Quick Guide for the Identification of

Maryland Scarabaeoidea

Mallory Hagadorn Dr. Dana L. Price

Department of Biological Sciences Salisbury University

This document is a pictorial reference of Maryland Scarabaeoidea genera (and sometimes species) that was created to expedite the identification of Maryland Scarabs. Our current understanding of Maryland Scarabs comes from “An Annotated Checklist of the Scarabaeoidea (Coleoptera) of Maryland” (Staines 1984). Staines reported 266 species and subspecies using literature and review of several Maryland Museums. Dr. Price and her research students are currently conducting a bioinventory of Maryland Scarabs that will be used to create a “Taxonomic Guide to the Scarabaeoidea of Maryland”. This will include dichotomous keys to family and species based on historical reports and collections from all 23 counties in Maryland.

This document should be cited as: Hagadorn, M.A. and D.L. Price. 2012. Quick Guide for the Identification of Maryland Scarabaeoidea. Salisbury University. Pp. 54.

Questions regarding this document should be sent to: Dr. Dana L. Price - [email protected]

*All photographs are linked to their copyright holder. 1

Table of Contents

Families of Scarabaeoidea of Maryland……………………………………...... 6

Geotrupidae……………………………………………………………………………...... 7 Subfamily Bolboceratinae…………………………………………………………… 7 Genus Bolbocerosoma…………………………………………………...... 7 Genus Eucanthus……………………………………………………...... 7 Subfamily Geotrupinae……………………………………………………………… 8 Genus Geotrupes………………………………………………………...... 8 Genus Odonteus...………………………………………………………….. 9

Glaphyridae...... 9 Genus Lichnanthe.…………………………………………………………... 9

Hybosoridae………………………………………………………………………………… 10 Genus Germarostes…………………………………………………………. 10

Lucanidae…………………………………………………………………………………… 10 Subfamily Aesalinae……………………………………………………………...... 10 Genus Nicagus………………………………………………………………. 10 Subfamily Luncaninae………………………………………………………………. 10 Genus Dorcus…………………………………………………...... 10 Genus Lucanus…………………………………………………………...... 11 Genus Platycerus……………………………………………………………. 12 Subfamily Syndesinae………………………………………………………………. 12 Genus Ceruchus…………………………………………………………….. 12

Scarabaeidae……………………………………………………………………………….. 13 Subfamily Aphodiinae……………………………………………………………….. 13 Tribe Aegialiini…………………………………………………………………… 13 Genus Aegialia……………………………………………………………….. . 13 Genus Caelius………………………………………………...... 13

*All photographs are linked to their copyright holder. 2

Tribe Aphodiini………………………………………………………………...... 14 Genus Acrossus………………………………………………...... 14 Genus Agoliinus………………………………………………...... 14 Genus Aphodius………………………………………………...... 15 Genus Blackburneus…………………………………………...... …… 15 Genus Blackburneus…………………………………………...... 15 Genus Calamosternus………………………………………………………. 15 Genus Chilothorax…………………………………………………………… 16 Genus Colobopterus………………………………………………………… 16 Genus Dialytellus…………………………………………………………….. 16 Genus Dialytes……………………………………………………………….. 17 Genus Flaviellus……………………………………………………………... 17 Genus Irrasinus………………………………………………………………. 17 Genus Labarrus……………………………………………………………… 18 Genus Lechorodius………………………………………………………….. 18 Genus Melinopterus…………………………………………………………. 18 Genus Oscarinus……………………………………………...... 19 Genus Otophorus……………………………………………………………. 19 Genus Oxyomus……………………………………………………………... 19 Genus Pardalosus…………………………………………………………… 20 Genus Planolinellus…………………………………………………………. 20 Genus Planolinus……………………………………………………………. 20 Genus Pseudagolius………………………………………...... 21 Genus Stenotothorax………………………………………...... 21 Genus Strigodius…………………………………………………………….. 21 Genus Teuchestes…………………………………………...... 22 Tribe Didactyliini…………………………………………………………………. 22 Genus Aidophus……………………………………………………………... 22 Tribe Eupariini……………………………………………………………………. 22 Genus Aphotaenius………………………………………………………….. 22 Genus Ataenius……………………………………………………………… 23 Genus Pseudataenius………………………………………………………. 23 Genus Saprosites……………………………………………………………. 24 Tribe Psammodiini……………………………………………………………….. 24 Genus Pammodius…………………………………………...... 24 Genus Rhyssemus…………………………………………...... 25 Genus Pleurophorus………………………………………………………… 25 Subfamily Cetoniinae……………………………………………………………….. 26 Tribe Cetoniini…………………………………………………………………… 26 Genus Euphoria……………………………………………………………… 26

