Coleoptera: Scarabaeidae - Salisbury University

Illustrated Keys to the Scarabaeinae (Coleoptera: Scarabaeidae) of Maryland Author(s): Simone N. Nemes and Dana L. Price Source: Northeastern Naturalist, 22(2):318-344. Published By: Eagle Hill Institute DOI: http://dx.doi.org/10.1656/045.022.0208 URL: http://www.bioone.org/doi/full/10.1656/045.022.0208

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Northeastern Naturalist NORTHEASTERN NATURALIST S.N. Nemes and D.L. Price

Vol. 22, No. 2 22(2):318–344

Illustrated Keys to the Scarabaeinae (Coleoptera: Scarabaeidae) of Maryland Simone N. Nemes1 and Dana L. Price1,* Abstract - This paper provides illustrated taxonomic keys for the identification of Maryland dung beetles. Twenty-one species are recognized from 5 tribes, including 7 genera. Keys are supported by 37 figures, including 7 habitus illustrations.

Introduction Dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) are widely known as beneficial insects for their habits of removal and burial of dung. Ecological services they provide include, but are not limited to, nutrient cycling, bioturbation, plant growth enhancement, and secondary seed dispersal in tropical forests (Nichols et al. 2008). Tunnelers and rollers transfer dung below the soil, thereby increasing the likelihood of micro-organismal and chemical changes in the soil and thus improving plant height and leaf production (Miranda et al. 2000, Nichols et al. 2008). These beetles are also economically important for the suppression of dung-breeding flies and nematodes that reduce livestock productivity. Losey and Vaughn (2006) suggested that dung beetles save North American cattle farmers an estimated 380 million dollars a year. Our current knowledge of Maryland Scarabaeinae is limited to a checklist provided by Staines (1984) that noted 20 species of Scarabaeinae previously reported from Maryland and surrounding states. Currently there is no comprehensive identification guide for the Mid-Atlantic Region. Our research provides brief descriptions and keys to 21 species known from Maryland including Onthophagus taurus (Schreber), a Eurasian species that was first reported in Florida in 1975 (Fincher and Woodruff 1975). Methods We surveyed available literature for existing records of Maryland dung beetles. Specimens examined include those held at Salisbury University (personal collection of Dana L. Price) as well as specimens at the National Museum of Natural History (Smithsonian Institution, Washington, DC). Keys and species descriptions have been modified from several resources including The Scarab Beetles of Florida (Woodruff 1973), Scarab Beetles of South Carolina (Harpootlian 2001), The Beetles of Northeastern North America (Downie and Arnett 1996), and The Scarabaeoid Beetles of Nebraska (Ratcliffe and Paulsen 2008). Additional monographic works regarding Maryland species are noted throughout. Department of Biological Sciences, Salisbury University, 1101 Camden Avenue, Salisbury, MD, 21801. *Corresponding author - [email protected]. 1

Manuscript Editor: Christopher M. Heckscher 318

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All figures were outlined using a Leica M125 Camera Lucida Microscope, and then details were provided as needed. Dorsal and ventral aspects of a Scarabaeinae dung beetle are shown in Figures 1 and 2, with key structures named.

Figure 1. Dorsal aspect of a dung beetle.

Figure 1. Ventral aspect of a dung beetle.

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Results and Discussion The behavior and ecology of the subfamily Scarabaeinae are better known than for any other group of scarabaeoids. These beetles range in size from 2 to 60 mm, although most US species do not exceed 30 mm (Ratcliffe and Paulsen 2008). They can be distinguished from most other scarabs using several characteristics including: sternites that are shorter in length than the metasternum; 8–9 antennal segments; a posterior tibia with a single spur (except for Melanocanthon which has two; Harpootlian 2001, Ratcliffe and Paulsen 2008). Key to the Tribes of Adult Scarabaeinae in Maryland (after Ratcliffe and Paulsen 2008) 1. Metatarsus with basal segment longer than next 3 segments combined (Fig. 3) . ..............................................................................................ONTHOPHAGINI Metatarsus with basal segment shorter than next 3 segments combined (Fig. 4)....2 2. Meso- and metatarsi lacking claws (Fig. 4)....................................PHANAEINI Meso- and metatarsi with claws (Fig. 5)................................................................3 3. Meso- and metatibiae slender, barely enlarged at apex (Fig. 5). Head and pronotum lacking horns or protuberances................................... CANTHONINI Meso- and metatibiae enlarged apically (Fig. 4). Head variable, often with horns or protuberances.................................................................................................4 4. Protibiae with inner apical region truncate, curved (Fig. 6). Head often with horn or protuberance. Length > 10 mm............................................... COPRINI Protibiae with inner apical region right-angled, flat (Fig. 7). Head simple. Length < 10 mm.........................................................................................ATEUCHINI

Figures 3–7. Tribe characteristics: (3) Onthophagini—metatibia and tarsus; (4) Phanaeini— metatibia and tarsus; (5) Canthonini—metatibia and tarsus; (6) Coprini—protibia and tarsus; and (7) Ateuchini—protibia and tarsus. 320


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Tribe ONTHOPHAGINI Burnmeister, 1846 Genus Onthophagus Latreille, 1802 Keys to the Onthophagus species occurring north of Mexico were provided by Howden and Cartwright (1963). The genus Onthophagus has over 1500 species worldwide with 37 species occurring in the United States (Ratcliffe and Paulsen 2008). This genus is the most speciose of the dung beetles in Maryland with 9 species present. Species range in size from 3.0 to 10.5 mm. Onthophagus orpheus (Panzer) is shown in Figure 8.

