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University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln

Center for Systematic Entomology, Gainesville, Insecta Mundi Florida

March 2002

An annotated checklist of Wisconsin (Coleoptera)

Nadine A. Kriska University of Wisconsin-Madison, Madison, WI

Daniel K. Young University of Wisconsin-Madison, Madison, WI

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Kriska, Nadine A. and Young, Daniel K., "An annotated checklist of Wisconsin Scarabaeoidea (Coleoptera)" (2002). Insecta Mundi. 537. https://digitalcommons.unl.edu/insectamundi/537

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 3 1

An annotated checklist of Wisconsin Scarabaeoidea (Coleoptera)

Nadine L. Kriska and Daniel K. Young Department of Entomology 445 Russell Labs University of Wisconsin-Madison Madison, WI 53706

Abstract. A survey of Wisconsin Scarabaeoidea (Coleoptera) conducted from literature searches, collection inventories, and three years of field work (1997-1999), yielded 177 representing nine families, two of which, and , represent new state family records. Fifty-six species (32% of the Wisconsin fauna) represent new state species records, having not previously been recorded from the state. Literature and collection distributional records suggest the potential for at least 33 additional species to occur in Wisconsin.

Introduction however, most of Wisconsin's scarabaeoid species diversity, life histories, and distributions were vir- The superfamily Scarabaeoidea is a large, di- tually unknown. verse group of , comprising more than 30,000 species in 13families worldwide. Over 1,530 species Methods occur in the continental United States. Despite this superfamily's popularity among scientists and hob- To determine what species had been taken in byists, and its incredible diversity (or perhaps, Wisconsin, historical collection and literature because of it), few scarabaeoid surveys have been records, as well as data from private regional collec- conducted in the United States. Blatchley (1910) tions [e.g. University of Wisconsin-Madison surveyed the Coleoptera of Indiana; Dawson (1922b) Research Collection (WIRC) (including the collec- catalogued the scarabs of Nebraska; Blatchley (1927 tion of Daniel K. Young), Milwaukee Public Muse- a,b, 1928 a-d, 1929 a-c, 1930 a,b) catalogued the um (MPMC), and Field Museum of Natural History scarabs of Florida; Rogers (unpublished M.S. the- (FMNH)] were compiled. Field sampling focused on sis, 1946) surveyed the scarabs of Washington State, regions of Wisconsin that had unique records or which served as the basis of Hatch's (1971) treat- were thought to be historically under-sampled. A ment of scarabs in his volumes on Coleoptera of the variety of collecting methods was employed, includ- Pacific Northwest; Helgesen and Post (1967) sur- ing blacklight, flight intercept traps (Malaise and veyed the saprophagous scarabs of North Dakota, window-pane), Lindgren funnel, unbaited and bait- which was followed by the phytophagous scarabs ed pitfall traps, leaf litter and sand sifting, and and trogids (Lago et al. 1979). Following Blatchley's hand collecting from flowers and foliage during works, Woodruff (1973) and Woodruff and Beck both daylight and evening hours. To expand geo- (1989) published more detailed surveys of Florida graphical and temporal species distributions, spe- laprostict scarabs and June beetles, respectively. cific sites were chosen for more in-depth examina- Expanding on Dawson's catalogue, Ratcliffe (1991 tions due to apparent species rarity or uniqueness a,b) published the most comprehensive (Nebras- of habitat types. Collecting efforts were modified kan) scarabaeoid surveys to date. Harpootlian (2001) for specific taxa, and natural history information surveyed the scarabaeoids of South Carolina. was gathered for each species. No comprehensive scarabaeoid surveys have Field samples from broader, ongoing surveys been conducted in Wisconsin, much less the Great [e.g. joint projects with the Wisconsin Department Lakes region. Rauterberg (1885-1889) recorded a of Natural Resources, Nature Conservancy surveys passalid and some stag beetles from Milwaukee (e.g. HemlockDraw), Fort McCoy inventory project, County; Snyder (1897) recorded 30 species from and inventory work at the Necedah National Wild- Dodge County; Dahl and Mahr (1991) conducted a life Refuge] also contributed significantly to this limited survey of June beetles ( spp.); study. Katovich (1995) and Katovich et al. (1998) surveyed the larvae of Wisconsin Phyllophaga. Until now, 32 Volume 16, No. 1-3, March-September, 2002, INSECTA MUNDI

Results Oconto; WC: Monroe; C: Wood; SW: Richland; SC: Green. This survey yielded 177 Wisconsin scarabaeoid Family species representing nine families. The familial phylogenetic arrangement of Scarabaeoidea adopt- Odontotaenius disjunctus (Illiger) - Schuster ed herein is based primarily on the works of Browne (1983)". Rauterberg (1889) reported this species to and Scholtz (1999). Subfamilial, tribal, and generic be widely encountered in Wisconsin, however, he arrangements are primarily based on Lawrence provided no further locality data, and no Wisconsin and Newton (1995) and Ratcliffe (1991b), respec- specimens have been found. It is possible that this tively. Vouchers of specimens taken during this species has been extirpated from Wisconsin due to survey are housed in the WIRC. A website has been the loss of mature, old growth forests with downed developed for Wisconsin scarabaeoids, and it in- timber of moderate to large diameter in the south- cludes information on their distributions in the ern half of Wisconsin. form of printable maps; it can be accessed at the following address: Family Lucanidae

Subfamily Lucaninae

Dorcus parallelus (Say) - Benesh (1960). Hand Additionally, a specimen-level database for all collected at night, on the trunlc of a standing Wisconsin records has been completed as part of a dead hackberry tree (Celtis sp.) and on the larger project for Wisconsin's Coleoptera fauna. trunk of Quercus rubra (Linnaeus) (red oak). The data reside with the authors and the WIRC June-July. NC: Lincoln, Taylor; SW: Grant, La under the BIOTA software (Colwell 1996). Crosse; SC: Dane, Sauk; SE: Milwaukee. In the following checklist of Wisconsin Scara- Lucanus capreolus (Linnaeus) - Benesh (1960). baeoidea, the 56 new state species records are Taken at UV light, hand collected at night from indicated in boldface. Species previously recorded porches, sidewalks, and sandy roads. May-Au- in the literature from Wisconsin are followed by the gust. WC: Monroe; SW: Crawford, Grant, Rich- pertinent literature reference. Those species with a land, Vernon; SC: Columbia, Dane, Iowa, Rock; literature record only (no specimens found and no SE: Jefferson, Kenosha, Milwaukee, Racine, locality data available) are marked with an asterisk Waukesha. (*) following the literature citation. To simplify Lucanus placidus Say - Benesh (1960). Shenefeldt county associations, Wisconsin has been divided and Simkover (1950) reported conifer damage into nine, 8-county regions (Map 1) after Hilsenhoff in tree nurseries in Wisconsin by the larvae (1995). Life history and phenological information feeding on the roots. On one occasion, an aggre- pertain solely to adult Wisconsin scarabaeoid records gation of about 15 males and females was found and has been taken directly from labels accompany- under Quercus velutina Lamarck (black oak) ing specimens. Thus, in reporting plant associa- shortly after dusk. May-August. NW: Burnett, tions, we report the data as indicated by specimen Rusk; NE: Marinette, Oconto; WC: Monroe, labels. However, we also commonly indicate an Pierce; C: Adams, Juneau, Portage, Waupaca, appropriate vernacular name when the specimen Wood; EC: Winnebago; SW: Grant, La Crosse, bears a latin binomial, and vice versa [e.g., on the Richland, Vernon; SC: Columbia, Dane, Lafay- trunk of a standing dead hackberry tree (Celtis sp.)] ette, Rock, Sauk; SE: Racine, Walworth. or [on the trunk of Quercus rubra (Linnaeus) (red Platycerus piceus (Kzrby) - Benesh (1960). Found oak)]. Many scarabaeoid species are long-lived as along the shore of Lake Superior. May-July. adults, and many phenologies are likely artifacts of NW: Bayfield, Douglas; NE: Marinette. human activity. We still lack sound natural history Platycerus virescens (Fabricius) - Benesh (1960). information for some species in Wisconsin. On the surface of rotting oak logs. April-Sep- tember. NW: Bayfield; NC: Oneida; NE: Door, Family Glaresidae Oconto; WC: Eau Claire; C: Waupaca, Waush- ara; SW: Grant; SC: Dane, Green, La Fayette, inducta Horn - Gordon (1970). All speci- Saulc; SE: Ozaukee, Racine. mens taken at UV light. June-September. NE: INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 3 3

Map 1. Regional divisions (nine, 8-county areas) of Wisconsin (after Hilsenhoff 1995)