*All photographs are linked to their copyright holder. 3

Tribe Cremastocheilini…………………………………………………………... 28 Genus Cremastocheilus…………………………………………………….. 28 Genus Psilocnemis………………………………………………………….. 28 Tribe Gymnetini………………………………………………………………….. 29 Genus Cotinis………………………………………………...... 29 Tribe Trichiini……………………………………………………………………... 29 Genus Gnorimella……………………………………………………………. 29 Genus Osmoderma………………………………………………………….. 29 Genus Trichiotinus…………………………………………………………… 30 Genus Trigonopeltastes…………………………………………………….. 31 Tribe Valgini………………………………………………………………………. 31 Genus Valgus………………………………………………...... 31 Subfamily Dynastinae………………………………………………...... 31 Tribe Cyclocephalini……………………………………………...... 31 Genus Cyclocephala………………………………………………………… 31 Genus Dyscinetus…………………………………………………………… 32 Tribe Dynastini…………………………………………………………………… 32 Genus Dynastes……………………………………………………………... 32 Tribe Oryctini……………………………………………………………………... 32 Genus Strategus…………………………………………………………….. 32 Genus Xyloryctes……………………………………………………………. 33 Tribe Pentodontini………………………………………………...... 33 Genus Aphonus……………………………………………………………… 33 Genus Euetheola…………………………………………………………….. 34 Genus Tomarus……………………………………………………………… 34 Tribe Phileurini…………………………………………………………………… 35 Genus Phileurus……………………………………………………………... 35 Subfamily Melolonthinae……………………………………………………………. 35 Tribe Diplotaxini…………………………………………………...... 35 Genus Diplotaxis……………………………………………...... 35 Tribe Hopliini……………………………………………………………………… 36 Genus Hoplia…………………………………………………………………. 36 Tribe Macrodactylini……………………………………………………………... 37 Genus Dichelonyx…………………………………………………………… 37 Genus Macrodactylus………………………………………...... 38 Tribe Melolonthini………………………………………………………………... 38 Genus Phyllophaga…………………………………………...... 38 Genus Polyphylla…………………………………………………………….. …………….. 40 Tribe Sericini……………………………………………………………………… 41 Genus Maladera……………………………………………………………... 41

*All photographs are linked to their copyright holder. 4

Genus Nipponoserica……………………………………………………….. 41 Genus Serica…………………………………………………………………. 41 Subfamily Rutelinae…………………………………………………...... 42 Tribe Anomalini...... 42 Genus Anomala……………………………………………………………… 42 Genus Popillia………………………………………………………………... 44 Genus Strigoderma…………………………………………...... 45 Tribe Rutelini……………………………………………………………………... 45 Genus Cotalpa……………………………………………………………….. 45 Genus Parastasia……………………………………………………………. 45 Genus Pelidnota……………………………………………………………... 46 Subfamily Scarabaeinae……………………………………………...... 46 Tribe Ateuchini…………………………………………………………………… 46 Genus Ateuchus……………………………………………...... 46 Tribe Canthonini…………………………………………………………………. 46 Genus Canthon………………………………………………...... 46 Genus Melanocanthon………………………………………………………. 47 Tribe Coprini……………………………………………………………………… 47 Genus Copris………………………………………………………………… 47 Genus Dichotomius…………………………………………...... 48 Tribe Onthophagini………………………………………………………………. 48 Genus Onthophagus………………………………………………………… 48 Tribe Phanaeini…………………………………………………………………... 49 Genus Phanaeus……………………………………………...... 49

Trogidae……………………………………………………………………………………... 50 Subfamily Troginae………………………………………………………………….. 50 Genus Omorgus……………………………………………………………… 50 Genus Trox…………………………………………………………………… 51

References………………………………………………………………………………….. 53

*All photographs are linked to their copyright holder. 5

Table of Contents Families of the Scarabaeoidea of Maryland Geotrupidae Glaphyridae Hybosoridae

Geotrupes hornii Lichnanthe vulpina Germarostes globosus

Lucanidae Passalidae

Lucanus capreolus Odontotaenius disjunctus

Scarabaeidae Trogidae

Onthophagus taurus Trox spinulosus

*All photographs are linked to their copyright holder. 6

Table of Contents GEOTRUPIDAE Subfamily Bolboceratinae Genus: Bolbocerosoma Species: Bolbocerosoma farctum Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Bolboceratini Page 51

http://bugguide.net/node/view/255129/bgimage

Genus: Bolbocerosoma Species: Bolbocerosoma tumefactum Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Bolboceratini

Page 51

http://bugguide.net/node/view/429013/bgimage

Genus: Eucanthus Species: Eucanthus lazarus Similar to: E. impressus Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Bolboceratini

