://purl.oclc.org/estimates (accessed Beal, R. S. Jr. 2003. Annotated checklist of Nearc- 22 August 2020) tic Dermestidae with revised key to genera. Coulson R. N., D. A. Crossley, Jr., and C. S. The Coleopterists Bulletin 57: 391–404. Gist. 1971. Patterns of Coleoptera species di- Bousquet, Y. 2012. Catalogue of Geadephaga versity in contrasting white pine and coppice (Coleoptera, ) of America, north of canopy communities. The American Midland Mexico. ZooKeys 245: 1–1722. Naturalist 86: 145–151. Bousquet, Y., D. B. Thomas, P. Bouchard, A. Danks, H. V. 1978. Modes of seasonal adaptations D. Smith, R. L. Aalbu, M. A. Johnston, in the insects. l. Winter survival. The Cana- and W. E. Steiner, Jr. 2018. Catalogue of dian Entomologist 110: 1167–1205. Tenebrionidae (Coleoptera) of North Ameri- Dillon, E. S. and L. S. Dillon. 1972. A manual ca. ZooKeys 728: 1–455. of common beetles of eastern North America. Bouchard, P., A. B. T. Smith, H. B. Douglas, M. Vol. 1. Dover Publications, New York, NY. L. Gimmel, A. J. Brunke, and K. Kanda. 434 pp. 2017. Biodiversity of Coleoptera, pp. 337– Downie, N. M. and R. H. Arnett, Jr. 1996. The 417. In: R. G. Foottit and P. H. Adler (eds.), beetles of northeastern North America. Vol. 1

Published by ValpoScholar, 2020 13 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 151

and 2. The Sandhill Crane Press, Gainesville, Hatch, M. H. 1925. A list of Coleoptera from, FL. 1721 pp. Michigan. Papers of the Michigan Academy of Science, Arts, and Letters 4: 543–586. Durkis, T. J. and R. M. Reeves. 1982. Barriers increase efficiency of pitfall traps. Entomo- Herman, L. H. 2001. Catalogue of the Staphylin- logical News 93: 8–12. idae (Insecta: Coleoptera). 1758 to the end of the second millennium. Parts I-VII. Bulletin Evans, A. V. 2014. Beetles of eastern North Amer- of the American Museum of Natural History ica. Princeton University Press, Princeton, 265: 1–4218. NJ. 560 pp. Hubbard, H. G. and E. A. Schwarz. 1878. The Erbilgin, N. and K. F Raffa. 2002. Association Coleoptera of Michigan. Proceedings of the of declining red pine stands with reduced American Philosophical Society 17: 593–669. populations of bark beetle predators, season- al increases in root colonizing insects, and Hutcheson, J. 1990. Characterization of terres- incidence of root pathogens. Forest Ecology trial insect communities using quantified, and Management 164: 221–236. Malaise-trapped Coleoptera. Ecological Entomology 15: 143–151. Foottit, R. G. and P. H. Adler (Eds). 2017. Insect biodiversity: science and society. 2nd Hyvärinen, E., J. Kouki, and P. Martikain- edition. John Wiley & Sons, Hoboken, NJ. en. 2006. A comparison of three trapping 904 pp. methods used to survey forest dwelling Co- leoptera. European Journal of Entomology Gosling, D. C. L. 1973. An annotated list of the 103: 397–407. Cerambycidae of Michigan (Coleoptera) part I, introduction and the subfamilies Parandri- Jost, L. 2006. Entropy and diversity. Oikos 113: nae, Prioninae, Spondylinae, Aseminae, and 363–375. Cerambycinae. The Great Lakes Entomolo- Kost, M. A., D. A. Albert, J. G. Cohen, B. S. gist 6: 65–84. Slaughter, R. K. Schillo, C. R. Weber, and Gosling, D. C. L. 1980. An annotated list of the K. A. Chapman. 2007. Natural communities of Michigan: classification and description. checkered beetles (Coleoptera: Cleridae) of Michigan Natural Features Inventory, Re- Michigan. The Great Lakes Entomologist port No. 2007-21, Lansing, MI. 317 pp. 13: 65–76. Marske, K. A. and M. A. Ivie. 2003. Beetle Gosling, D. C. L. 1983. New state records of Cer­ fauna of the United States and Canada. The ambycidae from Michigan (Coleoptera). The Coleopterists Bulletin 57: 495–503. Great Lakes Entomologist 16: 187. Martikainen, P. and I. Kaila. 2004. Sampling Gosling, D. C. L., and N. M. Gosling. 1977. An saproxylic beetles: lessons from a 10-year annotated list of the Cerambycidae of Mich- monitoring study. Biological Conservation igan (Coleoptera). Part II, the subfamilies 120: 171–181. and . The Great Lakes Entomologist 10: 1–37. Mattson, W. J. and R. A. Haack. 1987. The role of drought in outbreaks of plant–eating GRNA (Grass River Natural Area). 2020. On- insects. BioScience 37: 110–118. line at: https://www.grassriver.org/ (accessed 22 August 2020) Millar, J. G. and L. M. Hanks. 2017. Chemical ecology of cerambycids, pp. 161–196. In Q. Graham, E. E., R. F. Mitchell, P. F. Reagel, Wang (ed.), Cerambycidae of the world: biol- J. D. Barbour, J. G. Millar, and L. M ogy and pest management. CRC Press/Taylor Hanks. 2010. Treating panel traps with a & Francis, Boca Raton, FL. fluoropolymer enhances their efficiency in capturing cerambycid beetles. Journal of Miller, D. R., C. M. Crowe, K. J. Dodds, L. D. Economic Entomology 103: 641–647. Galligan, P. De Groot, E. R. Hoebeke, A. E. Mayfield, T. M. Poland, K. F. Raf- Grey, L. and A. I. Cognato. 2019. Ciidae of fa, and J. D. Sweeney. 2015. Ipsenol, Michigan (Insects: Coleoptera). The Great ipsdienol, ethanol, and α-pinene: trap lure Lakes Entomologist 52: 161–168. blend for Cerambyc­ idae and Buprestidae Hackett, R. A., P. Higman, and L. May. 2017. (Coleoptera) in pine forests of eastern North Natural community delineation and floristic America. Journal of Economic Entomology quality assessments of Grass River Natural 108:1837–1851. Area, Antrim County, Michigan. Michigan Muona, J. 1999. Trapping beetles in boreal coni­ Natural Features Inventory, Report No. ferous forest – how many species do we miss? 2017-12, Lansing, MI. 64 pp. Fennia 177: 11–16. Hansen, J. E. and T. R. New. 2005. Use of O’Brien, C. W. and G. J. Wibmer. 1982. An- barrier pitfall traps to enhance inventory notated checklist of the weevils (Curculion- surveys of epigaeic Coleoptera. Journal of idae sensu lato) of North America, Central Insect Conservation, 9: 131–136. America, and the West Indies (Coleoptera:

https://scholar.valpo.edu/tgle/vol53/iss2/9 14 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

152 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4

Curculionoidea). Memoirs of the American and Curculionoidea excluding Scolytinae). Entomological Institute 34: 1–382. Journal of Insect Science 17: 1–28. Peck, S. B. and A. F. Newton. 2017. An anno- Skvarla, M. J., J. L. Larson, and A. G. P. tated catalog of the Leiodidae (Coleoptera) Dowling. 2014. Pitfalls and preservatives: a of the Nearctic region (continental North review. Journal of the Entomological Society America north of Mexico). The Coleopterists of Ontario 145: 15–43. Bulletin 71: 211–258. Spilman, T. J. 1973. A List of the Tenebrionidae Pelletier, G. and C. Hébert. 2014. The Canthar­ of Michigan (Coleoptera). The Great Lakes idae of eastern Canada and northeastern Entomologist 6(3): 85–91. United States. Canadian Journal of Arthro- Trieff, D. D. 2002. Composition of the Coleoptera pod Identification 25: 1–246. and associated insects collected by canopy Petrice, T. R. and R. A. Haack 2015. Compar- fogging of northern red oak (Quercus rubra ison of different trap colors and types for L.) trees in the Great Smoky Mountains capturing adult (Coleoptera: Bupres- National Park and The University of Ten- tidae) and other buprestids. The Great Lakes nessee Arboretum. M. S. thesis, University Entomologist 48: 45–66. of Tennessee, Knoxville. 87 pp. Rabaglia, R. J., A. I. Cognato, E. R. Hoebeke, Townsend, T. 1889. Contribution to a list of the C. W. Johnson, J. R. Labonte, M. E. Car- Coleoptera of the Lower Peninsula of Michi- ter, and J. J. Vlache. 2019. Early detection gan. Psyche 5: 231–325. and rapid response: a ten-year summary of Ulyshen, M. D.and J. Šobotník. 2018. An the USDA Forest Service program of sur- introduction to the diversity, ecology and veillance for non-native bark and ambrosia conservation of saproxylic insects, pp. 1–47. beetles. American Entomologist 65: 29–42. In: M. D. Ulyshen, (ed.), Saproxylic insects: Rassati, D., L. Marini, M. Marchioro, P. diversity, ecology and conservation. Springer, Rapuzzi, G. Magnani, R. Poloni, F. Di Heidelberg. Giovanni, P. Mayo, and J. Sweeney. 2019. van Klink, R., D. E. Bowler, K. B. Gongalsky, Developing trapping protocols for wood-bor- A. B. Swengel, A. Genyile, and J. M. ing beetles associated with broadleaf trees. Chase. 2020. Meta-analysis reveals declines Journal of Pest Science 92: 267–279. in terrestrial but increases in freshwater in- Riley, E. G., S. M. Clark and T. N. Seeno. sect abundance. Science 368 (6489): 417–420. 2003. Catalog of the leaf beetles of America Washington, H. G. 1984. Diversity, biotic and north of Mexico. Special Publication No. 1. similarity indices: a review with special rele- The Coleopterists Society, North Potomac, vance to aquatic ecosystems. Water Research MD. 290 pp. 18: 653–694. Samways, M. J. 2019. Insect conservation: a Wellso, S. G., G. V. Manley, and J. A. Jackman. global synthesis. CABI, Wallingford, Oxon, 1976. Key and notes on the Buprestidae UK. 560 pp. (Coleoptera) of Michigan. The Great Lakes Samways, M. J, M. A. McGeoch, and T. R. Entomologist 9: 1–22. New. 2010. Insect conservation: a handbook White R. E. 1983. A field guide to the beetles of of approaches and methods. Oxford Univer- North America. Houghton Mifflin, Boston, sity Press, Oxford. 441 pp. MA. Schroeder, L. M. and A. Lindelöw. 1989. At- Wolcott, A. B. 1909. Supplementary list of Isle traction of scolytids and associated beetles by Royale beetles, pp. 204-215 in C. C. Adams different absolute amounts and proportions (ed.), An ecological survey of Isle Royale, of a-pinene and ethanol. Journal of Chemical Lake Superior. Michigan Geological Survey Ecology 15: 807–817. Annual Report for 1908, Lansing, MI. Schwarz, E. A. 1876. List of Coleoptera collected Wolda, H. 1988. Insect seasonality: Why? Annual in Michigan in 1874. Psyche 1: 145–148. Review of Ecology and Systematics 19: 1–18. Skvarla, M. J. and A. P. G. Dowling. 2017. A Zhang, Z.-Q. 2011. biodiversity: an in- comparison of trapping techniques (Coleop- troduction to higher-level classification and tera: Carabidae, Buprestidae, Cerambycidae, taxonomic richness. Zootaxa 3148: 7–12.