Figure 8. Onthophagus orpheus. 321

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Key to species of the genus Onthophagus in Maryland 1. Pronotum distinctly tuberculate (Fig. 9) or centrally punctate with both small tubercles and punctures on lateral margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Pronotum uniformly smooth or punctate (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Head and pronotum black, dull to moderately shiny. Male majors without any distinct protuberances or horns. Body slightly to moderately setigerous. Length 3.0–9.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Head and pronotum bright shining emerald green or bluish, elytra usually bicolored, mottled brown and black. Male majors with long, forward-projecting bifurcate horn on pronomtum. Body densely setigerous. Length 5.0–8.1 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. concinnus Laporte 3. Head with 2 small tubercles between eyes, clypeal apex bidentate (Fig. 11), sometimes emarginated. Pronotum punctate centrally with both small tubercles and punctures on lateral margins; male pronotum without any protuberances or horns. Elytra usually with some orange-brown patches on lateral margins and apical region. Length 3.0–5.5 mm. . . . . . . . . . . . . . . . O. tuberculifrons Harold Head smooth between eyes; females with two transverse ridges on head (Fig. 12); clypeal apex of male sharply upturned as subtriangular tooth (Fig. 13), that of females smaller. Pronotum densely covered in oblong tubercles and short setae; male majors with forward-projecting, bifurcate, dorsoventrally flattened horn on pronotum (Figs. 13–14), horn reduced in male minors and without tooth. Elytra uniformly black. Length 5.2–9.5 mm. . . . . . . . . . . . . . . . . . . O. hecate (Panzer) 4. Clypeal apex broadly emarginate, usually bidentate (Fig. 15). Length 3.5–5.1 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. subaeneus (Palisot de Beauvois) Clypeal apex without emargination; rounded, bidentate, or truncate. . . . . . . . . . . 5 5. Elytra color yellow or tan, mottled with black patches or spots. Males with median horn on base of head between or just behind eyes (Fig. 16); female head with two transverse ridges (Fig. 17); female pronotum with small anterior bump. Length 6.3–8.1 mm. . . . . . . . . . . . . . . . . . . . . . . . . . O. nuchicornis (L.) Elytra color and pattern not as described as above. Males with either no horns or 2 horns on head, females variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Figures 9–10. Pronotal sculpturing (40x magnification): (9) pronotum tuberculate; (10) pronotum punctate.

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Figures 11–12. (11) Onthophagus tuberculifrons, head tuberculate (with two small tubercles between eyes), and clypeus usually bidentate; (12) Onthophagus hecate, head of female with two transverse ridges.

Figures 13–14. (13) Onthophagus hecate, clypeal apex curving upwards, (14) Onthophagus hecate pronotum.

Figure 15. Onthophagus subaeneus, clypeal apex broadly emarginate.

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6. Head, pronotum, and elytra metallic green to cupreous red. Pronotum of male major with flattened, bifurcate protuberance projecting over the head (compare to Onthophagus hecate; Fig. 14). Male minors with horns reduced or absent; pronotal horn shortened. Females with anterior pronotal ridge (Fig. 18). Length 5.0–9.0 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. orpheus (Panzer) Head, pronotum, and elytra color variable: either completely black or brown, or bicolored with black pronotum and brown elytra; luster variable, dull to shiny. Pronota of males and female without protuberance . . . . . . . . . . . . . . . . . . . . . . 7 7. Length 8.0–10.5 mm. Male majors with 2 long, curved horns on head extending posteriorly over pronotum; male minors with 2 pointed tubercles at base of head. ...................................................................... O. taurus (Schreber) Length 7.0 mm or less. Males with no horns or with 2 erect horns, one behind each eye............................................................................................................. 8

Figures 16–17. Onthophagus nuchicornis: (16) male, (17) female.

Figure 18. Onthophagus orpheus female anterior pronotal ridge.

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8. Head and pronotum shiny, dark brown to black, or black with emerald reflections; elytra uniformly brown or black, or with brownish-yellowish streaks or patches at base and/or apex. Males with slender, erect horn behind each eye (Figs. 19–20). Pronotum of males with median anterior swelling (Fig. 20), sometimes adorned with tubercles. Female pronotum with swelling less pronounced (Fig. 21), reduced to 2 bumps on small females. Length 4.2–6.8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. striatulus (Palisot de Beauvois) Head, pronotum, and elytra generally black. Males without horns on head. Pronotum of males and females without median swelling or tubercles. Length 3.5–5.0 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. pennsylvanicus Harold Onthophagus concinnus Laporte, 1840 Onthophagus concinnus Dejean 1836:157 (nomen nudum) Ontophagus concinnus Laporte 1840:87 (genus misspelled, headings are correct) Onthophagus viridicollis Sturm 1843:108 (nomen nudum) Onthophagus protensus Melsheimer 1845:134 Length 5.0–8.1 mm. This species is easily distinguished by color alone. Pronotum and head metallic green, rarely bluish green; elytra bicolored: light brown at base and/or apex and dark brown to black medially. Head notched at margins, notches more prominent in males. Males sometimes with clypeus elongated into a subtriangular horn, reflexed strongly, otherwise clypeus generally rounded in males and female. Head punctate, sometimes rugose towards apex. Females with two ridges that traverse the head medially and basally. Pronotum and elytra densely setigerous, setae color yellow. Pronotum tuberculate, males with anteriorly projecting flattened bifurcate horn extending from pronotum. Elytra shiny; intervals finely granulate with small, tubercles in rows. Striae with shallow punctation. Onthophagus concinnus is widely distributed, occurring in the spring and fall from New Jersey to Florida and westward to Mississippi (Howden and Cartwright 1963). In Maryland, the highest abundances are during April and May on cow

Figures 19–21. Onthophagus striatulus: (19) Frontal view of male, (20) dorsal view of male head and pronotum, and (21) dorsal view of female pronotum. 325