Page 51; Skip to 9(5)

http://bugguide.net/node/view/345282/bgimage

*All photographs are linked to their copyright holder. 7

Table of Contents Genus: Eucanthus Species: Eucanthus subtropicus Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Bolboceratini Page 51; Skip to 9(5)

http://bugguide.net/node/view/209128/bgimage

Subfamily Geotrupinae Genus: Geotrupes

Species: Geotrupes splendidus Similar to: G. balyi, G. semiopacus Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Geotrupini Page 56; Skip to 2(1)

http://bugguide.net/node/view/402659/bgimage

Genus: Geotrupes Species: Geotrupes blackburnii Similar to: G. egeriei

Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Geotrupini Page 56; Skip to 2(1)

http://bugguide.net/node/view/358581/bgimage

*All photographs are linked to their copyright holder. 8

Table of Contents Genus: Geotrupes Species: Geotrupes hornii Similar to: G. ulkei (Cited in Literature) Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Geotrupini Page 56; Skip to 2(1)

http://bugguide.net/node/view/284669/bgimage

Genus: Odonteus (Bolboceras) Species: Odonteus (Bolboceras) liebecki

Similar to: O. cornigerus (Cited in Literature) O. darlingtoni, O. falli, O. thoracicornis Dichotomous Key: Scarab Beetles of South Carolina

Key to Tribe Bolboceratini, Page 51; skip to 2(1) O. falli-The Scarabaeoid Beetles of Nebraska Key to Odonteus, Page 38

http://bugguide.net/node/view/19076/bgimage

GLAPHYRIDAE Genus: Lichnanthe Species: Lichnanthe vulpina Dichotomous Key: Species description in Scarab Beetles of South Carolina, Page 58

http://bugguide.net/node/view/547246/bgimage

*All photographs are linked to their copyright holder. 9

Table of Contents HYBOSORIDAE Subfamily Ceratocanthinae Genus: Germarostes (Cloetous) Species: Germarostes (Cloetous) globosus Similar to: Germarostes (Cloetous) aphodioides Dichotomous Key: The Scarabaeoid Beetles of Nebraska, Key to Germarostes, Page 130

http://bugguide.net/node/view/273801/bgimage

http://bugguide.net/node/view/400347/bgimage

LUCANIDAE Subfamily Aesalinae Genus: Nicagus Species: Nicagus obscurus Dichotomous Key: For Species Description see the link below

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoi dea/Lucanidae/AES/NIC/obscurus.html

Subfamily Lucaninae

Genus: Dorcus Species: Dorcus brevis (Cited in Literature) Dichotomous Key: For Species Description see the link below

http://www-

museum.unl.edu/research/entomology/Guide/Scarabaeoi dea/Lucanidae/LUC/DOR/brevis.html

http://bugguide.net/node/view/60176

*All photographs are linked to their copyright holder. 10

Table of Contents Genus: Dorcus Species: Dorcus parallelus Dichotomous Key: Species Description in The Scarabaeoid

Beetles of Nebraska, Page 108

http://bugguide.net/node/view/434738/bgimage

Genus: Lucanus (Pseudolucanus)

Species: Lucanus capreolus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Lucanus, Page 110

http://www.flickr.com/photos/pcoin/5914932906/in/set-

72057594074593321

Genus: Lucanus Species: Lucanus elaphus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Lucanus, Page 110

http://www.flickr.com/photos/entomopixel/5098352100/

*All photographs are linked to their copyright holder. 11

Table of Contents Genus: Platycerus Species: Platycerus virescens Similar to: Platycerus piceus Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 106

http://bugguide.net/node/view/390659/bgimage

Subfamily Syndesinae

Genus: Ceruchus Species: Ceruchus piceus Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 105

http://bugguide.net/node/view/278893/bgimage

PASSALIDAE Genus: Odontotaenius (Popilius) Species: Odontotaenius disjunctus Dichotomous Key: Species Description in The Scarabaeoid

Beetles of Nebraska, Page 25

http://bugguide.net/node/view/585405/bgimage

*All photographs are linked to their copyright holder. 12

Table of Contents SCARABAEIDAE Subfamily Aphodiinae Tribe: Aegialiini Genus: Aegialia Species: picture just representative of genus

Similar to: Aegialia blanchardi (cited in literature)

***Species found in Maryland*** Dichotomous Key: Scarab Beetles of South Carolina Key to Aegialia, Page 25

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoi dea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aegialiini /Aegialia/Aegialia.html

Genus: Caelius (Aegialia) Species: picture just representative of genus Similar to: Caelius humeralis (Aegialia humeralis) (Cited in the Literature)

***Species found in Maryland*** Dichotomous Key: Scarab Beetles of South Carolina Key to Aegialia, Page 25

http://www-

museum.unl.edu/research/entomology/Guide/Scarabaeoi dea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aegialiini /Caelius/Caelius.html

