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22001155 NORTHEASTERN NATURALIST 2V2(o2l). :2321,8 N–3o4. 42
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.
1Department of Biological Sciences, Salisbury University, 1101 Camden Avenue, Salisbury,
MD, 21801. *Corresponding author - dlprice@salisbury.edu.
Manuscript Editor: Christopher M. Heckscher
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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 n amed.
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 co mbined (Fig. 3) .
..............................................................................................ONTHOPHAGINI
Metatarsus with basal segment shorter than next 3 segments comb ined (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.
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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.
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Key to species of the genus Onthophagus in Maryland
1. Pronotum distinctly tuberculate (Fig. 9) or centrally puncta te 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, forw ard-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 biden tate (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 (F ig. 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 sp ots. Males with
median horn on base of head between or just behind eyes (Fig. 1 6); 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 horn s, 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-y ellowish
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 swelli ng 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.
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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 pu nctures.
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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).
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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,
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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
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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.
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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.
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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 gran -
ules (Fig. 27).....................................................................................................4
Pronotum scabrous (irregularly roughened) (Fig. 28); pygidium u niformly 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 (F ig. 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.
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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.
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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
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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.
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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.
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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 vi ew) (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.
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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.
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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.
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