Subfamily Nicaginae Subfamily Syndesinae

Nicagus obscurus (LeConte) - Katovich and Kriska Ceruchuspiceus (Weber) - Benesh (1960). Found in (2002). Dead and live specimens discovered oak logs in an advanced red-rot stage of decay. amongst driftwood and beach debris on the Active year around. NW. Bayfield, Polk; NC: shore of Lake Superior. May-July. NW: Dou- Price, Taylor; NE: Door, Florence, Oconto; WC: glas, Bayfield. Eau Claire, Jackson, Monroe; C: Juneau; SW: Grant, Richland; SC: Columbia, Dane, La Fay- ette, Rock, Sauk; SE: Milwaukee, Racine, Waukesha. 34 Volume 16, No. 1-3, March-September, 2002, INSECTA MUNDI

Family Trox unistriatus Beauvois - Vaurie (1955). At light and in pitfall traps baited with carrion or hu- Omorgus scabrosus (Beauvois) - In a pitfall trap man dung or pig dung. April-August. NW: baited with pig dung, malt, and molasses. June- Burnett, Douglas; NC: Oneida, Vilas; NE: Flo- August. SW: Grant, Richland. rence, Forest, Oconto; WC: Jackson, Monroe; Omorgus suberosus (Fabricius) - Vaurie (1955). At C: Juneau, Portage, Waushara, Wood; EC: light and in pitfall traps baited with pig dung, Fond du , Sheboygan; SW: Crawford, Grant, malt, and molasses. May-September. C: Wood; Richland, Vernon; SC: Columbia, Dane, Green, SW: Grant, Richland; SC: Columbia, Dane, Iowa, Rock, Sauk; SE: Kenosha, Racine. Sauk; SE: Jefferson. Trox variolatus Melsheimer - Vaurie (1955). On Trox aequalis Say -At light and in tree hole leaf dried carnivore dung containing hair and bone litter material. May-September. NW: Bayfield, fragments, on a dead raccoon, in pitfall traps Burnett; NC: Taylor; NE: Oconto; WC: Jack- baited with carrion or human dung or pig dung, son, Monroe, Pierce; C: Marquette, Waupaca, and in flight intercept traps. April-September. Wood; SW Crawford, Grant, Richland; SC: NE: Florence, Forest, Menominee; NC: Onei- Columbia, Dane, Green, Sauk; SE: Racine, da, Vilas; WC: Eau Claire, Monroe; C: Green Walworth, Waukesha. Lake, Waupaca, Waushara, Wood; SW: Grant, Trox atrox LeConte - At light. April-August. NW: La Crosse, Richland, Vernon; SC: Columbia, Burnett, Douglas; NE: Oconto; C: Waushara, Dane, Green, Iowa, La Fayette, Rock, Sauk; Wood; WC: Monroe; SW: Grant, Vernon; SC: SE: Racine, Walworth, Washington, Wauke- Columbia, Dane, Sauk; SE: Kenosha, Walworth, sha. Waukesha. Trox capillaris Say - On a dead dog in a ditch Family along a county highway. April-July. SW Rich- land; SC: Dane, Sauk. Bolboceras falli (Wallis) - Howden (1955). In un- Trox foveicollis Harold - Vaurie (1955). At light, in baited pitfall traps partially filled with propy- carrion-baited pitfall traps, and on dried carni- lene glycol and hand collected on a deer path in vore dung. May-August. WC: Monroe; C: Wood; northern mesic hardwood forest. June-July. SW: Richland; SC: Columbia, Dane, Iowa, Sauk. NC: Vilas; NE: Marinette; C: Wood. Trox hamatus Robinson - In flight intercept Bolboceras filicornis (Say) - Howden (1955). No traps, pitfall traps baited with carrion or pig ecological information available. May-Septem- dung/malt/molasses, and at light. May-August. ber. NW: Douglas; EC: Fond du Lac. NW: Douglas; NE: Marinette, Oconto; WC: Bolboceras liebecki (Wallis) - Howden (1955). Fly- Monroe, Pierce; C: Juneau, Marquette, Por- ing low over a footpath at dusk in small oak and tage, Waushara, Wood; SW: Grant, Richland, maple woods with organic, compacted, clay soil. Vernon; SC: Dane, Green, Rock, Sauk; SE: May-September. WC: Eau Claire, Jackson; SC: Waukesha. Dane, Green, Iowa, La Fayette, Sauk. Trox laticollis LeConte - No ecological informa- biplagiatum Dawson and McCol- tion available. April-June. SW: Crawford, SC: loch - Howden (1955)". No ecological informa- Dane. tion available. June-September. NC: Vilas. Trox robinsoni Vaurie - Vaurie (1955). In pitfall Bolbocerosoma bruneri Dawson and McColloch - traps baited with carrion or pig dung/malt/ Howden (1955). At UV lights near agriculture molasses. April-September. NW: Douglas; NC: stations, privately owned farms, and state- Vilas; NE: Door; C: Wood; SW: Grant. owned nurseries. April-October. SW: Craw- (Linnaeus) - Vaurie (1955). At light. ford, Grant, La Crosse, Vernon; SC: Columbia, May-August. NW: Douglas; NC: Vilas; NE: Dane, Sauk; SE: Racine. Oconto; C: Wood; SVV: Richland, Vernon; SC: fossatus (Haldeman) - At UV Dane, Sauk; SE: Jefferson. light near state-owned nursery near Boscobel, Trox sordidus LeConte - On a deer carcass. May- sandy savanna. May-August. SW: Grant, La June. SC: Iowa. Crosse. Trox striatus Melsheimer - At UV light. May- lazarus (Fabricius) - Howden (1955). At July. SW: Richland. UV light and in flight intercept traps. April- September. NW: Burnett; WC: Chippewa, Eau INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 3 5

Claire, Pierce; C: Wood; EC: Fond du Lac; SW: Douglas, Sawyer; WC: Monroe; SW: Richland; Grant, Richland; SC: Columbia, Dane, Iowa, SC: Columbia, Dane, Sauk. Sauk; SE: Racine, Walworth, Waukesha. Family Ceratocanthidae Family Germarostes aphodioides (Illiger) - At light Subfamily and hand collected at night from the bark of standing, dead oak trees. May-July. SW: Rich- Geotrupes balyi Jekel - Howden (1955). At light land; SC: Dane. and in carrion-baited pitfall traps. June-Sep- tember. NW: Bayfield, Burnett; NC: Iron, Mar- Family athon, Oneida, Vilas; NE: Forest, Marinette, Menominee, Oconto; WC: Eau Claire, Monroe; Subfamily C: Juneau, Marquette, Waupaca, Waushara, Wood; SW: Richland; SC: Dane, Iowa, La Fay- Tribe Aegialiini ette. Geotrupes blackburaii (Fabricius) - Howden conferta Horn - Among driftwood and (1955)". No ecological information available. beach debris along the shore of Lake Michigan. September-May. May-August. NE: Door; EC: Manitowoc; SE: Geotrupes hornii Blanchard-Howden (1955). Hand Milwaukee. collected in flight and in flight intercept traps. Aegialia criddlei Brown - In pitfall traps in May-September. NW: Bayfield; NC: Vilas; NE: sandy, oak-pine savanna. May-August. WC: Shawano; WC: Eau Claire, Monroe; C: Juneau, Jackson; C: Wood. Wood; SC: Sauk. Aegialia latispina LeConte - Sifted from leaf Geotrupes opacus Haldeman - Howden (1955). In litter in sandy, oak barrens. May-August. SW: pitfall traps baited with huinan dung or pig Richland. dung. March-June and July-September. C: Aegialia rufa (Fabricius) - Found dead amongst Adams, Marquette; SW: Grant, Richland; SC: beach debris on the shore of Lake Michigan. Dane, Rock, Sauk. May-June. EC: Sheboygan. Geotrupes semiopacus Jekel - Howden (1955). In pitfall traps baited with huinan dung or pig Tribe dung. May-September. NW: Burnett, Polk; NC: Vilas; NE: Menoininee; WC: Eau Claire; C: alterlzatus Horn - Rogers (1997)". No Juneau, Waupaca, Wood; EC: Sheboygan; SW: ecological information available. January-May. Crawford, Richland, Vernon; SC: Dane, Green, SC: Columbia. Iowa, La Fayette, Rock, Sauk; SE: Waukesha. Aphodius badipes Melsheimer - In a gray squir- Geotrupes sple~zdidus(Fabricius) - Howden (1955). rel nest in a hollowed, fallen tree and in tree In pitfall traps baited with carrion or human hole leaf litter in oak barrens and savanna. dung or pig dung, on stems of fungi, and at March-November. NE: Oconto; C: Waupaca, light. March-October. NW: Burnett, Douglas, Wood; SW: Grant, La Crosse, Richland; SC: Polk; NC: Vilas; NE: Forest, Marinette, Me- Sauk, Green. nominee, Oconto; WC: Eau Claire, Monroe; C: Aphodius bicolor Say - Under scat, in human Juneau, Marquette, Portage, Waupaca, Waush- dung-baited pitfall traps, and in flight intercept ara, Wood; SW: Grant, La Crosse, Richland; traps. September-April. SC: Green, La Fay- SC: Dane, Dodge, Green, Iowa, Rock, Sauk; ette, Sauk. SE: Racine. Aphodius distinctus (Muller) - Gordon (1983). In pitfall traps baited with human dung or pig Family Ochodaeidae dung and among tall grasses in barrens-type habitat. March-December. NW: Burnett, Dou- Ochodaeus musculus Say - At W light, in a win- glas, Polk; NE: Door; WC: Monroe, St. Croix; C: dow-pane trap and a Malaise trap, all located in Green Lake, Marquette, Waushara, Wood; SW: sandy, savanna habitat. June-September. NW: Grant, Pepin, Richland, Vernon; SC: Colum- 3 6 Volume 16, No. 1-3, March-September, 2002, INSECTA MUNDI