*All photographs are linked to their copyright holder. 13

Table of Contents Tribe Aphodiini Genus: Acrossus (Aphodius) Species: Acrossus rubripennis (Aphodius rubripennis) Dichotomous Key: Aphodiini of the United States and Canada, Key to Acrossus, page 254

http://bugguide.net/node/view/110525

Genus: Acrossus (Aphodius) Species: Acrossus rufipes (Aphodius rufipes) Dichotomous Key: Aphodiini of the United States and Canada, Key to Acrossus, page 254

http://typhaeus.chez-alice.fr/galaph.htm

Genus: Agoliinus (Aphodius) Species: picture just representative of genus Similar to: Agoliinus leopardus (Aphodius leopardus) Agoliinus manitobensis (A. manitobensis) ***Species found in Maryland*** Dichotomous Key: Aphodiini of the United States and Canada, Key to Agoliinus, Page 125

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/Scara baeidae/Aphodiinae/AphodiinaeTribes/Aphodiini/Agoliinus/Agoliinus .html

*All photographs are linked to their copyright holder. 14

Table of Contents Genus: Aphodius Species: Aphodius fimetarius Dichotomous Key: Aphodiini of the United States and Canada, Page 106

The Scarabaeoid Beetles of Nebraska, Key to Aphodius, Page 147

http://bugguide.net/node/view/355100

Genus: Blackburneus (Aphodius) Species: Blackburneus aegrotus (A. campestris) Similar to: Blackburneus rubeolus (A. rubeolus) B. lentus (A. lentus) B. stercorosus (A. stercorosus) Dichotomous Key: Aphodiini of the United States and Canada, Key to Blackburneus, page 351

The Scarabaeoid Beetles of Nebraska, Key to Aphodius, Page 147

http://bugguide.net/node/view/496691/bgpage

Genus: Calamosternus (Aphodius) Species: Calamosternus granarius (A. granarius) Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 258

The Scarabaeoid Beetles of Nebraska, Key to Aphodius, Page 147

http://bugguide.net/node/view/390683/bgimage

*All photographs are linked to their copyright holder. 15

Table of Contents

Genus: Chilothorax (Aphodius) Species: Chilothorax distinctus (Aphodius distinctus) Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 258

The Scarabaeoid Beetles of Nebraska, Key to Aphodius, Page 147

http://bugguide.net/node/view/436765/bgimage

Genus: Colobopterus (Aphodius) Species: Colobopterus erraticus (Aphodius erraticus) Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 92

The Scarabaeoid Beetles of Nebraska, Key to Aphodius, Page 147 http://museum.unl.edu/research/entomology/Guide/Scarabae oidea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aphodiini/ Colobopterus/Colobopterus.html

Genus: Dialytellus (moved from Tribe Eupariini) Species: Dialytellus dialytoides Similar to: Dialytellus tragicus (D. humeralis) Dichotomous Key: Aphodiini of the United States and Canada, Key to Dialytellus, Page 78

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/ Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aphodiini/Dialyte llus/Dialytellus.html

*All photographs are linked to their copyright holder. 16

Table of Contents Genus: Dialytes (moved from Tribe Eupariini) Species: Dialytes truncatus Similar to: Dialytes striatulus, D. ulkei Dichotomous Key: Aphodiini of the United States and Canada, Key to Dialytes, Page 82

http://bugguide.net/node/view/565984/bgimage

Genus: Flaviellus (Aphodius) Species: Flaviellus phalerioides (A. phaleroides) Similar to: Dialytes striatulus, D. ulkei Dichotomous Key: Aphodiini of the United States and Canada, Key to Flaviellus, Page 335

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoi dea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aphodiin i/Flaviellus/Flaviellus.html

Genus: Irrasinus (Aphodius) Species: Irrasinus stupidus (Aphodius stupidus) Dichotomous Key: Aphodiini of the United States and Canada, Key to Irrasinus, Page 368

http://bugguide.net/node/view/369547/bgimage

*All photographs are linked to their copyright holder. 17

Table of Contents Genus: Labarrus (Aphodius) Species: Labarrus pseudolividus (Aphodius lividus) Dichotomous Key: Aphodiini of the United States and Canada, Key to Labarrus, Page 261

http://museum.unl.edu/research/entomology/SSSA/SS SA-Taxa/Aphodiinae/Aphodius/pseudolividus.html

Genus: Lechorodius (Aphodius) Species: picture just representative for genus Similar to: Lechorodius terminalis (A. terminalis) Lechorodius lutulentus (A. lutulentus) ***Species found in Maryland*** Dichotomous Key: Aphodiini of the United States and Canada, Key to Lechorodius, Page 371