Published by ValpoScholar, 2020 15 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 153 6 Pal Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Eur USA Native to

0 0 0 0 0 0 0 5 2 0 1 4 12 Total (Continued on next page) 5

0 0 0 0 0 0 0 0 0 0 0 1 0 Ips

0 0 0 0 0 0 0 3 0 0 0 4 1 ETOH Funnel trap collections

0 0 0 0 0 0 0 2 2 0 1 7 3 a-p

4

S S S S S S S F F P F F F Methods

3

V VI VI VII VII VII V VI VI IX VI VII VI VII Months VI VII IX collected VII VIII IX V VI VII VIII

2

P P P G G G G H GS GS HP HP GPS Habitats

(LeConte)

(Say)

(Boheman) (Germar) * (Olivier)

(Say) (Ménétriés) (Randall) (Say) (Say) Casey (Say) (LeConte)

1

family Species

Elonus basalis Notoxus desertus Euparius marmoreus Trigonorhinus rotundatus Trigonorhinus sticticus Prostephanus punctatus Perapion curtirostre Agrilus cuprescens Agrilus politus Taphrocerus gracilis Anthaxia inornata divaricata alternatus Sub Notoxinae Anthribinae Bostrichinae Apioninae Agrilinae Buprestinae Chrysochroinae Byrrhinae Appendix 1. Coleoptera collected at Grass River Natural Area in 2017, using funnel traps (F) and pitfall (P), by sweeping (S), with details on habitats where specimens were collected, months when and numbers of individuals collected in funnel traps by bait type. Additional information is presented on whether each species considered a potential new state record for Michigan (species with an * after the species name), and if each is considered native or exotic to continental USA. FAMILY ADERIDAE ANTHICIDAE ANTHRIBIDAE BOSTRICHIDAE BRENTIDAE BUPRESTIDAE BYRRHIDAE

https://scholar.valpo.edu/tgle/vol53/iss2/9 16 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

154 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes USA Native

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 0 0 0 0 2 0 0 0 7 5 34 Total 5

0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 1 Ips

0 0 0 0 0 0 0 0 0 0 0 8 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 2 1 ETOH Funnel trap collections

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 4 3 25 a-p

4

S S S S S S S S S S S S S S S S F P P P P P P P P F FS FS FS Methods

3

V IX IX VI VI VI VI VI VI VI VI VI VII VII VII VII VII VII VII VIII V VI V VIII V VIII VI VII VIII IX VI VIII VII VIII Months collected VI VII VIII VI VII VIII IX

2

S S S S S P P P G G G O G G H H H H CP SO HP CH GO HO GPS CHP CHP HSO GHS Habitats

Dejean

(DeGeer)

(Green) *

(Green) * (Eschscholtz)

(Say) (Say)

(LeConte) (LeConte)

(Say) (Fall)

Brown (LeConte) (Say) Lindroth Fall (LeConte) LeConte (Say) (Say) (Mannerheim) (Fabricius) (Melsheimer) (Say) (LeConte) LeConte (Casey) Casey Dejean

1

family Species

Atalantycha bilineata Atalantycha neglecta tuberculata Dichelotarsus piniphilus brevicollis Podabrus diadema Podabrus frater Podabrus rugosulus Podabrus tricostatus fraxini Rhagonycha imbecillis Rhagonycha nanula Rhagonycha oriflava Rhagonycha recta Rhagonycha scitula Rhagonycha vilis Chauliognathus pensylvanicus Polemius canadensis goryi stenostomus lecontei Agonum fidele Agonum gratiosum angustata Amara ellipsis interstitialis Anisodactylus agricola Anisodactylus kirbyi Rhagonycha hirticula Rhagonycha septentrionis Rhagonycha Sub Chauliognathinae Silinae Carabinae Appendix 1. Continued. FAMILY CANTHARIDAE CARABIDAE

Published by ValpoScholar, 2020 17 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 155 NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Eur

0 0 0 0 0 0 1 0 0 0 3 0 0 0 0 3 0 0 0 0 0 0 9 0 0 0 1 0 0 0 0 0 1 9 0 9 (Continued on next page)

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 2 0 4

0 0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 1 0 0 0 0 0 0 4 0 3

0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 1 0 0 0 0 0 0 6 0 0 0 0 0 0 0 0 0 1 3 0 2

S S S S P P P F P P P F P P P P P P P P P P F P P P F P P P P F P F FS FPS

V V IX IX IX VI VI VI VI VI VI VI VI VII VII VII VII VII VIII VIII V VI VI VII VI VII VI VII VI VII VIII IX VI VIII VII VIII VII VIII VI VII IX VI VIII IX VII VIII IX VII VIII IX V VI VII VIII VI VII VIII IX VI VII VIII IX

P P P P P P P P C C O G H H H H H H H H H H H H CP PO HP HP CG HP GO CH CHP CHP CHP CHP

(Motschulsky)

(Motschulsky)

(Illiger) Notman

(Say) Eschscholtz Dejean (LeConte) (Say) (Newman) (LeConte) (Dejean) (Germar) (Fabricius)

(Dejean) (Say)

(Say) Dejean Dejean (Say) (Say) Casey (Say) (Say) (Dejean) Hayward Dejean LeConte (Say)

Say Dejean Motschulsky Dejean

Dejean Say sp.

leida punctata Calathus gregarius Cal Cyclotrachelus sodalis Cymindis americana Cymindis limbata Cymindis platicollis Dromius piceus Harpalus erythropus Harpalus providens Harpalus somnulentus Lebia fuscata Lebia viridis Myas cyanescens Platynus decentis Poecilus lucublandus Pterostichus adstrictus Pterostichus commutabilis Pterostichus coracinus Pterostichus corvinus Pterostichus luctuosus Pterostichus melanarius Pterostichus mutus Pterostichus stygicus Pterostichus tristis Selenophorus opalinus Synuchus impunctatus Bembidion affine Bembidion fortestriatum Bembidion mimus Bembidion semicinctum Bembidion Trechus apicalis Neoclytus acuminatus Xylotrechus sagittatus Xylotrechus undulatus Astylopsis sexguttata Cerambycinae Lamiinae

CERAMBYCIDAE `

https://scholar.valpo.edu/tgle/vol53/iss2/9 18 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

156 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 Pal Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes USA EurAs Native

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6 0 0 1 0 1 0 0 0 1 5 4 66 Total 5

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 3 40 Ips

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 6 0 0 0 0 0 0 0 0 0 0 5 1 ETOH Funnel trap collections