bia, Dane, Iowa, Rock, Sauk; SE: Jefferson, son, Monroe; SW: La Crosse, Richland; SC: Milwaukee, Waukesha. Sauk. Aphodius erraticus (Linnaeus) - Gordon (1983). In Aphodius iowensis Wickham- In a Malaise trap pitfall traps baited with human dung or pig and in pig dung/maltlmolasses-baited pitfall dung and in flight intercept traps. April-May. traps established in pocket gopher burrow sys- NW: Burnett; NC: Vilas; NE: Door, Oconto; tems. September-December. NW: Bayfield, WC: Eau Claire, Monroe; C: Marquette, Waush- Burnett, Polk; WC: Jackson; SW: Richland, ara; SW: Grant; SC: Columbia, Dane, Iowa, SC: Sauk. Rock; SE: Jefferson, Walworth. Aphodius kirni Cartwright - In window pane (Linnaeus) - Gordon (1983). traps and in pig dung/malt/molasses-baitedpit- In pitfall traps baited with human dung or pig fall traps established in pocket gopher burrow dung, in flight intercept traps, and found dead systems. April-December. NW: Burnett, Polk; in beach debris on the shore of Lake Michigan. WC: Jackson, Monroe; SW: Richland; SC: Sauk. March-October. NW: Polk; NC: Marathon; NE: Aphodius lentus Horn -At light, in pitfall traps Door, Oconto; WC: Clark, Jackson, Monroe; C: baited with human dung or pig dung, in flight Marquette, Portage, Waupaca, Wood; EC: Ke- intercept traps, and on deer dung. April-Au- waunee, Manitowoc; SW: Crawford, Grant, gust. NW: Douglas; NE: Oconto; WC: Monroe; Vernon; SC: Columbia, Dane, Green, Iowa, La C: Waushara; SW. Buffalo, Grant, Richland; Fayette; SE: Jefferson, Walworth, Waukesha. SC: Columbia, Dane, Green, Iowa, Rock. (Linnaeus) - Gordon (1983). In Aphodius leopardus Horn- Gordon (1983). On deer pitfall traps baited with human dung or pig dung, in pitfall traps baited with human dung dung and in cow dung in open pastures. April- or pig dung, and in flight intercept traps. Au- August. NW: Burnett; NE: Door, Oconto; C: gust-June. NW. Burnett, Douglas, Polk; NE: Waupaca, Wood; EC: Kewaunee; SC: Dane; Florence; WC: Eau Claire, Monroe; C: Wau- SE: Walworth. paca, Wood; SC: Sauk. (Linnaeus) - Gordon (1983). Aphodius magnificens Robinson - In pig dung1 At light, in pitfall traps baited with human malt/molasses-baited pitfall traps established dung or pig dung, in flight intercept traps, on a in pocket gopher burrow systems. August-De- dead raccoon and a dead deer, and among beach cember. NW: Polk; WC: Jackson, Monroe; SW: debris on the shore of Lake Michigan. April- La Crosse, Richland; SC: Sauk. September. NW: Bayfield, Douglas; NC: Tay- Aphodius peculiosus Schmidt - Larvae and lor; NE: Oconto; WC: Dunn, Jackson, Monroe; adults were sifted from a pocket gopher mound. C: Juneau, Marquette, Waupaca, Wood; EC: September-May. WC: Monroe. Manitowoc, Sheboygan; SVV: Buffalo, Craw- (Brahm) - Gordon (1983). In ford, Richland; SC: Columbia, Dane, Grant, scat, under a dead raccoon, and in pitfall traps Iowa; SE: Milwaukee, Ozaukee, Racine, Wal- baited with carrion or human dung or pig dung. worth, Waukesha. April-November. NC: Iron; NE: Oconto, Sha- Aphodius haemorrhoidalis (Linnaeus) -In cow wano; EC: Manitowoc; WC: Dunn, Monroe; C: dung in open pasture, in pitfall traps baited Green Lake, Marquette, Waupaca, Waushara, with human dung or pig dung, and among Wood; EC: Sheboygan; SW: Grant, Richland; beach debris on the shore of Lake Superior. SC: Columbia, Dane, Green, Iowa, La Fayette, May-September. NW: Douglas; NE: Marinette; Rock, Sauk; SE: Racine, Walworth. SC: Iowa, Sauk; SE: Jefferson, Walworth. Aphodius punctissumus Brown - In pig dung1 Aphodius hyperboreus LeConte - About a foot malt/molasses-baited pitfall traps established inside the opening of a woodchuck burrow lo- in pocket gopher burrow systems. October. NW: cated in open sandy grassland/oak barrens. Burnett. May-September. NW: Douglas; NC: Iron; NE: Aphodius rubeolus Beauvois - In pig dunglmaltl Door. molasses-baited pitfall traps in oak savanna. Aphodius insolitus Brown - Taken in pig dung/ April-September. SC: Columbia. malt/molasses-baited pitfall traps established Aphodius rubripennis Horn - In pitfall traps in pocket gopher burrow systems. August-De- baited with human dung or pig dung, in flight cember. NW: Burnett, Douglas, Polk; WC: Jack- intercept traps, and on deer dung. June-Octo- ber. NW: Burnett, Polk; NC: Vilas; NE: Mari- INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 37

nette, Oconto; WC: Clark, Eau Claire, Jackson, Tribe Monroe; C: Juneau, Marquette, Waupaca, Wood; SC: Columbia, Iowa, Sauk. fattigi Cartwright - Cartwright (1974)". Aphodius rusicola Melsheimer - Gordon (1983). At No ecological information available. February- light, in pitfall traps baited with human dung October. C: Wood. or pig dung, and in flight intercept traps. April- Ataeniusgracilis (Melsheimer) - Cartwright (1974). November. NW: Burnett, Douglas; NC: Taylor, At UV light. May-October. NC: Price; WC: Vilas; NE: Florence, Oconto; WC: Clark, Dunn, Monroe; C: Wood; SW: Richland, Vernon; SC: Eau Claire, Jackson, Monroe, Pierce; C: Ju- Dane. neau, Waupaca, Waushara, Wood; SW: Grant, Ataenius itnbricatus (Melsheimer) - Cartwright La Crosse, Richland; SC: Columbia, Dane, (1974). At light and sifted from leaf litter in Green, Iowa, La Fayette, Racine, Rock, Sauk; sandy, oak barrens. April-September. NW: SE: Walworth, Waukesha. Douglas; NE: Oconto; WC: Monroe; C: Wood; Aphodius stercorosus Melsheimer - Gordon (1983). SW: Grant, Richland, Vernon; SC: Columbia, At light, in pitfall traps baited with human Dane, Green. dung or pig dung, and in leaf litter near a fallen Ataenius miamii Cartwright - Sifted from leaf tree in mesic hardwood forest. April-August. litter in sandy oak barrens and among drift- NW: Douglas; NC: Vilas; WC: Jackson, Mon- wood and beach debris along the shore of Lake roe, Pierce; SW: Grant, Richland, Vernon; SC: Michigan. March-December. SW: Grant, Rich- Dane, Green, Rock, Sauk. land; SC: Sauk; SE: Milwaukee. Aphodius terminalis Say - On bones and hide of Ataenius punctifrons Cartwright - Cartwright a deer carcass. January-April, November. SW: (1974). At light, in pan traps and flight inter- Grant. cept traps, and among beach debris on the Aphodius walshii Horn - In pitfall traps baited shore of Lake Michigan. April-September. NE: with carrion or human dung or pig dung. May- Door, Oconto; WC: Monroe; C: Wood; SW: Grant, June. WC: Monroe; C: Marquette, Waushara; La Crosse, Richland, Vernon; SC: Dane, Green, SW: Grant; SC: Dane, Rock. Iowa, Sauk; SE: Kenosha, Milwaukee, Ozau- Dialytellus dialytoides (Fall) - In rotting fungi, kee, Racine. leaf litter, and flight intercept traps. June- Ataenius robustus Horn - Cartwright (1974). No August. NE: Marinette; WC: Eau Claire, Jack- ecological information available. March-July. son; C: Waupaca. SC: La Fayette. striatulus (Say) - In pitfall traps baited (Haldeman) - Cartwright (1974). with human dung or pig dung and in flight At light and hand collected in flight from culti- intercept traps. June-October. NW Burnett, vated grasses (sports fields, golf courses, ). Polk; NE: Florence; WC: Eau Claire, Monroe; March-September. NE: Oconto; C: Wood; SW: C: Juneau, Marquette, Waupaca, Waushara, Richland,Vernon; SC: Dane, Green, Iowa, Sauk; Wood; SW: Grant; SC: Iowa, Sauk. SE: Kenosha, Racine. Dialytes truncatus (Melsheimer) - Gordon (1983). (Say) - Cartwright (1974). At In pitfall traps baited with human dung or pig light, in pitfall traps baited with human dung dung and in flight intercept traps. June-Octo- or pig dung, sifted from leaf litter, and hand ber. NW: Burnett, Polk; NE: Florence, Oconto; collected in flight at dusk over old fields. March- WC: Monroe, Pierce; C: Juneau, Wood; SW: October. NW: Bayfield, Douglas; NE: Door, Grant, Richland; SC: La Fayette, Sauk. Oconto; WC: Monroe; C: Waupaca, Wood; EC: Dialytes ulkei Horn -Under scat, in pitfall traps Fond du Lac, Outagamie, Sheboygan, Win- baited with human dung or pig dung, and in nebago; SW: Crawford, Grant, Richland, Ver- flight intercept traps. June-September. WC: non; SC: Dane, Columbia, Green, Iowa, La Eau Claire, Monroe; C: Waupaca, Wood; SC: Fayette, Rock, Sauk; SE: Jefferson, Kenosha, Green, La Fayette, Sauk. Milwaukee, Ozaukee, Walworth, Waukesha.