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoi dea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aphodiin i/Lechorodius/Lechorodius.html

Genus: Melinopterus (Aphodius) Species: Melinopterus femoralis (Aphodius femoralis)

Similar to: Melinopterus prodromus (A. prodromus)

Dichotomous Key: Aphodiini of the United States and Canada, Key to Melinopterus, Page 320

http://bugguide.net/node/view/496735

*All photographs are linked to their copyright holder. 18

Table of Contents Genus: Oscarinus (Aphodius) Species: picture just representative of genus Similar to: Oscarinus silvanicus (Aphodius silvanicus) Oscarinus rusicola (A. ruricola) ***Species found in Maryland***

Dichotomous Key: Aphodiini of the United States and Canada, Key to Oscarinus, Page 165

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoi dea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aphodii ni/Oscarinus/Oscarinus.html

Genus: Otophorus (Aphodius) Species: Otophorus haemorrihoidalis (A. haemorrihoidalis)

Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 89

The Scarabaeoid Beetles of Nebraska Key to Aphodius, Page 147

http://bugguide.net/node/view/496747

Genus: Oxyomus Species: Oxyomus sylvestris (O. silvestris) (Cited in Literature) Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 87 http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/S carabaeidae/Aphodiinae/AphodiinaeTribes/Aphodiini/Oxyomus/ Oxyomus.html

*All photographs are linked to their copyright holder. 19

Table of Contents Genus: Pardalosus (Aphodius) Species: picture just representative for the genus Similar to: Pardalosus serval (Aphodius serval) ***Species found in Maryland*** Dichotomous Key: Aphodiini of the United States and Canada, Key to Pardalosus, Page 377

http://bugguide.net/node/view/494555

Genus: Planolinellus (Aphodius) Species: Planolinellus vittatus (Aphodius vittatus)

Dichotomous Key: Species Description in Aphodiini of the

United States and Canada, Page 122

The Scarabaeoid Beetles of Nebraska Key to Aphodius, Page 147

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/Scarab aeidae/Aphodiinae/AphodiinaeTribes/Aphodiini/Planolinellus/Planolin ellus.html

Genus: Planolinus (Aphodius)

Species: Planolinus tenellus (Aphodius tenellus) (Cited in Literature) Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 117

The Scarabaeoid Beetles of Nebraska Key to Aphodius, Page 147

http://bugguide.net/node/view/496753/bgimage

*All photographs are linked to their copyright holder. 20

Table of Contents

Genus: Pseudagolius (Aphodius) Species: Pseudagolius bicolor (Aphodius bicolor)

Dichotomous Key: Aphodiini of the United States and Canada, Key to Pseudagolius, Page 311

http://bugguide.net/node/view/262887/bgimage http://bugguide.net/node/view/245071/bgimage

Genus: Stenotothorax (Aphodius) Species: Stenotothorax badipes (A. badipes)

Dichotomous Key: Aphodiini of the United States and Canada, Key to Stenotothorax, Page 189

http://www - museum.unl.edu/research/entomology/Guide/Scarabaeoide a/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Aphodiini/Ste notothorax/Stenotothorax.html

Genus: Strigodius (Aphodius) Species: Strigodius robinsoni (A. robinsoni) Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 237

The Scarabaeoid Beetles of Nebraska Key to Aphodius, Page 147 http://www-

museum.unl.edu/research/entomology/Guide/Scarabaeoidea/Sca rabaeidae/Aphodiinae/AphodiinaeTribes/Aphodiini/Strigodius/Stri godius.html

*All photographs are linked to their copyright holder. 21

Table of Contents

Genus: Teuchestes (Aphodius) Species: Teuchestes fossor (Aphodius fossor)

Dichotomous Key: Species Description in Aphodiini of the United States and Canada, Page 94

The Scarabaeoid Beetles of Nebraska Key to Aphodius, Page 147 http://www.galerie- insecte.org/galerie/view.php?adr=image%2Fdos6%2Ftemp %2Fteuchestes_fosso.jpg

Tribe Didactyliini Genus: Aidophus (Aphodius) Species: Aidophus parcus (A. parcus) (Cited in Literature)

Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 225

http://bugguide.net/node/view/531667

Tribe Eupariini Genus: Aphotaenius (moved from Tribe Psammodiini) Species: Aphotaenius carolinus Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 227

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/ Scarabaeidae/Aphodiinae/AphodiinaeTribes/Eupariini/Aphota enius/Aphotaenius.html

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Table of Contents

Genus: Ataenius Species: Ataenius strigatus Similar to: Ataenius apicalis, A. brevis, A. exiguous, A. fattigi, A. glaseri, A. imbricatus, A. insculptus (cited in literature), A. ovatulus, A. spretulus, A. wenzelii ***Black/Really Dark Brown color***