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 1 4 0 21 a-p

4

S S S S S S S S S S S S S S S S S S S S S S F F F F FS FS FS Methods

3

V V IX VI VI VI VI VI VI VII VII VII VII VII VIII VIII VIII VIII V VI VI VII VI VIII VI VIII VI VIII VI VIII VII VIII VII VIII Months V VI VII collected VII VIII IX VI VII VIII IX

2

S S P P P G O O G O G G O O G G G G H GS GS PO PO GO GO GPS CPO CHP CHP Habitats

#

Newman Kirby (Say) Casey

(Newman) (Forster)

Fabricius (Say)

(Germar) Bechyne (Schaeffer) (Kirby) (Fabricius) (Webar) (Newman) Haldeman Schoenherr (Fabricius)

(Olivier) (Forster) Fåhraeus (Swederus) (Say) Pierce Brown (Linnaeus) (Crotch) (Fabricius) Say

1

family Species

Monochamus scutellatus Oberea tripunctata Pogonocherus mixtus Saperda mutica Tetraopes melanurus Tetraopes tetrophthalmus Brachyleptura champlaini Grammoptera haematites Strangalepta abbreviata Typocerus velutinus Tetropium cinnamopterum brachialis Chalepus walshii Microrhopala vittata Sumitrosis inaequalis Calligrapha multipunctate Chrysolina hyperici Chrysomela knabi Chrysomela mainensis Gonioctena americana Labidomera clivicollis Bassareus mammifer Cryptocephalus quadruplex Cryptocephalus venustus Diachus auratus Exema canadensis Lexiphanes saponatus Sub Lepturinae Bruchinae Cassidinae Chrysomelinae Appendix 1. Continued. FAMILY CHRYSOMELIDAE

Published by ValpoScholar, 2020 19 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 157 Pal Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Eur

0 0 0 0 0 0 0 0 0 0 1 0 1 3 0 0 9 1 0 0 9 0 0 6 8 2 50 29 69 108 (Continued on next page)

0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 3 4 0 0 0 0 1 20 58 79

0 0 0 0 0 0 0 0 0 0 0 0 1 2 0 0 2 0 0 0 9 1 0 0 0 1 0 5 10 22

0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 7 1 0 0 4 0 0 6 7 1 6 7 20 17

S S S S S S S S S S S S S S S S F F F F F F F FS FS FS PS FS FS FS

V IX IX VI VI VI VI VI VII VII VIII V VI VI IX VI VII VI VII VIII XI VIII IX VIII IX VIII IX VII VIII V VI VII VI VII IX VI VIII IX VII VIII IX VII VIII IX VII VIII IX V VI VII IX V VI VII VIII VI VII VIII IX V VI VII VIII IX

S S S P P P G G G G G G G CP SO HP CH HO HO GPS GPS GSO CHP GPO CHP GPO CHP CHO GHO GHO

Barber

(Spinola)

(LeConte) *

(Brown) (Koch) *

(Say)

(Melsheimer)

(Say) Parry Horn * (Say)

Schaeffer ‡ (Say) Melsheimer (Say) (Brown) (Say) (Fabricius) (Melsheimer) (Horn) Blake (LeConte)

(Say) (Say) Blatchley * (Melsheimer) Wilcox Balsbaugh sp. Mellié

Neochlamisus comptoniae Neochlamisus eubati Pachybrachis othonus Pachybrachis trinotatus Donacia tuberculifrons Plateumaris rufa Paria fragariae Paria pratensis Paria thoracica Xanthonia decemnotata Xanthonia villosula Altica gloriosa Capraita subvittata Crepidodera longula Crepidodera nana Crepidodera populivora Crepidodera violacea Diabrotica undecimpunctata howardi Luperaltica nigripalpis Mantura chrysanthemi Neogalerucella Ophraella conferta Tricholochmaea decora Trirhabda borealis Cis fuscipes nigrifrons Enoclerus nigripes rufiventris dubius Madoniella dislocata Phyllobaenus pallipennis Donaciinae Eumolpinae Ciinae Clerinae Epiphloeinae Hydnocerinae CIIDAE CLERIDAE

https://scholar.valpo.edu/tgle/vol53/iss2/9 20 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

158 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 NA NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes USA Asian EurAs Native

3 0 0 0 0 0 9 1 0 0 0 0 4 1 1 1 1 0 1 1 1 2 19 21 Total 5

1 0 0 0 0 0 5 0 3 0 0 0 0 0 1 0 0 0 0 0 4 0 0 0 Ips

0 0 0 0 0 0 0 1 1 0 0 0 0 2 0 0 0 0 0 0 2 1 1 1 ETOH Funnel trap collections

2 0 0 0 0 0 4 0 0 0 0 0 2 0 1 1 1 0 1 0 0 1 15 15 a-p

4

S S S S S S S S S S F F F F F F F FS FS FS FS FS FS FPS

Methods

3

V V V VI VI VI VII VII VIII VIII VIII VIII VI IX VI VIII VII VIII Months VI VIII IX collected VII VIII IX VI VII VIII VI VII VIII V VI VII VIII VI VII VIII IX VI VII VIII IX V VI VII VIII IX V VI VII VIII IX

2

S S P P P C C O G G G O G H PS GS PO GP HP CH GPS GPO GPO CGPSO Habitats

Mulsant (Melsheimer) Melsheimer

(Say)

Linnaeus

(Say)

(Say) (Park) *

(Say) Casey (Say) Mulsant (Gyllenhal) (Say) (Say) (LeConte) (Pallas) (Say) (Say)

(Say) (Say)

(Say)

1 sp.

sp.