Tribe Didactylini Tribe

Aidophusparcus (Horn) - No ecological informa- interruptus (Say) - Sifted from tion available. SW: Grant. leaf litter and in pan traps and barrier pitfall 3 8 Volume 16, No. 1-3, March-September, 2002, INSECTA MUNDI

traps, in sandy, oak barrens and in dune habi- non; SC: Columbia, Dane, Iowa, Rock, Sauk; tat near Lake Michigan. May-July. WC: Mon- SE: Kenosha, Walworth, Waukesha. roe; SW: Richland; SE: Kenosha. carolinus (Linnaeus) - No ecolog- notialis (Cartwright) - Sifted from ical information available. August. SC: Dane. leaf litter and in barrier pitfall traps in oak barrens and oak savanna habitats. June-Octo- Tribe ber. NW: Burnett; WC: Monroe; SW. Grant, Richland; SC: Columbia, Green, Sauk; SE: hecate (Panzer) - Howden and Cart- Kenosha. wright (1963). At W light, in pitfall traps caesus (Creutzer) - Jerath (1960). In baited with carrion or human dung or pig dung, beach debris along the shore of Lalre Michigan and in flight intercept traps. March-October. and the Wisconsin River. April-December. NE: NW: Burnett, Douglas, Polk, Washburn; NC: Door; EC: Manitowoc; SC: Dane, Iowa; SE: Vilas; NE: Florence, Marinette, Oconto, Me- Milwaukee. nominee, Shawano; WC: Clark, Eau Claire, Rhyssemusgermanus (Linnaeus) -Among beach Jaclrson, Monroe; C: Juneau, Marquette, Por- debris along the shore of Lalre Michigan. May- tage, Waushara, Wood; EC: Brown, Sheboyg- August. EC: Sheboygan. an; SW: Buffalo, Grant, La Crosse, Richland, Vernon; SC: Columbia, Dane, Green, Iowa, La Subfamily Fayette, Rock, Sauk; SE: Jefferson, Kenosha, Milwaukee, Ozaukee, Racine, Walworth, Tribe Canthonini Waukesha. Olzthophagus nuchicor7zis (Linnaeus) - Howden chalcites (Haldeman) - In pitfall traps and Cartwright (1963). At W light and in baited with human dung or pig dung and in pitfall traps baited with human dung or pig flight intercept traps. May-August. NW: Bur- dung. April-October. NW: Burnett, Douglas; nett, Douglas; NE: Oconto; WC: Eau Claire, NE: Door, Oconto, Shawano; WC: Clark, Eau Jaclrson, Monroe; C: Juneau, Washburn, Wood. Claire, Jackson, Monroe; C: Adams, Juneau, Canthon vigilans LeConte - No ecological infor- Marquette, Waushara; SW: Grant, Richland; mation available. May-July. C: Wood; EC: Fond SC: Columbia, Dane, Iowa, Rock, Sauk. du Lac. Onthophagus orpheus (Panzer)-Howden and Cart- (Beauvois) - In pitfall traps wright (1963). In deer dung, pitfall traps baited baited with human dung or pig dung and in with human dung or pig dung, flight intercept flight intercept traps. April-June. SW: Grant, traps, and Lindgren funnel traps. April-Sep- Richland; SC: Sauk. tember. NVV: Burnett, Douglas, Polk; NE: Oc- nigricornis (Say) - Williams and onto; WC: Eau Claire, Jackson, Monroe; C: Kriska (2001). In pitfall traps baited with hu- Juneau, Marquette, Waupaca, Waushara, man dung or pig dung, on a dead lizard, ob- Wood; SW: Grant, La Crosse, Richland; SC: served rolling a ball of carrion, and observed Columbia, Dane, Green, Iowa, La Fayette, Rock, breaking down, bundling, and rolling away Sauk; SE: Walworth. pieces of a gill cap mushroom. May-September. Onthophagus pennsylvanicus Harold - Howden NW: Burnett, Douglas; NE: Oconto, Shawano; and Cartwright (1963). Inflight intercept traps, WC:Monroe; C:Marquette, Portage; SW: Grant, in pitfall traps baited with carrion or human Richland; SC: Columbia, Dane, Iowa, Sauk. dung or pig dung, and on turkey dung and dog feces. April-September. WC: Eau Claire, Jack- Tribe son, Monroe; C: Juneau, Marquette, Wausha- ra, Wood; SW: Grant, La Crosse, Richland; SC: fricator (Fabricius)-Matthews (1961). Hand Columbia, Dane, Iowa Rock, Sauk; SE: Kenos- collected from tall grass stems at night, at light, ha. in pitfall traps baited with carrion or human Onthophagus striatulus (Beauvois) - In flight dung or pig dung, and in flight intercept traps. intercept traps, in pitfall traps baited with April-October. NW: Burnett; NE: Oconto; WC: human dung or pig dung, and in a jug trap Eau Claire, Monroe, Pierce; C: Green Lake, baited with pinene and 75% EtOH. May-July. Wood; SW: Crawford, Grant, Richland, Ver- INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 3 9