Dichotomous Key: The Scarabaeoid Beetles of Nebraska, Key to Ataenius, Page 229

http://bugguide.net/node/view/306014/bgimage

Genus: Ataenius Species: Ataenius gracilis Similar to: Ataenius alternates (cited in literature), A. abditus, A. cylindrus, A. glaseri, A. miamii ***Lighter Brown Color*** Dichotomous Key: The Scarabaeoid Beetles of Nebraska, Key to Ataenius, Page 229

http://bugguide.net/node/view/498543/bgimage

-Species A. figulator and A. simulator found in Staines but not in current taxonomic classification-

Genus: Pseudataenius (Pseudoataenius) Species : picture just representative of genus Similar to: Pseudataenius contorus (cited in literature), P. walterhorni (cited in literature) ***Species found in Maryland***

Dichotomous Key: Species Description in Scarab Beetles of South Carolina, Page 36

http://bugguide.net/node/view/496778/bgimage

*All photographs are linked to their copyright holder. 23

Table of Contents Genus: Saprosites (Moved from Tribe Psammodiini) Species : pictures just representative for genus Similar to: Saprosites ventralis ***Species found in Maryland*** Dichotomous Key: Species Description in Scarab Beetles of South Carolina, Page 37

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/ Scarabaeidae/Aphodiinae/AphodiinaeTribes/Eupariini/Sapros ites/Saprosites.html

Tribe Psammodiini Genus: Psammodius Species : Psammodius basalis Dichotomous Key: For genus description see the link below Species Description in Scarab Beetles of South Carolina, Page 37

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoid ea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Psammodii ni/Psammodius/Psammodius.html

Genus: Psammodius Species : Psammodius pierotti (P. sulcicollis) (cited in literature) Dichotomous Key: For genus description see the link below

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoid ea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Psammodii ni/Psammodius/Psammodius.html

http://coleoptera.ksib.pl/kfp/search.php?img=6954

*All photographs are linked to their copyright holder. 24

Table of Contents Genus: Psammodius Species : Psammodius laevipennis Dichotomous Key: For genus description see the link below

http://www-

museum.unl.edu/research/entomology/Guide/Scarabaeoid ea/Scarabaeidae/Aphodiinae/AphodiinaeTribes/Psammodii ni/Psammodius/Psammodius.html

http://www.entomologiitaliani.net/public/forum/phpBB3/viewtopi c.php?t=29232 -Species P.bidens and P. interruptus found in Staines but not in current taxonomic classification-

Genus: Rhyssemus Species : pictures just representative for genus Similar to: Rhyssemus scaber ***Species found in Maryland*** Dichotomous Key: Scarab Beetles of South Carolina, Key to Species of Psammodiini, Page 44

http://bugguide.net/node/view/357036

Genus: Pleurophorus Species : Pleurophorus caesus

Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 244 -Link below for additional species description

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/ Scarabaeidae/Aphodiinae/AphodiinaeTribes/Psammodiini/Ple urophorus/Pleurophorus.html

*All photographs are linked to their copyright holder. 25

Table of Contents Genus: Pleurophorus Species : Pleurophorus atlanticus (cited in literature)

Dichotomous Key: Genus description in The Scarabaeoid

Beetles of Nebraska, Page 244

http://unibio.unam.mx/irekani/handle/123456789/25622?proyec to=Irekani

Subfamily Cetoniinae Tribe Cetoniini Genus: Euphoria Species: Euphoria areata (Stephanucha areata) Dichotomous Key: Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Centoniinae), Key to areata Species-Group, Page 98

Species Description in Scarab Beetles of South Carolina, Page 116

http://bugguide.net/node/view/57375

Genus: Euphoria

Species: Euphoria flugida Dichotomous Key: Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Centoniinae), Key to flugida Species-Group, Page 41

The Scarabaeoid Beetles of Nebraska Key to Euphoria, Page 469

http://bugguide.net/node/view/28508/bgimage

*All photographs are linked to their copyright holder. 26

Table of Contents Genus: Euphoria Species: Euphoria herbacea Dichotomous Key: Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Centoniinae), Key to herbacea Species-Group, Page 47

Scarab Beetles of South Carolina Key to Euphoria, Page 117

http://bugguide.net/node/view/24068/bgimage

Genus: Euphoria Species: Euphoria inda

Dichotomous Key: Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Centoniinae), Key to inda Species-Group, Page 92

Scarab Beetles of South Carolina Key to Euphoria, Page 117

The Scarabaeoid Beetles of Nebraska Key to Euphoria, Page 469

http://bugguide.net/node/view/634709/bgimage

Genus: Euphoria Species: Euphoria sepulchralis Dichotomous Key: Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Centoniinae, Key to sepulchralis Species-Group, Pg. 92