family Species

Zenodosus sanguineus Cymatodera bicolor Chilocorus stigma labiculata Anatis mali Anisosticta bitriangularis monticola Coccinella septempunctata Coccinella trifasciata perplexa Cycloneda munda Harmonia axyridis Hippodamia parenthesis Psyllobora vigintimaculata B Brachiacantha quadripunctata Hyperaspis binotata Hyperaspis proba Hyperaspis undulata Scymnus rubricaudus Scymnus Clypastraea lunata Sericoderus lateralis Atomaria Henoticus mycetoecus Henoticus Sub Thaneroclerinae Tillinae Chilocorinae Corylophinae Cryptophaginae Appendix 1. Continued. FAMILY COCCINELLIDAE rachiacantha decempustulata CORYLOPHIDAE CRYPTOPHAGIDAE

Published by ValpoScholar, 2020 21 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 159 Pal Pal Pal Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Eur Eur Eur Eur Eur Eur Eur EurAs

0 0 0 0 0 0 0 0 0 7 0 1 2 0 1 1 1 0 1 9 2 0 0 1 1 0 0 1 0 11 (Continued on next page)

0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 1 2 0 0 0 1 0 0 0 0 0

0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 1 0 0 1 1 5 1 0 0 0 0 0 0 0 0

0 0 0 0 0 0 0 0 0 5 0 1 0 0 1 0 1 0 0 7 4 1 0 0 0 1 0 0 1 0

S S S S S S S S S P F F P F F P F F P P F PS FS FS FS FS PS PS FP FP

V V VI VI VI VI VI VI VI VI VI VI VII VII VII VIII VIII VIII V VI VI IX VI VII VI VII VI VII VIII IX VI VIII VII VIII V VI VII VII VIII IX VII VIII IX V VI VII VIII IX

P P P P C C C G O O G G G H H H GP HP HP GO GO HO HO GPS CHP CHP GHP HPO GHSO GHPO

(Horn) *

(Goeze)

Horn *

Bedel (Casey) LeConte

Boheman (Fabricius) (Herbst) Say (Boheman) (Fabricius) (Bonsdorff) (Mayer) (Say) (Linnaeus) Casey * (LeConte) (Fåhraeus) (Linnaeus) Buchanan (Fabricius) Dalla Torre et al. (Linnaeus) LeConte LeConte (Fabricius) (Bonsdorff) (Fabricius)

Dirabius rectirostris Odontocorynus salebrosus Odontocorynus umbellae Stethobaris ovata (LeConte) Rhinoncus castor Acoptus suturalis Anthonomus subfasciatus Orchestes pallicornis Rhinusa tetra Tachyerges ephippiatus Tachyerges salicis Tychius picirostris Dryophthorus americanus pellucidus Mesagroicus minor Otiorhynchus ovatus Otiorhynchus rugosostriatus Otiorhynchus sulcatus Phyllobius oblongus Polydrusus formosus Sciaphilus asperatus Sitona cylindricollis Sitona lineellus Magdalis gentilis nenuphar Conotrachelus posticatus Hylobius congener Sibariops sectator Sibariops protractus Trichacorynus scoparius Sphenophorus Baridinae Ceutorhynchinae Conoderinae Cossoninae Curculioninae Dryophthorinae Entiminae Mesoptiliinae CURCULIONIDAE

https://scholar.valpo.edu/tgle/vol53/iss2/9 22 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

160 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Eur USA Asian EurAs Native

4 2 7 8 2 2 0 0 1 2 3 1 9 8 6 4 5 18 72 57 17 17 44 14 25 20 23 17 22

Total 5

3 3 1 0 0 4 1 1 0 0 0 8 1 2 0 1 0 6 4 2 0 6 0 0 2 0 0 1

12 Ips

1 2 5 1 0 1 0 0 2 0 0 0 0 2 3 1 4 1 5 1 2 0 6 11 70 14 17 41 11 ETOH Funnel trap collections

4 0 1 0 2 3 1 0 0 0 0 7 0 0 3 2 1 6 2 5 3 1 2 5 31 21 22 14 15 a-p

4

S S F F F F F F F F F F F F F F F F F F F F F FS FS FS FP FP FP

Methods

3

VI VI VI VII V VI V VI V VI V VI V VI V VI V VII V VII VI VII

VIII IX VII VIII VII VIII Months V VI VII V VI VII V VI VII V VI VII V VI VII VI VII IX collected VI VII VIII V VI VII IX V VI VII IX V VI VII VIII VI VII VIII IX V VI VII VIII IX V VI VII VIII IX

2

P P P P P O H CP CP GP PO HP HP HP HP HP HP CH CH CHP CHP CHP CHP CHP CHP CHP CHP CHP CHPO

Habitats

(Say)

(Fitch) (Herbst)

(Zimmermann)

(Say)

LeConte

(LeConte)

(Ratzeburg) (Say) (Blandford) Hopkins

(Harris) (Eichhoff) Erichson (Say) (Ratzeburg) (Say) Swaine (LeConte) (Linnaeus) (Say) (LeConte) (Kirby) (Fitch) (Say) Hopkins (Eichhoff) Randall

1

(Say)

family Species

Pissodes affinis Pissodes approximatus Rhyssomatus lineaticollis Anisandrus obesus Anisandrus sayi Corthylus punctatissimus Crypturgus borealis Dryocoetes autographus Gnathotrichus materiarius Ips grandicollis Ips pini Lymantor decipiens Monarthrum mali Orthotomicus caelatus Phloeotribus liminaris Xyleborinus saxeseni Xylosandrus germanus Anthrenus verbasci Cardiophorus convexulus Cardiophorus convexus Cardiophorus gagates acanthus Athous brightwelli Elathous discalceatus Gambrinus plebejus Hemicrepidius memnonius Limonius basilaris Pseudanostirus hieroglyphicus Pseudanostirus propolus Sub Scolytinae Megatominae Cardiophorinae Appendix 1. Continued. FAMILY DERMESTIDAE ELATERIDAE

Published by ValpoScholar, 2020 23 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 161 NA NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes

0 0 0 3 1 2 0 0 1 6 3 1 5 4 4 3 3 5 5 23 48 16 19 (Continued on next page)

0 0 0 2 0 2 0 0 0 1 0 0 1 2 2 3 1 3 0 2 6 10 22

0 0 0 0 1 0 0 0 0 2 1 0 3 2 6 1 0 2 6 1 2 10 10

0 0 0 1 0 0 0 0 1 3 3 2 1 1 0 1 0 0 7 4 1 3 20

S S S F F P P F F F F F F F F F F FS FS FS FS FS FS

V IX VI VI VI VI VI VI VI VI V VI VI VII VI VII VIII IX V VI VII V VI VII V VI VII VII VIII IX VII VIII IX VII VIII IX VI VII VIII V VI VII VIII V VI VII VIII

S S P P P C G H H H H H PO HP HP HP CH CHS CHP CHP CHPS HPSO CHPO

(Randall)

(LeConte)

LeConte (Say)

Otto

LeConte ‡ LeConte Newman ‡ (Brown) (Paykull) Otto, Muona & McClarin -complex (Newman)

LeConte Van Zwaluwenberg Germar (Herbst) Brown Say Brown

Casey

sp. sp.