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pastures. It has been taken from the dung of small mammals, as well as human dung, cow dung, and pitfall traps with no bait (Price et al. 2012; Staines 1984). Onthophagus hecate (Panzer, 1794) Scarabaeus hectate Panzer 1794:5 Copris hastator Fabricius 1798:28 Copris latebrosis Fabricius 1801:34 Copris obtectus Palisot de Beauvois 1805:25 Onthaphagus furcicollis Dejean 1836:157 (nomen nudum) Onthaphagus lama Dejean 1836:157 (nomen nudum) Onthaphagus scabricollis Kirby 1837:126 Onthaphagus sayi Laporte 1840:87 Length 5.2–9.5 mm. Overall body black. Surface densely setigerous, setae white to pale yellowish-white to dark yellow. Head variably textured in females and male minors; in male majors either smooth, punctate, rugose, or rugopunctate—usually a combination of at least two of the aforementioned. Pronotum with pronounced, large, oblong tubercles, tubercles shiny, with moderately dense setae. Male majors on pronotum with flattened, forward-projecting horn, horn usually expanded apically, bifurcate in most specimens. Male minors with pronotal horn reduced, or with a low, ridged or rounded swelling. Female with tubercles reduced or absent at apex of pronotum. Male majors with clypeus sharply reflexed upward into a subtriangular tooth. Females with two transverse ridges on head; clypeus rounded and reflexed. Six elytral striae between humerus and suture, striae impressed and sometimes punctate. Intervals granulate, with raised tubercles. Pygidium punctate, punctures becoming more noticeable at apex, faint to obsolete at base. Onthophagus hecate is the most widely distributed and one of the most common North American species. It occurs in southern Canada and all of the United States except Washington, Oregon, Nevada, and California (Howden and Cartwright 1963). It is abundant throughout the state of Maryland from April to October, and has been taken from dung, fungi, carrion, rotten vegetables, and malt traps as well as at lights (Howden and Cartwright 1963, Staines 1984). Onthophagus nuchicornis (Linnaeus, 1758) Scarabaeus nuchicornis Linnaeus 1758:347 Onthophagus rhinoceros Melsheimer 1846:134 Length 6.3–8.1 mm. Head and pronotum black, dully shining; both granulate and punctate; head with small punctures, pronotum with large punctures. Clypeus broadly emarginate, may be rounded in some specimens. Females with two transverse ridges on head; the basal ridge being wider. Males with a single short horn at base of head; horn widened at base. Pronotum with a median protuberance, protuberance usually more pronounced in females and in such cases with a ridge. Elytra yellowish tan to brown, mottled with black patches; base with a black margin; elytra intervals finely granulate interspersed with finely setigerous punctures. 326

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Onthophagus nuchicornis is a Eurasian species that was accidentally introduced into North America prior to the 1840s (Hoebeke and Beucke 1997, MacRae and Penn 2001). This species has since become widely established in North America, with populations on both the east and west coasts of Canada and the United States, probably by separate introductions (Howden and Cartwright 1963). In Maryland, it has been taken from cow, horse, and human dung, April through August (Price et al. 2012; Staines 1984). Onthophagus orpheus (Panzer, 1794) Scarabaeus orpheus Panzer 1794:5 Length 5.0–9.0 mm. Overall color metallic green, purplish brown with emerald reflections, reddish, or cupreous. Males with head distinctly notched at margins, separating frons from clypeus. Female head only marginally notched, with two transverse ridges situated basally and medially; medial ridge traverses the entire width of head. Male majors with one short horn behind each eye; clypeus reflexed upward. Pronotum smooth with moderate dense punctures. Male pronotum with flattened bifurcate horn projecting forward above head. Male minor with horn reduced or absent. Female with pronotal ridge, no horn. Elytral striae weakly impressed, punctate. Elytral intervals smooth with irregular punctures, moderately setigerous. Onthophagus orpheus has been reported in the eastern United States except the far Northeast, west to Texas and Kansas (Harpootlian 2001). This species is found across Maryland from May to October and has been taken from cow and human dung, carrion, and mammal’s nests. Onthophagus pennsylvanicus Harold, 1871 Onthophagus moeris Sturm 1826:178 (nomen nudum) Onthophagus pennsylvanicus Dejean 1836:158 (nomen nudum) Onthophagus pennsylvanicus Harold 1871:115 Onthophagus falcipes Harold 1871:115 Length 3.5–5.0 mm. Head, pronotum, and elytra dull black. Setae yellowish brown. Head finely granulate with small, setigerous punctures. Male clypeus truncate to rounded, reflexed at center; clypeus lightly to moderately punctate, punctures small to moderately large. Female clypeal apex similar to males but not as strongly reflexed. Females with one to two transverse ridges on head; front ridge rounded anteriorly. Pronotum finely granulate with setigerous punctures. Elytra with 6 punctate striae between humerus and suture. Elytral intervals with rows of small setigerous tubercles. Onthophagus pennsylvanicus occurs in all states east and south of South Dakota and Colorado except New Mexico, Vermont, and Maine (Howden and Cartwright 1963). This species is found across Maryland from May to September and has been taken from dung, carrion, rotting fruit, and fungi (Price et al. 2012, Staines 1984). 327

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Onthophagus striatulus (Palisot de Beauvois, 1809) Copris striatulus Palisot de Beauvois 1809:92 Onthophagus cervicornis Kirby 1825:565 Onthophagus cavicornis Haldeman and LeConte 1853:54 Onthophagus castaneus Melsheimer 1846:134 Onthophagus viridicollis Gemminger and Harold 1869:1030 Length 4.2–6.8 mm. Head color dark brown to black, with faint cupreous sheen. Elytra color variable, frequently dark brown to black with light brown, yellow, or amber patches at base and/or apex; or light brown, mottled with dark brown patches. Some specimens may be entirely orangish brown, dark brown, or black with cupreous reflections. Setae brownish-yellow, moderate to dense on pronotum and elytra, sparse to moderate on head. Pronotum densely punctate, black with dark green or copper reflections. Male majors easily identifiable by 2 vertical, erect horns present basally on head. Male minors with horns reduced to pointed tubercles. Female head rugose or punctate, sometimes both, and with two transverse ridges, one at the base of head, the other present medially, which completely traverses entire width of head. Elytra with 6 impressed, punctate striae between suture and humerus. Elytral intervals 1–3 usually with irregularly spaced, setigerous tubercles. Onthophagus striatulus occurs in the eastern half of the United States, westward to Nebraska, and Texas. This species is found across Maryland from April to October, and has been taken from dung, fungi, carrion, and sugar lure (Stains 1984). This species is commonly taken from rotting fungi (Howden and Cartwright 1963), and Halffter and Matthews (1966) suggest it is almost exclusively mycetophagous. Onthophagus subaeneus (Palisot de Beauvois, 1811) Copris subaneus Palisot de Beauvois 1811:105 Onthophagus cribricollis Horn 1881:76 Length 3.3–5.1 mm. General color variable: brownish red, dark brown, black with metallic reflections, or dark green, shiny. Head smooth, interspersed with setigerous punctures. Head with two small transverse ridges, basal ridge wider, more pronounced. Clypeus reflexed, broadly emarginate, nearly dentate either side of middle. Pronotum covered with large, setigerous punctures. Elytra granulate with setigerous tubercles. Onthophagus subaeneus occurs in the eastern United States from Pennsylvania south to Florida and west to Kansas, Oklahoma, and Texas (Howden and Cartwirght 1963). This species has been reported from western Maryland (West of the Chesapeake Bay), from June to October. It has been taken from fungi, dung, debris, and carrion, and prefers moist woodlands (Staines 1984). Onthophagus taurus (Shreber, 1759) Scarabaeus taurus Schreber 1759:7 Length 8.0–10.5 mm. Color mostly black to dark brown; elytra may be dark brown, or lighter. Head of male generally finely granulate with shallow punctures, 328