WC: Monroe; SW: La Crosse, Richland; SC: sis sp. (tickseed). May-October. WC: Pierce; Dane, Green, La Fayette, Sauk; SE: Walworth. SC: Crawford, Grant, Richland; SC: Sauk. Onthophagus tuberculifrons Harold - Howden and harperi Blanchard - Vaurie (1956). At Cartwright (1963)". No ecological information UV light and hand collected feeding on Cornus available. April-October. racemosa Lamarck (dogwood). April-August. SC: Dane, Green; SE: Racine, Waukesha. Tribe Phanaeini LeConte -At UVlight. June- August. NW: Burnett, Douglas; WC: Jackson; vindex MacLeay - In pitfall traps C: Waushara. baited with human dung or pig dung. April- Diplotaxis sordida (Say) - Vaurie (1956). At UV September. WC: Monroe; C: Marquette, Waush- light and in pig dung/malt/molasses-baited pit- ara; SW: Grant, Richland, Trempealeau, Ver- fall traps. May-August. NW: Burnett, Douglas; non; SC: Sauk. NE: Oconto; WC: Monroe; C: Marquette, Waushara, Wood; SW: Grant, Richland; SC: Subfamily Dane, Iowa, Sauk. Kirby - Vaurie (1956). At light Tribe Dichelonychini and found under a stone partially buried in the sand in oak savanna habitat. May-August. NW: albicollis (Burmeister) - At UV Douglas; NE: Door, Oconto; WC: Clark, Mon- light, in flight intercept traps and Lindgren roe, C: Wood; SW: Grant, La Crosse. funnel traps, and hand collected feeding diur- nally on needles of Pinus strobus Linnaeus Tribe Hopliini (white pine) in sandy oaWpine forest. May- July. Cornell (unpublished Ph.D. thesis, 1972) modesta Haldeman - Hardy (1977). Found also listed this species from Wisconsin. NW: amongst beach debris on the shore of Lake Burnett, Douglas, Sawyer; NC: Oneida; NE: Michigan and hand collected from Rosa sp., Marinette, Oconto; WC: Clark, Jackson, Mon- Sambucus racemosa Linnaeus (red elder), Rhus roe; C: Marquette, Waushara; SW: Grant, Rich- sp. (sumac), and Cornus racemosa Lamarck land; SC: Iowa, Sauk; SE: Waukesha. (dogwood). June-July. NW: Bayfield, Burnett, Dichelonyx canadensis (Horn) - No ecological Douglas; NE: Oconto; WC: Jackson, Monroe; information available. May-July. C: Wausha- C: Juneau, Washburn, Wood; EC: Manitowoc; ra. SW: Grant, Richland, Vernon, SC: Dane, Iowa, Dichelonyx elongatula (Schonherr) - At UV Sauk. light, in flight intercept traps, and hand collect- Hoplia trifasciata Say - Hardy (1977). Found nec- ed feeding diurnally on Acer rubrum Linnaeus taring on Prunus sp. (cherry) flowers. May- (red maple), Lupinus perennis Linnaeus (lu- July. NW: Burnett; NE: Brown, Marinette, pine), and Geranium sp. (geranium). April- Oconto, Shawano; WC: Monroe; C: Juneau, June. Cornell (unpublished Ph.D. thesis, 1972) Waupaca; SW: Richland, Vernon; SC: Rock, also listed this species from Wisconsin. NW: Sauk. Bayfield, Douglas; NE: Oconto; WC: Monroe; Hoplia trivialis Harold- Hardy (1977)". No ecolog- SC: Sauk; SE: Walworth. ical information available. May-July. Dichelonyx subvittata (LeConte) -At UV light. May-July. Cornell (unpublished Ph.D. thesis, Tribe 1972) also listed this species from Wisconsin. WC: Jackson; SW: Grant, La Crosse, Richland, subspinosus (Fabricius) - Carrillo Vernon; SC: Dane, Green, Sauk; SE: Wal- and Gibson (1960). In dung-baited pitfall traps, worth. hand collected feeding diurnally on Rosa sp. (rose), Salix sp. (willow), Quercus sp. (oak), Tribe Diplotaxini Apocynum sp. (dogbane), Verbascum sp. (mul- lein), and on ornamental composites in a flower Diplotaxis frondicola (Say) - Vaurie (1956). At UV garden, and a mating aggregation was observed light and hand collected at night from Coreop- during late summer on common milkweed (As- clepias sp.). June-July. NW: Burnett; NC: Onei- 40 Volume 16, No. 1-3, March-September, 2002, INSECTA MUNDI

da; NE: Brown, Door, Oconto, Shawano; WC: roe; C: Adams, Juneau, Marquette, Washburn, Eau Claire, Jackson, Monroe; C: Juneau, Mar- Waushara, Wood; EC: Fond du Lac, Sheboyg- quette, Portage, Wood; EC: Sheboygan, Win- an; SW: Crawford, Grant, La Crosse, Richland; nebago; SW: Grant, La Crosse, Richland, Ver- SC: Columbia, Dane, Green, Rock, Sauk; SE: non; SC: Columbia, Dane, Iowa; SE: Jefferson, Ozaukee, Waukesha. Sauk. Phyllophaga forsteri (Burmeister) - Luginbill and Tribe Painter (1953). At W light. April-July. SW: Crawford; SC: Columbia, Dane. Phyllophagaanxia (LeConte)- Luginbill and Paint- Phyllophaga fraterna Harris - Luginbill and Paint- er (1953). At UV light; larvae were the only er (1953). No ecological information available. Phyllophaga species occurring in Wisconsin March-July. C: Portage. cranberry beds (Katovich 1995; Katovich et al. Phyllophaga fusca (Froelich) - Luginbill and Paint- 1998), and they were also found in turf grasses er (1953). At UV light and found feeding on and irrigated sivicultural sites (balsam fir and Cornus racemosa Lamarck (dogwood); larvae of white pine plantations). March-August. NW: P. fusca are the primary species responsible for Barron, Bayfield, Douglas, Polk; NC: Ashland, damage to turf grass (Katovich 1995; Katovich Marathon, Price, Vilas; NE: Door, Florence, et al. 1998). March-July. NW: Polk; NC: Mara- Forest, Oconto; WC: Jackson, Monroe; C: Wau- thon; NE: Door, Florence, Marinette, Oconto; paca, Wood; EC: Brown, Fond du Lac; SW: WC: Chippewa; C: Marquette, Waupaca, Crawford, Grant, Richland; SC: Dane, Green, Waushara, Wood; EC: Fond du Lac; SW: Craw- Iowa, La Fayette, Sauk; SE: Jefferson, Milwau- ford, Grant, La Crosse, Richland; SC: Colum- kee, Racine, Waukesha. bia, Dane, Green, Iowa, Rock, Sauk; SE: Kenos- Phyllophaga balia (Say) - Luginbill and Painter ha, Milwaukee, Washington, Waukesha. (1953). At UV light. April-July. WC: Monroe; C: Phyllophaga futilis (LeConte)-Luginbill and Paint- Washburn; SW: Crawford, Grant; SC: Dane, er (1953). At light; adults are common in gar- Iowa, Rock. dens and at porch lights. March-July. NVV: Phyllophaga barda (Horn) - Luginbill and Painter Burnett; NE: Brown, Door, Oconto; WC: Mon- (1953)". No ecological information available. roe; C: Portage; EC: Fond du Lac; SW: Grant, April-July. Richland; SC: Columbia, Dane, Green, Iowa, Phyllophaga bipartita (Horn) - Luginbill and Paint- La Fayette; SE: Kenosha, Walworth. er (1953). No ecological information available. Phyllophaga gracilis (Burmeister) - Luginbill and May-July. NW: Barron. Painter (1953). At UV light and in flight inter- Phyllophaga corrosa (LeConte) - Luginbill and cept traps, in oak barrens and savanna habitat. Painter (1953). No ecological information avail- June-August. NW. Douglas; WC: Monroe, able. March-July. SC: Dane. Pierce; SW: Grant; SC: Dane. Phyllophaga crassissima (Blanchard) - Luginbill Phyllophaga hirticula (Knoch) - Luginbill and and Painter (1953)". No ecological information Painter (1953). At UV light. March-July. C: available. March-August. Wood; EC: Fond du Lac; SW: Crawford, Grant, Phyllophaga crenulata (Froelich) - Luginbill and Vernon; SC: Columbia, Dane, Iowa, La Fay- Painter (1953). At UV light, in flight intercept ette, Rock, Sauk. traps, and feeding on oak leaves at night in Phyllophaga hornii (Smith) - Luginbill and Paint- sandy, oak barrens and savanna habitat. March- er (1953)*. No ecological information available. August. NW: Barron, Douglas; WC: Jackson, March-June. Monroe, Pierce; C: Waushara, Wood; EC: Fond Phyllophaga ilicis (Knoch) - Luginbill and Painter du Lac; SW: Grant, Richland, Vernon; SC: (1953). At UV light. April-July. WC: Pierce, Columbia, Dane, Green, Iowa, Rock, Sauk; SE: Wood; SW. Crawford, Grant; SC: Dane, Iowa, Washington. Sauk. Phyllophaga drakii (Kirby) - Luginbill and Painter Phyllophaga implicita (Horn) -Luginbill and Paint- (1953). At W light and in flight intercept traps; er (1953). At UV light. March-July. SW: Craw- larvae were found in turf grass and sivicultural ford, Grant, La Crosse, Richland; SC: Dane, sites (Katovich 1995; Katovich et al. 1998). Green, Iowa; SE: Jefferson. March-July. NW: Burnett, Douglas, Polk; NC: Iron; NE: Florence, Oconto; WC: Jackson, Mon- INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 41