Scarab Beetles of South Carolina Key to Euphoria, Page 117

The Scarabaeoid Beetles of Nebraska Key to Euphoria, Page 469

http://bugguide.net/node/view/152750/bgimage

*All photographs are linked to their copyright holder. 27

Table of Contents Tribe Cremastocheilini Genus: Cremastocheilus Species: Cremastocheilus canaliculatus Similar to: Cremastocheilus castaneae, C. variolosus Dichotomous Key: Scarab Beetles of South Carolina Key to Species of Cremastocheilini , Page 118

http://bugguide.net/node/view/641649/bgimage

Genus: Cremastocheilus Species: Cremastocheilus harrisii Dichotomous Key: Scarab Beetles of South Carolina Key to Species of Cremastocheilini , Page 118

http://bugguide.net/node/view/360964/bgimage

Genus: Psilocnemis Species: Psilocnemis leucosticte (cite in literature)

Dichotomous Key: Scarab Beetles of South Carolina

Key to Species of Cremastocheilini, Page 118

http://bugguide.net/node/view/360964/bgimage

*All photographs are linked to their copyright holder. 28

Table of Contents Tribe Gymnetini Genus: Cotinis Species: Cotinis nitida

Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 466

http://bugguide.net/node/view/44539/bgimage

Tribe Trichiini Genus: Gnorimella Species: Gnorimella maculosa Dichotomous Key: Species Description in The Scarab Beetles of South Carolina, Page 122

http://www.chebucto.ns.ca/Environment/NHR/Gnorimella_ma culosa.html

Genus: Osmoderma Species: Osmoderma eremicola

Similar to: Osmoderma scabra Dichotomous Key: Scarab Beetles of South Carolina Key to Osmoderma, Page 121

http://bugguide.net/node/view/47113

*All photographs are linked to their copyright holder. 29

Table of Contents Genus: Trichiotinus Species: Trichiotinus affinis Similar to: Trichiotinus piger Dichotomous Key: Scarab Beetles of South Carolina Key to Trichiotinus, Page 122

http://bugguide.net/node/view/534025/bgimage

Genus: Trichiotinus Species: Trichiotinus lunulatus Dichotomous Key: Scarab Beetles of South Carolina Key to Trichiotinus, Page 122

http://bugguide.net/node/view/57332

Genus: Trichiotinus Species: Trichiotinus bibens

Dichotomous Key: Scarab Beetles of South Carolina Key to Trichiotinus, Page 122

http://bugguide.net/node/view/448300/bgimage

*All photographs are linked to their copyright holder. 30

Table of Contents Genus: Trigonopeltastes Species: Trigonopeltastes delta Dichotomous Key: Species Description in Scarabs Beetles of South Carolina, Page 122

http://bugguide.net/node/view/277919/bgimage

Tribe Valgini Genus: Valgus Species: Valgus canaliculatus Similar to: Valgus squamiger Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Valgus, Page 502

http://bugguide.net/node/view/477528/bgimage http://bugguide.net/node/view/514769/bgimage

Subfamily Dynastinae Tribe Cyclocephalini

Genus: Cyclocephala Species: Cyclocephala borealis Similar to: Cyclocephala immaculata

Dichotomous Key: Scarab Beetles of South Carolina Key to Cyclocephala, Page106

http://bugguide.net/node/view/427813/bgimage

*All photographs are linked to their copyright holder. 31

Table of Contents Genus: Dyscinetus Species: Dyscinetus morator Similar to: Dyscinetus trachypygus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Dyscinetus, Page 443

http://bugguide.net/node/view/269765/bgimage

Tribe Dynastini Genus: Dynastes Species: Dynastes tityus Dichotomous Key: Species Description in Scarab Beetles of South Carolina, Page 113

http://bugguide.net/node/view/44808/bgimage http://bugguide.net/node/view/535056/bgimage

Tribe Oryctini Genus: Strategus Species: Strategus antaeus Dichotomous Key: Scarab Beetles from South Carolina Key to Strategus, Page 113

http://bugguide.net/node/view/24830 http://bugguide.net/node/view/73318

*All photographs are linked to their copyright holder. 32

Table of Contents Genus: Xyloryctes Species: Xyloryctes jamaicensis Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 460

Species Description in Scarab Beetles of South Carolina, Page 112

http://bugguide.net/node/view/318374/bgimage http://bugguide.net/node/view/318382/bgimage