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sparsely setigerous at apex; clypeus rounded, elongated, reflexed. Larger males with two long, prominently curved horns projecting posteriorly. Head of females with two transverse ridges, basal ridge more prominent; clypeus rounded, reflexed but not as strongly as in males; surface punctate. Pronotum with deep, lateral impressions for reception of horns in males and evenly convex in females. Pronotum finely granulate with large punctures, smaller punctures may also be present. Elytral striae shallowly impressed, punctate. Elytral intervals granulate, with very small punctures, sparsely setigerous, setae usually restricted to apex and outer margins. Onthophagus taurus is reported from Europe, Africa, India, United States, and Australia. This species was initially introduced in Florida but has since expanded its range to include the eastern United States from New York and Ohio, and west to Texas; it has also been intentionally introduced into California (Hoebeke and Beuke 1997). This species is statewide in Maryland from April to October and prefers cow and horse dung. Onthophagus tuberculifrons Harold, 1871 Onthophagus tuberculifrons Sturm 1843:108 (nomen nudum) Onthophagus tuberculatus Gemminger and Harold 1869:1038 (nomen nudum) Onthophagus tuberculifrons Harold 1871:115 Length 3.0–5.5 mm. Color dull black or faintly cupreous with orange-brown spots at the apices of the elytra. Head with two basal tubercles, which distinguishes it from all other species. Head finely granulate with moderately large punctures interspersed. Clypeus dark brown, reflexed, emarginate or conspicuously bidentate. Pronotum finely granulate with large, densely spaced punctures. Elytra striae shallowly impressed, punctate. Elytra intervals finely granulate, intervals with two rows of setigerous tubercles (except for interval 1), elytra sometimes with faint cupreous sheen. Head, pronotum, and elytra moderately setigerous, setae golden to orange. Onthophagus tuberculifrons is found from Connecticut to Florida and west to Texas, Oklahoma, and Kansas (Howden and Cartwright 1963). This species prefers sandy soils and is most common in the eastern parts of Maryland from March to October. It has been taken from dung, fungi, and rotten vegetables. Tribe PHANAEINI Kolbe, 1905 Genus Phanaeus Say, 1823 Only 1 of the 7 species of Phanaeus known north of Mexico is found in Maryland. The genus was revised by Edmonds (1994). A male minor of Phanaeus vindex MacLeay is shown in Figure 22. Phanaeus vindex Macleay, 1819 Phanaeus vindex MacLeay 1819:133 Length 11–22 mm. Color metallic green with gold or red iridescence. Head transversely rugose, color metallic green with gold or red iridescence. Head of 329

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male with single acuminate horn, curving up and over the pronotum; variable in length. Head of female with erect, traverse tubercle with apex usually slightly bifid. Pronotum metallic green-red with gold reflections and metallic gold reflections at margins. Surface coarsely, transversely rugose; with two impressions present on each side of pronotal disc. Male majors with pronotal disc in shape of large, flat

Figure 22. Phanaeus vindex, male minor. 330

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subtriangular shield; posterior angles acutely produced, elevated. Females with slight longitudinal depression at base of disc; front with transverse ridge, more pronounced in larger individuals. Elytra metallic green, longitudinally rugose. Surface coarsely rugose; striae weakly impressed. Phanaeus vindex occurs throughout the eastern parts of the United States ranging north to Massachusetts in the east and South Dakota in the west (Ratcliffe and Paulsen 2008). This species is found throughout Maryland from May to October (Staines 1984) and has been taken from dung, carrion, rotting fruit, and traps with no bait (Price et al. 2012). Tribe CANTHONINI Lansberge, 1874 Key to Species of the Tribe Canthonini 1. Metatibiae each with 2 terminal spurs............................. Genus Melanocanthon Metatibiae each with a single terminal spur....................................Genus Canthon Genus Melanocanthon Halffter, 1958 Only one species of Melanocanthon is present in Maryland. This genus is easily distinguished from all other genera of Scarabaeinae by the presence of 2 apical spurs on the posterior tibiae (i.e. the metatibiae). Melanocanthon bispinatus (Robinson) is shown in Figure 23. Melanocanthon bispinatus (Robinson, 1941) Canthon bispinatus Robinson 1941:128 Length 6.0–10.0 mm. General color gray to black, dull. Clypeus quadridentate; outer 2 teeth shorter and blunter than medial teeth. Head, pronotum, elytra, and pygidium finely granulate, interspersed with larger, raised granules which are moderate to dense in distribution. Metatibia with 2 apical spurs. Elytral striae obsolete. Melanocanthon bispinatus is found along the coastal eastern United States from New York to Mississippi (Harpootlian 2001). In Maryland, it is most commonly found from May to September in the coastal counties (Worcester and Wicomico), although Staines (1984) reported it from Anne Arundel. It has been taken from dung, dry fungi, dead animals, rotting fruit, and malt (Price et al. 2012, Staines 1984). Harpootlian (1995) reported it burying dead Carabids in a sandy road. Genus Canthon Hoffmannsegg, 1817 The genus Canthon is represented by 5 species known to occur in Maryland. Canthon are generally large beetles (length 12.0–24.0 mm), with the exception of Canthon viridis (Palisot de Beauvois) (length > 5 mm). Canthon chalcites (Haldeman) is shown in Figure 24. 331

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Figure 23. Melanocathon bispinatus.

Figure 24. Canthon chalcites.