Pseudanostirus triundulatus Ampedus mixtus Ampedus pedalis Dalopius vagus Dalopius vernus Elater abruptus Melanotus castanipes Melanotus communis Melanotus hyslopi Melanotus Phymaphora pulchella Lycoperdina ferruginea Acropteroxys gracilis Triplax frosti Eucinetus strigosus Onichodon canadensis Dirrhagofarsus ernae Entomophthalmus rufiolus Microrhagus carinicollis Microrhagus pectinatus Microrhagus subsinuatus Microrhagus triangularis Leiestinae Lycoperdininae Languriinae Erotylinae Macraulacinae Melasinae Bolboceratinae ENDOMYCHIDAE EROTYLIDAE EUCINETIDAE EUCNEMIDAE GEOTRUPIDAE

https://scholar.valpo.edu/tgle/vol53/iss2/9 24 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

162 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 NA NA NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes USA Native

0 0 0 1 1 0 1 0 0 0 2 4 0 1 53 10 20 10 54 Total 5

0 0 0 0 7 1 1 0 1 0 0 0 0 2 1 0 1 30 20 Ips

0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 0 0 10 15 ETOH Funnel trap collections

0 0 0 1 3 0 0 0 0 0 0 2 5 2 0 0 13 19 19 a-p

4

S S S F F P F P P F P F F FS FS FS PS FS FS Methods

3

V V IX VI VI VI VI VI VI VII VII VIII VI VII VIII IX Months collected VI VII VIII V VI VIII IX V VI VIII IX V VI VII VIII VI VII VIII IX

2

S P P P C C G H H CP CH GPS HPS CHP HSO HSO CGHSO CGHPO CGHPSO Habitats

(Comolli)

(Mannerheim)

Casey * (LeConte) (Say)

(Say) (LeConte) Erichson

(LeConte) * (Paykull) (LeConte) (Harris) (Linnaeus) (LeConte) Say

(Olivier)

sp. 1 sp.

sp.

family Species

Paromalus bistriatus Atholus sedecimstriatus Euspilotus assimilis Helocombus bifidus Ellychnia corrusca Lucidota atra Photinus indictus Photinus Pyractomena angulata Pyropyga decipiens Photuris americana Melanophthalma distinguenda Melanophthalma pumila Stephostethus liratus basilaris Chaetarthria atra Chaetarthria fervidus Laemophloeus Sub Dendrophilinae Histerinae Saprininae Hydrophilinae Lampyrinae Photurinae Corticariinae Latridiinae Appendix 1. Continued. FAMILY HISTERIDAE HYDROPHILIDAE LAEMOPHLOEIDAE LAMPYRIDAE LATRIDIIDAE LEIODIDAE

Published by ValpoScholar, 2020 25 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 163 NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes

0 8 2 0 0 2 2 0 1 0 2 2 3 0 1 0 1 2 3 15 (Continued on next page)

0 1 1 0 0 2 1 0 0 0 1 0 2 4 0 0 0 0 1 3

0 3 1 0 0 0 0 0 1 0 1 0 1 2 0 0 0 1 1 0

0 4 0 0 0 0 1 0 0 0 0 2 0 9 0 1 0 0 0 0

S S S S S S F F F F F F F P F F F FS FS FP

IX VI VI VI VII VII VII VII VII VIII VI IX VII IX VI VII VI VII VI VII VI VIII VII VIII VI VIII IX VI VIII IX VI VII VIII

P P P P P P P O O O H CP GP PO HP HP HP CH GPO CGPSO

(Melsheimer) (Fabricius)

Hentz

Haldeman (Say) (Fabricius) LeConte Kirby (Say) Melsheimer (Say) Melsheimer Germar (LeConte) (Weber) LeConte Melsheimer (LeConte) Marshall *

sp.

Leiodes assimilis Leiodes punctostriata Ceruchus piceus Calochromus perfacetus Calopteron reticulatum Calopteron terminale Dircaea liturata Orchesia castanea Rushia longula Serropalpus substriatus Nemognatha nemorensis Attalus terminalis Collops quadrimaculatus Hypebaeus Rhizophagus remotus Mordella atrata Mordella marginata Mordella melaena Mordellistena fuscipennis Attalus frosti Attalus Leiodinae Calochrominae Melandryinae Nemognathinae Rhizophaginae Mordellinae LUCANIDAE LYCIDAE MELANDRYIDAE MELOIDAE MELYRIDAE MONOTOMIDAE MORDELLIDAE

https://scholar.valpo.edu/tgle/vol53/iss2/9 26 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

164 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes USA Native

0 0 7 1 1 0 2 6 1 0 3 3 0 0 0 0 27 27 Total 5

0 0 7 6 0 1 0 1 1 1 0 0 1 0 0 0 0 0 Ips

0 0 3 0 1 0 0 1 2 0 0 1 1 0 0 0 0 25 ETOH Funnel trap collections

0 0 1 0 0 0 0 3 0 0 2 1 2 0 0 0 0 17 a-p

4

S S S S S F F F F F F P P P FS FS FP FP

Methods

3

V V V IX IX VII VII VII VII VII VIII V VI VI VII Months collected VII VIII IX V VI VIII IX VI VII VIII IX VI VII VIII IX V VI VII VIII IX