Phyllophaga inversa (Horn) - Luginbill and Paint- Phyllophaga vehemens (Horn) -Luginbill and Paint- er (1953). At UV light. April-June. NW: Craw- er (1953)". No ecological information available. ford, Grant, Richland; SC: La Fayette, Sauk. March-June. Phyllophaga longitarsa (Say) -Luginbill and Paint- Phyllophaga vilifrons (LeConte) - Luginbill and er (1953). At W light, in sandy oak barrens and Painter (1953). At UV light. April-June. NW: savanna habitat. May-July. NW: Grant; SC: Bayfield; SC: Dane, Green, Iowa; SE: Ozaukee. Iowa. hammondi LeConte -Young (1988). At Phyllophaga marginalis (LeConte) - Luginbill and UV light in mid-summer, active from 11 p.m. to Painter (1953). At UV light in forested habitats. about 2 a.m., in sandy oak savanna habitats March-July. NE: Florence; WC: Monroe; C: near the Mississippi and Wisconsin rivers. May - Wood; SC: Columbia, Dane, Iowa. September. SW: La Crosse; SC: Sauk. Phyllophaga micans (Knoch) - Luginbill and Painter (1953)". No ecological information available. Tribe March-June. Phyllophaga nitida (LeConte) -Luginbilland Paint- atracapilla (Kirby) -Dawson (1919a). At UV er (1953). At UV light. April-July. NW: Dou- light. May-July. NW: Bayfield, Douglas, Polk; glas, Sawyer; WC: Eau Claire, Pierce; EC: NC: Lincoln, Oneida, Price, Vilas; NE: Door, Fond du Lac; SW: Crawford, Grant; SC: Co- Florence, Oconto; WC: Jackson; C: Juneau, lumbia, Dane, Iowa, Rock. Monroe, Waupaca, Wood; SC: Sauk. Phyllophaga prunina (LeConte) - Luginbill and Serica georgiana Leng - Dawson (1932). At light Painter (1953). At UV light in sandy oak bar- and in flight intercept traps in sandy oak bar- rens and oak savanna habitat. March-July. rens and savanna. May-August. NW: Bayfield, WC: Monroe; C: Wood; SW: Grant, Richland; Douglas, Sawyer; NC: Lincoln; NE: Brown, SC: Columbia, Dane, Iowa, Sauk. Oconto; WC: Clark, Eau Claire, Jackson, Mon- Phyllophaga rugosa (Melsheimer) - Luginbill and roe; C: Juneau, Marquette, Wood; SW: Buffalo, Painter (1953). At light, in pan and flight inter- La Crosse, Richland; SC: Columbia, Dane, Sauk; cept traps, and among beach debris and leaves SE: Waukesha. on the shore of Lake Michigan; larvae are Serica intermixta Blatchley - Dawson (1932). At economically important in Wisconsin, feeding UV light, in flight intercept and unbaited in turf grasses and sivicultural sites (Katovich Lindgren funnel traps, and under leaf litter in 1995; Katovich et al. 1998). April-July. NW: a sandy blow. May-August. NW: Bayfield, Bur- Burnett, Douglas; NC: Ashland, Iron; NE: nett, Douglas; NC: Oneida, Vilas, NE: Brown, Brown, Door, Florence, Marinette, Oconto; WC: Florence; WC: Dunn, Eau Claire, Jackson, Jackson, Monroe; C: Portage, Waushara, Wood; Monroe; C: Marquette, Portage, Waupaca, EC: Fond du Lac, Manitowoc; SW: Crawford, Waushara, Wood; SW: Grant, La Crosse, Rich- Grant, La Crosse, Richland, Vernon; SC: Co- land; SC: Dane; SE: Waulresha. lumbia, Dane, Iowa, Rock, Sauk; SE: Kenosha, Serica loxia Dawson - Dawson (1920). At UV light Milwaukee, Ozaukee, Walworth. in sandy oak barrens and savanna habitats. Phyllophaga spreta (Horn) - Luginbill and Painter June-July. NW: Bayfield, Douglas; NE: Ocon- (1953). No ecological information available. to. April-May. SW: Crawford. Serica mystaca Dawson - At UV light in sandy Phyllophaga subtonsa (LeConte) - Luginbill and oak barrens and savanna habitats. April-June. Painter (1953). No ecological information avail- SW: La Crosse, Pierce; SC: Dane, Sauk. able. June-July. SC: Dane. Casey - Dawson (1922a). At W Phyllophaga tristis (Fabricius) - Luginbill and light, in flight intercept and unbaited Lindgren Painter (1953). At UV light in sandy oak bar- funnel traps, and observed feeding and mating rens and savanna habitats. March-June. NW: on leaves of Quercus velutina Lamarck (black Burnett, Polk, Washburn; WC: Jackson, Ju- oak), Prunus serotina Ehrhart (black cherry), neau, Monroe, Pierce; C: Marquette, Waupaca, and an aster at night. April-July. NE: Brown, Waushara, Wood; SW: Crawford, Grant, Rich- Oconto; WC: Jackson, Monroe, Pierce; C: Ju- land; SC: Columbia, Dane, Dodge, Iowa, La neau, Portage, Wood; SW: Grant, La Crosse, Fayette, Rock, Sauk; SE: Waukesha. Richland, Vernon; SC: Dane, Green, Sauk; SE: Walworth, Waukesha. 42 Volume 16, No. 1-3, March-September, 2002, INSECTA MUNDI

Serica sericea (Illiger) - Dawson (1922a). At UV push-up. March-May. WC: Monroe; C: Wood; light, in flight intercept and unbaited Lindgren SW: Grant, Richland; SC: Dane, Sauk. funnel traps, and observed feeding and mating Exomala orientalis (Waterhouse) - A series of on leaves of Prunus serotina Ehrhart (black adults was collected in a Japanese trap. cherry) at night. April-August. NVV: Burnett, June-August. SE: Milwaukee. Douglas, Washburn; NC: Taylor, Vilas; NE: japonica Newman - Large series col- Door, Florence, Langlade, Oconto; WC: lected in traps, and found feed- Chippewa, Clark, Jackson, Monroe, Pierce; C: ing on ornamental flowers in gardens and nurs- Adams, Juneau, Marquette, Waupaca, Waush- eries. June-August. NW: Polk; NE: Door; WC: ara, Wood; SW. Crawford, Grant, La Crosse, Eau Claire; SW: Grant; SC: Dane, Rock; SE: Richland; SC: Dane, Green, Sauk; SE: Kenos- Kenosha. ha, Milwaukee, Racine, Walworth, Waukesha. Strigoderrna arbicola (Fabricius) - Blackwelder Serica sponsa Dawson - Dawson (1919b). At UV and Arnett (1974). In pan, flight intercept, and light, in flight intercept traps, and observed barrier pitfall traps, all in sandy, open habitats, feeding on leaves of Acer rubrum Linnaeus (red and observed feeding on flowers of Monarda sp. maple) and Prunus serotina Ehrhart (black (horsemint), Rosa sp. (rose), and nectaring on cherry). April-July. SW. Crawford, La Crosse, flowers of Asclepias sp. (milkweed). June-July. Pierce, Richland, Vernon; SC: Dane, Green, NVV: Barron, Burnett, Douglas, Washburn; NC: Sauk; SE: Racine, Wauliesha. Ashland; NE: Oconto, Shawano; WC: Dunn, Serica tristis LeConte - Dawson (1922a). At W Eau Claire, Jackson, Monroe; C: Adams, Green light and in flight intercept traps. April-Au- Lake, Juneau, Portage, Waushara, Wood; SW: gust. NW: Bayfield, Washburn; NC: Vilas; NE: Crawford, Grant, Pepin, Richland; SC: Colum- Florence, Langlade, Oconto, Shawano; WC: bia, Dane, Green, Iowa, Rock, Sauk; SE: Jeffer- Jackson, Pierce; SC: Sauk; SE: Racine. son, Kenosha, Waukesha. Serica vespertina (Gyllenhal) - Dawson (1921). At UV light in sandy oak barrens and savanna Tribe habitats. May-June. WC: Pierce; C: Wood; SW: Crawford; SC: Sauk. lanigera (Linnaeus) - Saylor (1940). At UV light and found dead on sandy trails and roads. Subfamily April-July. NVV: Bayfield, Burnett, Douglas, Sawyer; NC: Ashland; NE: Door, Marinette, Tribe Oconto; WC: Monroe; C: Adams, Green Lake, Juneau, Marquette, Portage, Waushara, Wood; binotata (Gyllenhal) - Potts (1977). In SW: Crawford, Richland, Vernon; SC: Colum- pitfall and flight intercept traps, found along bia, Dane, Sauk; SE: Milwaukee, Ozaukee, sandy hiking trails, found in the mound of a Waukesha. pocket gopher push-up, and observed nectaring punctata (Linnaeus) - Hardy (1975). At on flowers of Prunus sp. (cherry). May-July. light, and three adults reared from pupae found WC: Monroe; C: Marquette, Wood; SW: Grant, in an unidentified, advancedly decayed tree Pepin, Richland; SC: Columbia, Dane, Iowa, stump. June-August. NE: Oconto; EC: Fond du Rock, Sauk. Lac, Winnebago; C: Wood; SW: Buffalo, Craw- Anomala innuba (Fabricius) - Potts (1977). Occur- ford, Grant, Richland; SC: Dane, Rock; SE: ring in prairie habitats, on Apocynum sp. (dog- Racine, Walworth, Waukesha. bane) and in tall grass. June-July. SW: Craw- ford; SC: Rock. Subfamily Anomala ludouiciana Schaeffer - Potts (1977). In sandy blows and on sandy trails in oak barrens Tribe Oryctini and savanna habitats. June-July. NW: Bur- nett; WC: Jackson, Monroe; C: Portage, Wood; jamaicensis (Drury) - Endrodi (1985). SW: Richland; SC: Columbia, Dane, Iowa, Sauk. At light and found dead on sidewalks. July- Melsheimer - Potts (1977). At August. NC: Vilas; NE: Door; C: Wood; SW: UV light in sandy oak barrens and savanna Grant, Richland; SC: Dane, Iowa, SE: Milwau- habitats, and in a mound of a pocket gopher kee, Racine. INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 43