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Key to Species of the Genus Canthon 1. Clypeus bidentate (Fig. 25)................................................................................2 Clypeus quadridentate (Fig. 26)................................................C. probus (Germar) 2. Surface shining, bright green or purplish bronze. Length 2.0–4.0 mm................ ..........................................................................C. viridis (Palisot de Beauvois) Surface dull or shining, black to bronze or greenish. Length 12.0–20.4 mm..........3 3. Pronotum and pygidium with larger granules raised above field of smaller granules (Fig. 27).....................................................................................................4 Pronotum scabrous (irregularly roughened) (Fig. 28); pygidium uniformly finely granulate. Length 13.0–21.0 mm.................................C. chalcites (Handelman) 4. Eyes slender, width less than half of length as seen from above (Fig. 29). Length 12.0–17.0 mm. .............................................................C. pilularius (L.) Eyes large, suboval, width half or more of length as seen from above (Fig. 30). Length 13.0–20.4 mm. ......................................................C. vigilans (LeConte)

Figures 25–26. (25) clypeus bidentate, (26) Canthon probus clypeus quadridentate. Figures 27–28. Pronotal sculpturing (40x magnification): (27) Canthon pilularius pronotum and pygidium with larger granules raised above surface of smaller granules, (28) Canthon chalcites pronotum scabrous.

Figures 29–30. Dorsal view of head: (29) Canthon pilularius, (30) Canthon vigilans. 333

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Canthon chalcites (Haldeman, 1843) Coprobius chalcites Dejean 1836:151 (nomen nudum) Coprobius chalcites Haldeman 1843:304 Length 13.0–21.0 mm. Color cupreous or black. Clypeus bidentate, reflexed. Head densely scabrous. Head and gena smoothly joined, continuous, lacking emargination. Eyes narrow, length more than twice the width. Pronotum scabrous. Protibial spur bifurcate in males and acute in females. Elytra densely granulate with mixed small and large granules. Elytral striae faint between humerus and suture. Pygidium with small, dense granules. Similar in appearance to C. pilularius but can be differentiated by its scabrous pronotum and finely granulate pygidium when viewed at high magnification. Canthon chalcites occurs throughout most of the eastern half of the United States (Ratcliffe and Paulsen 2008). In Maryland, it is reported across the state from April to October. It has been taken from dung, rotting fruit, and road kill (Price et al. 2012; Staines 1984). Canthon pilularius (Linnaeus, 1758) Scarabaeus pilularius Linnaeus 1758:349 Scarabaeus laevis Drury 1770:79 Scarabaeus hudsonias Forster 1771:24 Scarabaeus volvens Fabricius 1792:66 Coprobius obtusidens Ziegler 1844:45 Length 12–17.0 mm. Overall color black, sometimes with weak bronze, blue, or copper hue. Head, pronotum, elytra, and pygidium granulate; granules small and large mixed. Clypeus reflexed, apex bidentate, teeth broadly emarginate, rounded. Eyes narrow, length more than twice width. Protibial spur bifurcate in males, acute in females. Elytra with 6 weakly impressed striae between humerus and suture. Similar in appearance to Canthon chalcites, but it can be differentiated by the textures on the pronotum and pygidium as noted above. Canthon pilularius is found in all of the states east of Rocky Mountains (Harpootlian 2001, Ratcliffe and Paulsen 1998). In Maryland, it was reported from Anne Arundel, Howard, and Worcester counties during June and July. It has been taken from cow, horse, sheep, and human dung (Staines 1984). Canthon probus (Germar, 1824) Cathon probus Germar 1824:98 Canthon minor Sturm 1843:104 Length 3.8–6.5 mm. Color black. Head, pronotum, elytra, and pygidium finely granulate, sometimes with very small punctures, which in many cases are only visible at oblique view in high magnification. Clypeus distinctly quadridentate, weakly reflexed. Clypeus and gena separated by a deep triangular emargination. Elytral striae 1–6 feebly impressed between suture and humerus. 334

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Canthon probus occurs from New Jersey south to Florida and west to Utah and Arizona (Harpootlian 2001). In Maryland, it has been reported west of the Chesapeake Bay from May to August. It prefers sandy wooded areas and has been taken from deer, rabbit, and cow dung (Staines 1984). Canthon vigilans LeConte, 1858 Canthon vigilans LeConte 1858:16 Length 13.0–20.4 mm. Color black, frequently with weak green or purple reflections. Head, elytra, and pronotum densely granulate, granules mixed small and large. Clypeal apex deeply bidentate, teeth subtriangular, reflexed. Differentiated from C. chalcites and C. pilularius by the size of eyes when viewed dorsally (Fig. 30): eyes large, greatest width slightly shorter than length. Elytra with 6 faintly impressed striae between suture and humerus. Pygidium surface same as head except larger granules are less dense. Canthon vigilans occurs in the eastern United States west to Nebraska, Kansas, and Texas (Peck and Thomas 1998). This species has been reported west of the Chesapeake Bay (District of Columbia) from February to July. It was reported in dung and at lights (Staines 1984). Canthon viridis (Palisot de Beauvois, 1805) Copris viridis Palisot de Beauvois 1805:24 Ateuchus obsletus Say 1823:208 Onthophagus viridicatus Say 1835:173 Canthon metallicus Sturm 1843:104 Length 2.0–4.7 mm. Body color bright metallic green, deep purple, dark blue, or copper. Head, pronotum, and pygidium with surface finely granulate, interspersed with small punctures. Clypeus bidentate, teeth subtriangular, reflexed. Elytra finely granulate, with minor to obsolete punctures. Elytral striae 1–6 faint between suture and humerus. This species is easily identified by its size and coloration. Canthon viridis is widely distributed in the eastern United States and into northeastern Mexico (Ratcliffe and Paulsen 2008). In Maryland, this species has been collected in scattered places on the western shore of Maryland (Staines 1984). East of the Chesapeake Bay, it has only been collected in Talbot and Queen Anne’s counties (Price et al. 2012, unpubl. data). Staines (1984) suggests it prefers densely wooded areas and has been reported from mammal and bird dung. Tribe COPRINI Leach, 1815 Key to Genera of the Tribe Coprini 1. Body length > 19.5 mm. Clypeus broadly rounded; elytron with 6 impressed striae between suture and humerus.......................................Genus Dichotomius Body length < 18.5 mm. Clypeus notched at apex; elytra each with 8 distinct striae between suture and humerus................................................ Genus Copris 335