2

S P P P C G H H CS CP CP CP SO CHP CHP CHP CHP CGHPS Habitats

(Olivier)

(Say) (Say) W. J. Brown ‡ (Say)

LeConte Liljeblad Murray LeConte LeConte (Say) (LeConte) Liljeblad (LeConte) Melsheimer (Say) Casey

7

sp. 1

family Species

Mordellistena pulchra Mordellistena tosta Mordellistena trifasciata Mordellistena vera Mordellochroa scapularis Litargus tetraspilotus Cimberis elongata brachypterus Glischrochilus sanguinolentus Glischrochilus siepmanni Stelidota geminata Thalycra concolor Asclera ruficollis Acylomus ergoti Olibrus semistriatus Phalacrus politus Phalacrus Sub Mycetophaginae Cimberidinae Cryptarchinae Nitidulinae Oedemerinae Eustilbinae Olibrinae Phalacrinae Appendix 1. Continued. FAMILY MYCETOPHAGIDAE NEMONYCHIDAE NITIDULIDAE OEDEMERIDAE PHALACRIDAE

Published by ValpoScholar, 2020 27 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 165 NA NA NA NA Pal Pal Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Asian Asian

6 0 1 1 0 1 2 0 0 1 3 0 2 2 1 2 5 4 1 1 3 4 11 66 (Continued on next page)

0 0 0 0 0 0 0 0 0 0 2 7 0 1 1 1 1 2 3 1 1 2 0 16

0 0 0 0 0 0 1 0 0 0 1 0 0 0 1 0 0 1 1 0 0 1 2 18

6 0 1 1 0 1 1 0 0 1 0 4 0 1 0 0 1 2 0 0 0 0 2 32

S S F F F P P F F F P F F F F F F F F F FS FS FP FPS

V V IX IX VI VI VI VI VI VII VII VII VIII V VI V VI V VI VI IX VII IX VI VII VI VII V VI VII V VI VII VII VIII IX VII VIII IX

P P P P P P P P C C C O H H H H H CS HP HP HP GO GPO CHP

(Fabricius)

‡ (Fabricius)

(Say) (Froelich) (Linnaeus) (LeConte) (Schönherr)

Burmeister (Say) (LeConte) (Say) (Say) (Say) (Melsheimer)

Say Kirby Newman Brown 1 2

(Drury)

sp.

sp.

spp.

Ptiliola Ptilodactyla notatus sericeum Sculptotheca puberula Neopyrochroa flabellata Aegialia criddlei Colobopterus erraticus Dialytes striatulus Dialytes truncatus Pseudagolius bicolor Dichelonyx albicollis Dichelonyx backii Dichelonyx elongatula Diplotaxis sordida Hoplia trifasciata Macrodactylus subspinosus Phyllophaga crenulata Phyllophaga nitida Phyllophaga sp. Phyllophaga sp. Serica Popillia japonica Copris minutus Ptiliinae Ptilodactylinae Anobiinae Pyrochroinae Aphodiinae Melolonthinae Rutelinae Scarabaeinae PTILIIDAE PTILODACTYLIDAE PTINIDAE PYROCHROIDAE SCARABAEIDAE

https://scholar.valpo.edu/tgle/vol53/iss2/9 28 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

166 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes USA Native

0 0 0 1 4 1 0 3 1 1 0 1 0 7 5 2 65 18 24 17 Total 5

0 0 0 0 3 6 0 0 2 0 1 0 7 4 1 0 3 1 1 20 Ips

0 0 0 3 0 1 4 1 0 1 0 0 0 4 4 0 0 3 0 1 ETOH Funnel trap collections

0 0 0 1 0 8 0 0 0 1 0 0 9 0 0 1 4 0 42 13 a-p

4

S S P F F P F F F P F P F F FS FS FS FS FS FP

Methods

3

VI VI VI VI VI VII VII VII VII V VI VI IX VII IX VI VII VI VIII Months V VI VII V VI VIII collected VI VII VIII V VI VII VIII VI VII VIII IX V VI VII VIII IX

2

P P C C H H CP GS SO HP HP HP HP HP CHS CHP HSO GHO CHPS CHPSO Habitats

(Say)

(Forster)

(Thunberg) LeConte (Guérin-Méneville) (Linnaeus) Say LeConte (Klausnitzer) (Panzer) Haldeman Sharp * (Brown) (Guérin-Méneville)

(Haldeman) Laporte (Melsheimer) Guérin-Méneville unnamed nr. obscurus Say

1

family Species

Melanocanthon nigricornis Onthophagus hecate Contacyphon neopadi Contacyphon obscurus Contacyphon variabilis Contacyphon Nyholmia confusa Prionocyphon limbatus Sacodes thoracica Scirtes tibialis flavipennis Anaspis rufa Canifa pallipes Nicrophorus orbicollis Nicrophorus sayi Necrophila americana Oiceoptoma noveboracense Telephanus velox Sphindus americanus Silvanus muticus Silvanus Sub Scirtinae Anaspidinae Scraptiinae Nicrophorinae Silphinae Brontinae Silvaninae Sphindinae Appendix 1. Continued. FAMILY SCIRTIDAE SCRAPTIIDAE SILPHIDAE SILVANIDAE SPHINDIDAE

Published by ValpoScholar, 2020 29 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 167 NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA Pal NA Pal Pal Yes Yes Yes Yes Yes Yes Yes Yes Yes

0 1 0 0 1 0 6 0 0 0 4 1 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 (Continued on next page)

0 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0

0 1 0 0 1 0 5 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0

S P F P F P F P P P F F F P F F P P P P P P P P P P P P P F P P

IX IX IX IX IX IX IX IX VI VI VI VI VII VII VII VII VII VII VII VIII VIII VIII VIII V VI VI IX VII IX VII IX VII IX VI VII VIII IX VIII IX VI VII VIII IX

P P P P P P C C C C C C O H H H H H H H H H H CP HP HP HP CH CH CH CH label lost

(Mannerheim)

(Zetterstedt) (Bernhauer) (Gravenhorst) (Fabricius) (LeConte) Gravenhorst (Say) (Marsham) (Fabricius) (Gravenhorst)

Gravenhorst

sp.

sp.

sp.

sp.