Tribe Oconto; WC: Monroe; C: Marquette, Wausha- ra; SC: Rock. tridentatus (Say) -Gill and Howden (1985). harrisii Kirby - One adult At UV light. June-August. NW: Burnett, Dou- reared from a pupa taken in a soil sample, and glas; NE: Oconto; WC: Jackson, Monroe; C: hand collected in flight over a small sand dune Juneau, Wood; SW: Crawford; SC: Columbia, in oak barrens habitat. April-September. WC: Dane. Monroe; C: Wood. Ligyrusgibbosus (DeGeer) - Endrodi (1985). At UV Cremastocheilus Iznochi LeConte - Alpert (1994). light and found dead on sidewalks and under Found in a sandy pocket gopher push-up.March- gas station lights. April-October. WC: Monroe; September. WC: Monroe; C: Waushara; SC: C: Marquette, Waushara, Wood; SW: Grant, La Dane; SE: Milwaukee. Crosse, Richland; SC: Columbia, Dane, Sauk; Cremastocheilus wheeleri LeConte - Found SE: Jefferson. atop a Formica obscuripes Fore1 thatch nest, Ligyrus relictus (Say) - EndrAdi (1985). At UV the surface of the nest swarming with ants. light. April-October. NW: Polk; WC: Clark, June-August. C: Waushara; SC: Iowa. Jackson, Monroe; C: Wood; SW: Grant, Rich- Psilocnemis leucosticta Burmeister - One spec- land; SC: Columbia, Dane, Saulr, Jefferson, imen found dead in a backyard swimming pool Racine, Waukesha. and another found alive on a sandy road, adja- cent to a subterranean nest of the ant Doli- Subfamily Cetoniinae choderus taschenbergi (Mayr). May-July. WC: Jackson; C: Waushara. Tribe Subfamily Trichiinae fulgida (Fabricius) - Ratcliffe (1991a). In flight intercept traps and hand collected in Tribe Osmodermini flight through a sandy oak savanna site. May- July. NW: Polk; NC: Iron; NE: Door; WC: eremicola (Knoch) - Howden (1968). Monroe; SW: Crawford; SC: Dane, Iowa, Rock; At light; larvae commonly encountered in large, SE: Racine. rotted stumps and fallen tree trunks. March- (Linnaeus) - Ratcliffe (1991a). In September. NW: Burnett, Washburn; NE: Oc- flight intercept and baited pitfall traps, sifted onto; WC: Monroe; SW: Crawford, Grant, Rich- from pocket gopher mounds, hand collected in land; C: Green Lake, Wood; SC: Dane, Sauk; flight along sandy paths, and larvae and adults SE: Jefferson, Kenosha, Racine, Walworth. observed in the vicinity of Formica thatch ant Beauvois - Howden (1968). nests. March-September. NW: Burnett, Saw- Found on a gravel road in northern mesic yer, Washburn; NC: Iron; NE: Brown, Oconto; hardwood forest, and reared from larvae found WC: Jackson, Monroe; C: Adams, Portage, in a large rotted stump. April-September. NW: Waupaca, Waushara, Wood; EC: Sheboygan; Bayfield; NC: Oneida, Vilas; WC: Monroe; SW: SW: Crawford, Grant, Richland, Vernon; SC: Crawford; SC: Dane, Sauk. Columbia, Dane, Rock, Sauk; SE: Milwaukee, Washington, Waukesha. Tribe Trichiinj Euphoria selpulchralis Fabricius - No ecolog- icalinformation available. May-September. SE: Gnorimella maculosa (Knoch) - Howden (1968). In Racine. Lindgren funnel traps in forested areas, and a single adult was observed nectaring on ninebark Tribe (Physocarpus opulifolius (Linnaeus)) in an area of northern mesic hardwood forest. May-July. Cremastocheilus castaneus Knoch - Alpert (1994). WC: Jackson, Monroe; C: Wood; SC: Sauk. Observed in flight in oak savanna, found on the assimilis (Kirby) - Howden (1968). In surface of Formica sp. nests, and a single adult Malaise traps and observed feeding on flowers reared from a sandy mound of a pocket gopher of Monarda sp. (horsemint), Phlox sp. (phlox), burrow. April-September. NW: Bayfield; NE: Prulzus sp. (cherry), and Rosa sp. (rose). June- July. NW: Polk, Rusk, Sawyer, Washburn; NC: 44 Volume 16, No. 1-3, March-September, 2002, INSECTA MUNDI

Oneida, Vilas; NE: Marinette, Oconto; WC: Aphodius concavus Say: Iowa. Monroe, St. Croix; C: Wood; EC: Manitowoc; Aphodius femoralis Say: Illinois, Indiana, Michi- SE: Milwaukee. gan. (Fabricius) - Howden (1968). Aphodius manitobensis Brown: Southcentral Can- Observed feeding on a raspberry flower. May- ada (Ontario, Manitoba), possibly in northern August. NVV. Polk; WC: Monroe; C: Portage, Michigan, Minnesota. Waupaca, Wood; SW: Grant; SC: Columbia, Aphodius pinguis Haldeman: Michigan, Minneso- Dane, Iowa, Sauk; SE: Milwaukee, Outagamie. ta, North Dakota. Trichiotinus viridans (Kirby) - Observed feed- Aphodius serval Say: Illinois, Indiana, Iowa. ing on flowers of Anemone sp. (anemone), Ascl- Aphodius vittatus Say: Illinois, Indiana. epias sp. (milkweed), Physocarpus opulifolius (Linnaeus) (ninebark), Ceanothus americanus Tribe Psammodiini Linnaeus (New Jersey tea), Cornus racemosa Lamarck (dogwood), Rosa sp. (rose), and Rhus sonatus LeConte: Illinois, Iowa. sp. (sumac). May-July. NVV: Burnett, Polk; WC: Eau Claire, Jackson, Monroe; C: Juneau, Wood; Subfamily Scarabaeinae SVV: Grant, La Crosse; SC: Dane, Iowa, Sauk. Tribe The following species have distributions that could potentially extend into Wisconsin. This list is histeroides Weber: Illinois, Indiana, Iowa. based on known distributions, published litera- ture, and specimens examined during the course of Tribe Canthonini this study. The most pertinent literature relating to the development of this list includes Cornell (1972), (Linnaeus): Illinois, Indiana, Endrodi (1985), Gordon (1983), Gordon and Cart- Iowa, Michigan, Minnesota. wright (1980), Howden (1955, 1968), Luginbill and Painter (1953), Ratcliffe (1991a), and Vaurie (1956). Subfamily Melolonthinae Each species listed is followed by the state(s) and province(s) from which it is currently known that Tribe Dichelonychini show a reasonable proximity to Wisconsin. Dichelonyx diluta (Fall): Michigan. Family Bolboceratidae Dichelonyx fuscula (LeConte): Illinois. Dichelonyx macclayi Fraser: Illinois, Indiana. Bolboceras simi (Wallis): Indiana, Michigan, Min- nesota. Tribe Diplotaxini Bolboceras thoracicornis (Wallis): Illinois, Indiana, Iowa, Michigan. Diplotaxis atlantis Fall: Illinois, Iowa. Bolbocerosoma confusum Brown: Illinois, Indiana, Diplotaxis basalis Fall: Illinois, Indiana. Iowa. Diplotaxis blanchardi Vaurie: Illinois, Iowa.