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Genus Dichotomius Hope, 1838 Dichotomius carolinus (L.) is the only species of this genus found in Maryland. This species is easily identified by its large size: length > 19.5 mm. It is shown in Figure 31. Dichotomius carolinus (Linnaeus, 1767) Copris carolinus Linnaeus 1767:545 Pinotus bituberculatus Harold 1869:127 Length 19.8–28.2 mm. Overall color black, shining. Clypeus in both sexes coarsely trasversely rugose. Males with strong, transverse, bifid tubercle on center of clypeus (well in front of eyes), females with a similar tubercle (smaller) but more closely situated at the base of the clypeus (between eyes). Pronotum margined,

Figure 31. Dichotomius carolinus. 336

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extremely convex on basal half, with deep impression on each side at carina behind declivity. Elytra smooth, with 6 punctate striae present between suture and humerus, striae 1–3 widened posteriorly. Dichotomius carolinus occurs in the eastern United States from New York to Florida and west to southern Texas and southern Arizona (Ratcliffe and Paulsen 2008). In Maryland, it has been reported across the state from June to October (Staines 1984). This species has been taken from cow, horse, and human dung and has been collected at lights (Price et al. 2012; Staines 1984). Genus Copris Müller, 1764 Two species of Copris are known to occur in Maryland. This genus is easily distinguished from other Maryland species by the black color, distinct elytra striae, triangularly notched clypeus, and broad head. Copris fricator (Fabricius) is shown in Figure 32.

Figure 32. Copris fricator.

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Key to Species of the Genus Copris 1. Lateral pronotal carina absent; pronotal anterolateral angles rounded at apex (Fig. 33). Length 8.0–13.0 mm............................................. C. minutus (Drury) Lateral pronotal carina present; pronotal anterolateral angles sinuate at apex (Fig. 34). Length 10.5–18.2 mm....................................... C. fricator (Fabricius) Copris fricator (Fabricius, 1787) Scarabaeus fricator Fabricius 1787:15 Scarabaeus tullius Olivier 1789:118 Copris anaglypticus Say 1823:204 Length 10.5–18.2 mm. Color black. Similar in appearance to C. minutus, but larger. Head, pronotum, and elytra punctate; punctures largest on pronotum. Clypeus notched at apex and broadly rounded. Male majors with a long, erect, acuminate horn on center of head; male minors with a short horn or no horn. Females with rounded tubercle on center of head, occasionally lacking. Pronotum of male majors with up to 4 prominent tubercles at apex. Antennae reddish brown; club yellowish brown to reddish brown, pubescent. Elytra with punctate striae; intervals slightly convex. Copris fricator occurs in the eastern United States, reaching into Canada and westward to South Dakota (Matthews 1961). In Maryland, this species is restricted to the western shore, west of the Chesapeake Bay. Staines (1984) reports it collected during May. Copris minutus (Drury, 1770) Scarabaeus minutus Drury 1770:78 Scarabaeus silenus Fabricius 1775:21 Scarabaeus ammon Fabricius 1781:24 Scarabaeus lar Fabricius 1787:13 Copris reflexus Panzer 1794:7 Length 8.0–13.0 mm. Body color black. Similar in appearance to C. fricator but smaller. Clypeus deeply notched at apex. Horn developed in both of the sexes, but

Figures 33–34. Pronotum: (33) Copris minutus, (34) Copris fricator. 338

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longer in male majors, sometimes tuberculate to stout in females. Head moderate to densely punctate. Male majors with median pronotal hump at apex, between 2 small tubercles. Elytral intervals slightly convex and smooth with very fine punctures, punctures may be obsolete in some specimens. Elytral striae deep and punctate. Copris minutis occurs in the eastern United States from New Hampshire south to Florida, westward to northern Texas and Nebraska (Ratcliffe and Paulsen 2008). In Maryland, this species is found across the state from March to December. It has been taken from dung, carrion, and rotting fruit and has been collected at lights (Price et al. 2012, Staines 1984). Tribe ATEUCHINI Laporte, 1840 Genus Ateuchus Weber, 1801 Two species of Ateuchus are known to occur in Maryland. These species are easily distinguished from other Maryland Scarabaeinae by the inner apical angle of the protibia (Fig. 7), which is set at a right angle. Both species are similar in size and shape, but easily distinguished by the shape of the pronotum (Fig. 35–36). Ateuchus lecontei (Harold) is shown in Figure 37. Key to Species of the Genus Ateuchus 1. Posterolateral angles of pronotum below the level of elytra (lateral view) (Fig. 36). ........................................................................ A. histeroides (Weber) Posterolateral angles of pronotum level with elytra (lateral view) (Fig. 37)........... .......................................................................................... A. lecontei (Harold) Ateuchus histeroides Weber, 1801 Ateuchus histeroides Weber 1801:37 Ateuchus capistratus Fabricius 1801:62 Length 5.5–7.3 mm. Color dark reddish brown to purplish black. Clypeus emarginate or bidentate at apex. Head punctate, punctures vary in size from small at base to large at apex. Pronotum punctate, punctures larger at base; some specimens with

Figures 35–36. Lateral view of pronotum and elytra: (36) Ateuchus histeroides, (37) Ateuchus lecontei. 339

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a shallow medial line extending from the base to middle. Hind angles of pronotum formed below level of elytral margin (side view). Six elytral striae between suture and humerus, impressed and with large punctures. Elytral intervals alutaceous and moderately punctate with very small punctures. Ateuchus histeroides occurs in the eastern United States (Ratcliffe and Paulsen 2008). In Maryland, this species occurs across the state from May to September. It is most abundant in wooded areas and has been taken from dung, fungi, and dead fish and has been collected at lights (Staines 1984). Ateuchus lecontei (Harold, 1868) Choeridium lecontei Harold 1868:52–53 Length 6.0–7.0 mm. Appearance black, or red to reddish brown to purplish black. Clypeus bidentate. Head and pronotum punctate, punctures larger at base; some specimens with a shallow median line extending from the base of pronotum to middle half. Very similar in appearance to A. histeroides; can be differentiated