Aleochara lata Aleochara Unknown sp. 1 Unknown sp. 2 Unknown sp. 3 Unknown sp. 4 Unknown sp. 5 Unknown sp. 6 Unknown sp. 7 Unknown sp. 8 Unknown sp. 9 Unknown sp. 10 Unknown sp. 11 Unknown sp. 12 Unknown sp. 13 Unknown sp. 14 Unknown sp. 15 Unknown sp. 16 Eusphalerum orientale Phloeostiba lapponicus Oxytelus laqueatus Paederus littorarius Palaminus Batrisodes Baeocera Belonuchus rufipennis Bisnius blandus Dinothenarus badipes Nudobius cephalus Ontholestes cingulatus Philonthus caeruleipennis Quedius fulgidus Aleocharinae Omaliinae Oxytelinae Paederinae Pselaphinae Scaphidiinae STAPHYLINIDAE

https://scholar.valpo.edu/tgle/vol53/iss2/9 30 Haack and Ruesink: Coleoptera Collected at Grass River Natural Area

168 THE GREAT LAKES ENTOMOLOGIST Vol. 53, Nos. 3–4 to 6 NA NA NA Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Yes Eur USA EurAs Native

4 0 2 0 1 1 1 7 0 0 0 1 1 1 3 0 2 2 1 0 0 0 1 4 Total 5

0 0 2 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 Ips

0 0 0 0 0 0 0 0 0 0 0 1 1 0 2 0 2 0 1 0 0 0 0 2 ETOH Funnel trap collections

4 0 0 0 1 0 1 7 0 0 0 0 0 1 1 0 0 1 0 0 0 0 1 1 a-p

4

F P F P F F F P P P F F F F P F F P P P FS FS FS FS Methods

3

IX IX VI VI VI VI VII VII VII VIII VIII VI VII VI VII VI VII VI VII VIII IX VIII IX V VI IX VII VIII VII VIII Months VI VIII IX collected VII VIII IX VI VII VIII V VI VIII IX

2

P P P P P P C H H H H H H H H CP CP HP CO HP HP CH GPS CHP Habitats

*

(Fall) (Say) (Melsheimer) (Linnaeus) (Couper) Gravenhorst (Ziegler) * (Say)

Gravenhorst (Melsheimer) (Couper) (Say) (Heer) (Gravenhorst) (Stephens) Casey Mannerheim Casey (Gravenhorst) (Say)

(Melsheimer)

sp.

sp. 1

sp.

family Species

Quedius fulvicollis Quedius laticollis Quedius peregrinus Quedius plagiatus Quedius Tasgius melanarius Carphacis Sepedophilus littoreus Tachinus fimbriatus Tachinus fumipennis Tachinus memnonius Capnochroa fuliginosa Hymenorus obesus Hymenorus Isomira oblongula Isomira pulla Isomira quadristriata Isomira sericea Mycetochara bicolor Corticeus parallelus Corticeus praetermissus Arthromacra aenea Strongylium tenuicolle Anaedus brunneus Anaedus Sub Tachyporinae Alleculinae Lagriinae Stenochiinae Appendix 1. Continued. FAMILY TENEBRIONIDAE

Published by ValpoScholar, 2020 31 The Great Lakes Entomologist, Vol. 53, No. 2 [2020], Art. 9

2020 THE GREAT LAKES ENTOMOLOGIST 169 - Yes Yes Yes Yes Yes Yes Yes Yes Yes totals of individuals = European, EurAs f it occurring in

3 1 2 4 4 1 2 1 2

0 1 0 0 0 0 1 0 0

2 0 0 4 4 1 0 1 0

1 0 2 0 0 0 1 0 2

F F F F F F F F F

V VI VIII V VI VI VII VI VII VIII IX VI VIII VII VIII IX

P P H H H H H CP CP ‡ (N = 14) indicates species for which Michigan specimens have been deposited in the Albert J. Cook Arthropod Re

(Say) Blanchard * Chevrolat Say (Horn) (Say) (Schaeffer) Melsheimer (Audinet-Serville)

Eustrophus tomentosus Synstrophus repandus Aulonothroscus constrictor Trixagus carinicollis Thymalus marginicollis Airora cylindrica Bitoma quadriguttata Synchita fuliginosa Aulonothroscus distans Aulonothroscus Eustrophinae Throscinae Peltinae Trogossitinae Colydiinae A species typed in bold and followed by the symbol * is considered a new state record for Michigan (N = 18), i.e., we are not aware of any published mention o Habitats: C = cedar, G grassland, H hardwoods, O other but primarily the Rail Trail, P pine, and S Sedge Meadow Trail. See text for details. Months: V = May, VI June, VII July, VIII August, and IX September. Methods: F = funnel traps, P pitfall and S sweeping. Funnel traps were baited with one of three lures: a-p = alpha-pinene, ETOH ethanol, and Ips ipsenol. See text for details. Numbers represent season-long Species listed as “Yes” are considered native to the continental USA, and for those exotic their ranges given follows: Eur Some sources have elevated certain subfamilies of Nemonychidae to family status, such as Cimberidinae being Cimberidae. TETRATOMIDAE THROSCIDAE TROGOSSITIDAE ZOPHERIDAE 1 Michigan, either in print or online. A species name followed by search Collection (ARC) at Michigan State University (MSU) prior to our study and published on SCAN, but with no records in print. Similarly, a species name followed by # (N = 1) indicates a species for which photographs of Michigan specimens have been posted on BugGuide, but with no in the ARC-MSU collection, nor any published records in print. Species names not followed by a symbol (N = 336) already have published in-print record of occurring Michigan. 2 3 4 5 collected by lure type and the total for all funnel traps combined. If no values are listed, then these species were not in traps. 6 7 = Eurasian, and Pal Palearctic. NA not applicable.

https://scholar.valpo.edu/tgle/vol53/iss2/9 32