Family Scarabaeidae Tribe Melolonthinj

Subfamily Aphodiinae Phyllophaga congrua (LeConte): Illinois, Indiana, Iowa. Tribe Aegialiini Phyllophaga ephilida (Say): Illinois, Indiana, Iowa. Phyllophaga fervida (Fabricius): Illinois, Indiana, Aegalia humeralis Brown: Michigan. Iowa, Michigan, Minnesota. Phyllophaga hirtiventris (Horn): Indiana, Iowa. Tribe Aphodiini Phyllophaga lanceolata (Say): Iowa. Phyllophaga praetermissa (Horn): Illinois. Aphodius borealis Gyllenhal: Southcentral Cana- Phyllophaga rubiginosa (LeConte): Michigan. da (Ontario, Manitoba), possibly northern Mich- igan, Minnesota. INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 45

Subfamily Dynastinae associated with these habitats. These gophers 'rely on the deep soils for tunnelling, and they feed on Tribe roots of grasses in shallow feeding tunnels. Numer- ous subspecies of pocket gophers in the United borealis Arrow: Illinois, Indiana States are associated with aphodiine dung beetles Bland: Illinois, Indiana, Iowa endemic to their tunnel systems. Wisconsin pocket morator (Fabricius): Illinois, Iowa, In- gopher populations not only host five of these diana aphodiine species (Aphodius insolitus, A. iowensis, A. kirni, A. magnificens, A. peculiosis), but numer- Subfamily Cetoniinae ous other scarabaeoids take advantage of the mi- crohabitat provided by the pocket gophers in the Tribe Cetoniini form of sandy mounds or "push-ups" created when the gophers clear out their tunnels. The investiga- Stephanucapilipennis Kraatz: North Dakota, Min- tion of these unique habitats and associations using nesota. particular collecting methods contributed signifi- cantly to this survey, and it provides a springboard Subfamily Trichiinae for further studies both in Wisconsin as well as surrounding regions. Tribe Trichiinj Acknowledgments Trichiotinus affinis (Gory and Percheron): Illinois, Indiana, Michigan We thank our lab colleagues (Craig Brabant, Trichiotinus bibens (Fabricius): Illinois, Indiana, Jim Dunford, Jeff Gruber, Kerry Katovich, Michele Michigan Price, Alistair Ramsdale, and Andrew Williams) for assistance and contribution of material to this Discussion study. We also acknowledge curators and staff at the University of Wisconsin-Madison Insect Re- The scarabaeoid fauna of Wisconsin ranges search collection (Steven Krauth), Milwaukee Pub- from common, widespread species to species which lic Museum (Gary Noonan), and Field Museum of are uncommonly encountered and strongly influ- Natural History (Alfred Newton Jr. and Margaret enced by Wisconsin's unique abiotic and biotic Thayer) for assisting us during examination of the features, such as physical geography, climate, nat- collections under their care. Several col- ural boundaries of the Great Lakes and Mississippi leagues throughout the United States provided River, vegetation, and soils. Many Wisconsin spe- assistance with species identifications and nomen- cies are widespread throughout the United States, clatural issues; we specifically acknowledge Bruce but there are many species representing northern, Gill and Robert Gordon (scarabaeines and aphodi- southern, eastern, or western faunas whose ranges ines), Henry Howden (bolboceratids and geotrupids), include Wisconsin. Paul Lago (Serica), Chris Marshall (passalids), Certain portions of the state host a greater Scott McCleve (Diplotaxis), Paul Skelley (aphodi- number of species because of the vegetational and ines), and William Warner (Cremastocheilus). The soil components occurring there, particularly oak private collections of Byron Buchli, Herb Grimek, barrens and oak savanna with sandy and silty loam Michael Sabourin, Walter Suter, and Daniel Young soil. Of the 177 Wisconsin species, 108 were collect- proved to be very valuable sources of new state ed in areas containing oak barrens or savanna records and numerous county records. Several pri- habitat; 27 of those 108 species include new state vate landowners (Richard and Mary Norman, Ted records. The "dung beetles" (subfamilies Aphodii- and Donna Spencer, and Eric Strickler), allowed nae and Scarabaeinae) not only comprise the ma- access to their property to collect data. We thank jority of new state records, but they comprise the Thomas Meyers, Wisconsin Department of Natural majority of species encountered in the barrens and Resources (WDNR), for assisting us with acquiring savanna habitats. Many of these species are tied to permits to sample in Wisconsin's State Natural the mammals and birds that inhabit these areas. Areas, and other WDNR employees (especially One such mammal, Geomys bursarius (Shaw), the Kathy Kirk) who brought in scarabaeoids for iden- pocket gopher, is endemic to the rich, sandy soils tification. A special thanks to Dave Almquist and 46 Volume 16, No. 1-3, MarchSeptember, 2002, INSECTA MUNDI

Paul Skelley for reviewing this manuscript and Blatchley, W. S. 1930b. The Scarabaeidae of Flor- offering helpful feedback. This research was sup- ida. The Florida Entomologist 14: 25-35. ported in part by a Hatch grant (WIS-4056) to Browne, J. and C. H. Scholtz. 1999. A phylogeny Daniel K. Young. Additional agencies provided of the families of Scarabaeoidea (Coleoptera). funding to cover much-needed supplies and travel Systematic Entomology 24: 51-84. expenses, in particular, Lois Almon Small Grants Carrillo, J. L. and W. W. Gibson. 1960. Respao de Program, Milwaukee Zoological Society, and Natu- las especies mexicanus del ghnero Macrodacty- ral History Museums Council Small Grants. Fund- lus (Coleoptera, Scarabaeidae), con observa- ing for the specimen-level database and web-based ciones biol6gicas de algunas especies. Secretar- material came, in part, from a WDNR (ATRI:Aquatic ia de Agricultura y Ganaderia, Oficina de Estu- and Terrestrial Resource Inventory) grant to Daniel dios Especiales, Folleto Tecnico No. 39: 1-102. K. Young; for these resources we are grateful. Cartwright, 0.L. 1974. Ataenius, Aphotaeniz~s, and of the United States and Literature Cited Canada (Coleoptera: Scarabaeidae: Aphodiinae). Smithsonian Contributions to Zoology 154: 1- Alpert, 6. D. 1994. A comparative study of the 106. symbiotic relationships between beetles of the Colwell, R. 1996. BIOTA: The biodiversity data- Cremastocheilus (Coleoptera: Scara- base manager. Sinauer Associates, Sunderland, baeidae) and their host ants (Hymenoptera: Massachusetts. Formicidae). Sociobiology 25: 1-276. Cornell, J. F. 1972. A taxonomic review of the Benesh, B. 1960. Coleopterorum Catalogus Sup- beetle genus Dichelonyx Harris in North Amer- plementa, Pars 8. Lucanidae. W. Junk, Graven- ica (Scarabaeidae: Melolonthinae). Unpublished hage, Netherlands, 178 pp. Ph.D. dissertation, Oregon State University, Blackwelder, R. E. and R. H. Arnett, Jr. 1974. 253 pp. Checklist of the beetles of Canada, United States, Dahl, R. A. and Mahr, D. L. 1991. Light trap Mexico, Central America, and the West Indies. records of Phyllophagain Wisconsin 1984-1987. Vol. 1, part 3, the scarab beetles, ant-loving The Great Lakes Entomologist 24: 1-8. beetles, clown beetles, and related groups. Fam- Dawson, R. W. 1919a. New species of Serica (Scar- ilies separately paginated. Biological Research abaeidae), I. Journal of the New York Entomo- Institute of America, Latham, NY. logical Society 27: 32-39. Blatchley, W. S. 1910. Coleoptera or beetles known Dawson, R. W. 1919b. New species of Serica (Scar- to occur in Indiana. The Nature Publishing abaeidae), 11. Journal of the New York Entomo- Company, Indianapolis, Indiana, 1386 pp. logical Society 27: 223-225. Blatchley, W. S. 1927a. The Scarabaeidae of Flor- Dawson, R. W. 1920. New species of Serica (Scar- ida. The Florida Entomologist 11: 44-46. abaeidae), 111. Journal of the New York Ento- Blatchley, W. S. 192713. The Scarabaeidae of Flor- mological Society 28: 208-212. ida. The Florida Entomologist 11: 55-62. Dawson, R. W. 1921. New species of Serica (Scar- Blatchley, W. S. 1928a. The Scarabaeidae of Flor- abaeidae), IV. Journal of the New York Ento- ida. The Florida Entomologist 12: 9-14. mological Society 29: 160-168. Blatchley, W. S. 1928b. The Scarabaeidae of Flor- Dawson, R. W. 1922a. New species of Serica (Scar- ida. The Florida Entomologist 12: 22-30. abaeidae), V. Journal of the New York Entomo- Blatchley, W. S. 1928c. The Scarabaeidae of Flor- logical Society 30: 154-169. ida. The Florida Entomologist 12: 44-46. Dawson, R. W. 1922b. A synopsis of the Scara- Blatchley, W. S. 1928d. The Scarabaeidae of Flor- baeidae of Nebraska (Coleoptera). University of ida. The Florida Entomologist 12: 63-65. Nebraska, University Studies 22: 163-244. Blatchley, W. S. 1929a. The Scarabaeidae of Flor- Dawson, R. W. 1932. New species of Serica (Scar- ida. The Florida Entomologist 13: 33-37. abaeidae), VI. Journal of the New York Ento- Blatchley, W. S. 1929b. The Scarabaeidae of Flor- mological Society 40: 529-562. ida. The Florida Entomologist 13: 52-56. Endrodi, S. 1985. The Dynastinae of the world. Blatchley, W. S. 1929c. The Scarabaeidae of Flor- Dr. W. Junk, Dordrecht, 800 pp. ida. The Florida Entomologist 13: 69-77. Gill, B. D. and H. F. Howden. 1985. A review of Blatchley, W. S. 1930a. The Scarabaeidae of Flor- the North American genus Aphonus LeConte ida. The Florida Entomologist 14: 13-17. INSECTA MUNDI, Vol. 16, No. 1-3, March-September, 2002 47

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