Figure 37. Ateuchus lecontei. 340

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at lateral view (Fig. 35–36): hind angles of pronotum formed level with elytra. Elytral striae punctate. Elytral intervals alutaceous and moderately punctate with small punctures. Ateuchus lecontei occurs in the eastern coastal portions of the United States from New York to Mississippi (Harpootlian 2001). In Maryland, this species has only been collected from Worcester County, suggesting it may prefer pine forests and sandy habitats (Price et al. 2012, Staines 1984). Staines (1984) reports this species taken from cow dung and collected at lights. Summary This research is part of a larger project to survey Maryland scarabs (Coleoptera: Superfamily Scarabaeoidea) in all 23 counties. The superfamily Scarabaeoidea is a large, diverse, cosmopolitan group of beetles. Worldwide there are 35,000 species, with approximately 1500 species in North America (Ratcliffe and Paulsen 2008). Scarabaeoid beetles are prominent members of the entomofauna, and have been reported as important biocontrol agents, agricultural pests, and biological indicators. Other than the work of Staines (1984), Scarabaeinae research in the Mid-Atlantic has been limited to just a few studies that mainly focused on biodiversity (Price 2004, Price et al. 2012), food preference (Price 2006), and the perching behavior of Canthon viridis (Young 1982). With this research, we hope to greatly increase the knowledge of this beneficial group of beetles. Acknowledgments We thank the Guerrieri Undergraduate Summer Research Endowment and the Salisbury University Henson School of Science and Technology for funding this research. We thank Brett Ratcliffe (University of Nebraska, Lincoln, NE) for his review of this manuscript and Charlie Staines (National Museum of Natural History, Washington, DC) for his comments regarding the keys. We also thank the Northeastern Naturalist reviewers for their valuable comments and editing. Literature Cited Dejean. P.F.M.A. 1836. Catalogue des Coléoptères de la Collection de M. Le Comte Dejean. Troisieme Edition, Revue, Corrigee et Augmentee, Méquignon-Marvis, Paris, France. 384 pp. Downie, N.M., and R.H. Arnett Jr. 1996. The beetles of northeastern North America. Volume I: Introduction—suborders Archostemata, Adephaga, and Polyphaga, thru superfamily Cantharodea. The Sandhill Crane Press, Gainesville, FL. 880 pp. Drury, D. 1770–1782. Illustrations of Natural History, Volume 1 (1770): 130 pp., 50 plates; Volume 2 (1773): 90 pp., 50 plates; Volume 3 (1782-): 76 pp. 50 plates. B. White, London, UK. Edmonds, W.D. 1994. Revision of Phanaeus Macleay, a new world genus of scarabaeine dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Natural History Museum of Los Angeles County, Contributions in Science 443:1–105. Fabricius, J.C. 1775. Systema entomologiae. In Officina Libraria Kortii, Flensburgi et Lipsiae, Germany. 832 pp. 341

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Fabricius, J.C. 1781. Species insectorum, Volume 1. C.E. Bohnii, Hambugi et. Kilonii, Germany. 552 pp. Available online at http://www.biodiversitylibrary.org/ item/82398#page/1/mode/1up. Fabricius, J.C. 1787. Mantissa insectorum, Volume 1. C.G. Proft, Hafniae, Denmark. 348 pp. Available online at http://www.biodiversitylibrary.org/item/82321#page/5/ mode/1up. Fabricius, J.C. 1792. Entomologica systematica endemata et aucta. Secumdum Classes, Orines, Genera, Species adjectis synonimus, locis, observationibus, descriptionibus. Volume I Part II. C.G. Proft, Hafniae, Denmark. 538 pp. Available online at http://www. biodiversitylibrary.org/item/82436#page/3/mode/1up Fabricius, J.C. 1798. Supplementum Entomologiae Systematicae. C.G. Proft, Hafniae, Denmark. 572 pp. Fabricius, J.C. 1801. Systema Eleutheratorum, Volume 2. Bibliopolii Academici Novi, Kiliae, Germany. 687 pp. Fincher, G.T., and R.E. Woodruff. 1975. A European dung beetle, Onthophagus taurus Schreber, new to the US (Coleoptera: Scarabaeidae). Coleopterists Bulletin 29:349–350. Forster, J.R. 1771. Novae species insectorum. Centuria I. London, UK. 100 pp. Gemminger, M., and E. von Harold. 1869. Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus, Volume 4, Scarabaeidae. Monachii. Sumptu E.H. Gummi. Pp. 979–1346. Germar, E.F. 1824. Insectorum species novaeaut minus cognitae, descriptionibus illustratae. Halae. Hendel and Sons. 624 pp. Haldeman, S.S. 1843. Descriptions of North American species of Coleoptera, presumed to be undescribed. Proceedings of the Academy of Natural Sciences of Philadelphia 1:298–304. Haldeman, S.S., and J.E. LeConte. 1853. Catalogue of the described Coleoptera of the United States, by Friedrich Ernst Melsheimer, M.D., revised. Smithsonian Institution, Washington, DC. 174 pp. Halffter, G., and E.G. Matthews. 1966. The natural history of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae). Folia Entomologia Mexicana 12–14:1–312. Harold, E, von. 1868. Die Arten der Gattung Choeridium. Coleopterologische Hefte 4:32–76. Harold, E, von. 1869. Abanderungen vergebener Namen. Coleopterologische Hefte 5:122–125. Harold, E, von. 1871. Diagnosen neuer Coprophagen. Coleopterologishe Hefte 8:114–116. Harpootlian, P.J. 1995. Notes and records of Scarabaeidae from the southeastern United States. Coleopterists Bulletin 49:280. Harpootlian, P.J. 2001. Scarab Beetles (Coleoptera: Scarabaeidae) of South Carolina. Biota of South Carolina. Volume 2. Clemson University Public Service Publishing. Clemson, SC. Hoebeke, E.R., and K. Beuke. 1997. Adventive Onthophagus (Coleoptera: Scarabaeidae) in North America: Geographic ranges, diagnoses, and new distributional records. Entomological News 108(5):345–362. Horn, G.H. 1881. Descriptions of new species of North American Coleoptera. Transactions of the Kansas Academy of Science 7:74–77. Howden, H.F., and O.L. Cartwright. 1963. Scarab beetles of the genus Onthophagus Latreille North of Mexico (Coleoptera: Scarabaeidae). Proceedings of the United States National Museum 114(3467):1–135. 11 figures, 9 plates. 342

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Kirby, W. 1825. A description of such genera and species of insects, alluded to in the “Introduction to Entomology” of Messrs. Kirby and Spence, as appear not to have been before sufficiently noticed or described. Transactions of the Linnean Society of London 14:563–572. Kirby, W. 1837. The insects. Volume 4, In J. Richardson (Ed.). Fauna Boreali-Americana; or the Zoology of the Northern parts of British America. J. Fletcher, Norwich, UK. 325 pp. Laporte, F.L. 1840. Histoire naturelle des insectes Coléoptères. Avec une introduction Renferment l’Anatomie et la Physiologie des Animaux articulés, par M. Brullé, Volume 2. P. Duménil, Paris, France. 564 pp. LeConte, J.L. 1858. Catalogue of Coleoptera of the regions adjacent to the boundary line between the Unites States and Mexico. Journal of the Academy of Natural Sciences of Philadelphia (Series 2) 4:9–42. Linnaeus, C. 1758. Systemae Naturae, Editio Decima, Reformata. (Holmiae, fasc reprint British Museum of Natural history, London, UK 1939). 824 pp. Linnaeus, C. 1767. Systemae Naturae, Volume 1, Pars 2, Edito Duodecima Reformata. L. Salvii, Stockholm, Sweden. Pp. 533–1327. Losey, J.E., and M. Vaughan. 2006. The economic value of ecological services provided by insects. Bioscience 56:311–323. MacLeay, W.S. 1819. Horae Entomologicae: or Essays on the annulose animals, Volume 1. Part 1. S. Bagster, London, UK. 524 pp. MacRae, T.C., and S.R. Penn 2001. Additional records of adventive Onthophagus Latreille (Coleoptera: Scarabaeidae) in North America. Coleopterists Bulletin 55:49–50. Matthews, E.G. 1961. Revision of the genus Copris Müller of the western hemisphere (Coleoptera, Scarabaeidae). Entomologica Americana 41:1–139. Melsheimer, F.E. 1845. Descriptions of new species of Coleoptera of the United States. Proceedings of the Academy of Natural Sciences of Philadelphia 2:134–160. (volume for 1844 but published 1845 and cited as 1846 in Leng 1920). Melsheimer, F.E. 1846. Descriptions of new species of the United States. Proceedings of the Academy of Natural Sciences of Philadelphia 2:134–160. Miranda, C.B., J.D. Santos, and I. Bianchin. 2000. The role of Digitonthophagus gazella in pasture cleaning and production as a result of burial of cattle dung. Pasturas Tropicales, 22:14–18. Nichols, E., S. Spector, J. Louzada, T. Larsen, S. Amezquita, and M.E. Favila. 2008. Ecological functions and ecosystem services provided by Scarabaeinae dung beetles. Biological Conservation, 141:1461–1474. Olivier, A.G. 1789–1790. Entomologie, ou histoire naturelle des insects, avec leurs caractères génériques et spécifiques, leur description, leur synonymie, et leur figure enluminée. Coléoptères 1–2: (genera separately paged). Paris, France. Palisot de Beauvois, A.M.F.J. 1805. Insectes recuellis en Afrique et en Amérique, Vol. 3, pp. 25–40. Fain et Compagnie, Paris, France. Palisot de Beauvois, A.M.F.J. 1809. Insectes recuellis en Afrique et en Amérique, Vol. 6, pp. 89–100. Fain et Compagnie, Paris, France. Palisot de Beauvois, A.M.F.J. 1811. Insectes recuellis en Afrique et en Amérique, Vol. 7, pp. 101–120. Fain et Compagnie, Paris, France. Panzer, G.W.F. 1794. Faunae Insectorum americes borealis prodromus. Nürnberg, Germany. 8 pp. 1 plate. Peck, S.B., and M.C. Thomas. 1998. A distributional checklist of the beetles (Coleoptera) of Florida. Arthropods of Florida and Neighboring Land Areas 16:1–180. 343

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Price, D.L. 2004. Species diversity and seasonal abundance of scarabaeoid dung beetles (Coleoptera: Scarabaeidae, Geotrupidae and Trogidae) attracted to cow dung in Central New Jersey. Journal of the New York Entomological Society 112:334–347. Price, D.L. 2006. Notes on the scarabaeoid dung beetles (Coleoptera: Scarabaeidae, Geotrupidae, and Trogidae) of Hutcheson Memorial Forest, New Jersey, USA. Entomological news 117:347–350. Price, D.L., L.M. Brenneman, and R.E. Johnston. 2012. Dung Beetle (Coleoptera: Scarabaeidae and Geotrupidae) Communities of Eastern Maryland. Proceedings of the Entomological Society of Washington 114:142–151. Ratcliffe, B.C., and M.J. Paulsen. 2008. The scarabaeoid beetles of Nebraska (Coleoptera: Scarabaeoidea). University of Nebraska State Museum Bulletin 22:1–570. Robinson, M. 1941. Notes on some rare Scarabaeidae with the description of one new species (Coleoptera). Entomological News 52:227–232. Say, T. 1823. Descriptions of ceoleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long. Journal of the Academy of Natural Sciences of Philadelphia. 3:139–216. Say, T. 1835. Descriptions of new North American coleopterous insects, and observations on some already described. Boston Journal of Natural History 1:151–203. Schreber, J.C.D. 1759. Novae species insectorvm. Cvm figuris aeneis coloribvs pictis. Halae Magdebvrgicae. Schneider. Pp. 1–20. Staines, C.L. 1984. An annotated checklist of the Scarabaeoidea (Coleoptera) of Maryland. Maryland Entomologist 2:79–89. Sturm, J. 1826. Catalog Meiner Insecten-Sammlung. Erster Theil. Käfer J. Sturm, Nürnberg, Germany. 207 pp + 4 plates. Sturm, J. 1843. Catalog der Käfer-Sammlung von Jacob Sturm. Nürnberg, Germany. 386 pp. Weber, F. 1801. Observaciones entomologicae, continents novorum quae condidit generum characters, et nuper detectarum specierum descriptions. Kiliae. 116 pp. Woodruff, R.M. 1973. The scarab beetles of Florida (Coleoptera: Scarabaeidae). Part I. The Laprosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae, Acanthocerinae). Arthropods of Florida and Neighboring Land Areas. Volume 8:1–220. Young, O.P. 1982. Perching behavior of Canthon viridis (Coleoptera: Scarabaeidae) in Maryland. Journal of the New York Entomological Society 90:161–165. Ziegler, D. 1844. Descriptions of new North American Coleoptera. Proceedings of the Academy of Natural Sciences of Philadelphia 2:43–53.

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