Tribe Pentodontini

Genus: Aphonus (moved from Tribe Oryctini) Species: Aphonus castaneus Dichotomous Key: Scarab Beetles of South Carolina Key to Aphonus, Page 111

http://www.pbase.com/tmurray74/image/125304612

*All photographs are linked to their copyright holder. 33

Table of Contents Genus: Aphonus (moved from Tribe Oryctini) Species: Aphonus densicauda Dichotomous Key: Species Description in Scarab Beetles of South Carolina, Page 111

http://bugguide.net/node/view/454441/bgimage

Genus: Euetheola Species: picture just a representative for genus Similar to: Euetheola rugiceps ***Species found in Maryland*** Dichotomous Key: Species Description in Scarab Beetles of South Carolina, Page 112

http://bugguide.net/node/view/275139/bgimage

Genus: Tomarus (Ligyrus) (moved from Tribe Oryctini) Species: Tomarus gibbosus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Tomarus, Page 455

http://bugguide.net/node/view/431585/bgimage

* All photographs are linked to their copyright holder. 34

Table of Contents Genus: Tomarus (Ligyrus) (moved from Tribe Oryctini) Species: Tomarus gibbosus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Tomarus, Page 455

http://bugguide.net/node/view/559595 http://bugguide.net/node/view/893659

Tribe Phileurini Genus: Phileurus Species: Phileurus valgus (P. castaneus) Dichotomous Key: Scarab Beetles of South Carolina Key to Phileurus, Page 115

Species Description in The Scarabaeoid Beetles of Nebraska, Page 463

http://bugguide.net/node/view/285911/bgimage

Subfamily Melolonthinae Tribe Diplotaxini Genus: Diplotaxis

Species: Diplotaxis frondicola Similar to: Diplotaxis fulva, D. subcostata (cited in literature)

Dichotomous Key: Scarab Beetles of South Carolina Key to Diplotaxis, Page 73

http://bugguide.net/node/view/269786/bgimage

* All photographs are linked to their copyright holder. 35

Table of Contents Genus: Diplotaxis Species: Diplotaxis harperi Similar to: Diplotaxis atlantis, D. bidentata (cited in literature), D. liberta, D. punctatorugosa (cited in literature), D. sordid, D. tristis

Dichotomous Key: Scarab Beetles of South Carolina Key to Diplotaxis, Page 73

http://bugguide.net/node/view/449283/bgimage

Tribe Hopliini Genus: Hoplia Species: Hoplia trifasciata Similar to: Hoplia equina, H. modesta Dichotomous Key: Scarab Beetles of South Carolina Key to Hoplia, Page 96

http://bugguide.net/node/view/50872/bgimage

Genus: Hoplia Species: Hoplia trivialis Similar to: Hoplia equina Dichotomous Key: Scarab Beetles of South Carolina Key to Hoplia, Page 96

http://bugguide.net/node/view/263075/bgimage

*All photographs are linked to their copyright holder. 36

Table of Contents Tribe Macrodactylini

Genus: Dichelonyx Species: Dichelonyx albicollis Similar to: Dichelonyx fuscula Dichotomous Key: Scarab Beetles of South Carolina Key to Dichelonyx, Page 94

Dichelonyx albbicollis: Scarabaeoid Beetles of Nebraska, Key to Dichelonyx, Page 321 http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoidea/S carabaeidae/Melolonthinae/Melolonthinae-Generic- pages/Macrodactylini/Dichelonyx/Dichelonyx.html

Genus: Dichelonyx Species: Dichelonyx diluta Dichotomous Key: Scarab Beetles of South Carolina Key to Dichelonyx, Page 94

http://bugguide.net/node/view/405821/bgimage

Genus: Dichelonyx Species: Dichelonyx elongatula (Dichelonyx elongata) Similar to: Dichelonyx subvittata Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Dichelonyx, Page 321

http://bugguide.net/node/view/532932/bgimage

*All photographs are linked to their copyright holder. 37

Table of Contents Genus: Macrodactylus Species: Macrodactylus angustatus Dichotomous Key: Scarab Beetles of South Carolina Key to Macrodactylus, Page 95

http://bugguide.net/node/view/8041/bgimage

Genus: Macrodactylus Species: Macrodactylus subspinosus

Dichotomous Key: Scarab Beetles of South Carolina Key to Macrodactylus, Page 95

Species Description in The Scarabaeoid Beetles of Nebraska, Page 319

http://bugguide.net/node/view/192123/bgimage

Tribe Melolonthini Genus: Phyllophaga Species: Phyllophaga crenulata Similar to: Phyllophaga aemula Dichotomous Key: Scarab Beetles of South Carolina Key to Phyllophaga, Page 75

P. crenulata-Scarabaeoid Beetles of Nebraska, Key to Phyllophaga, Page 332

http://bugguide.net/node/view/445927/bgimage

*All photographs are linked to their copyright holder. 38

Table of Contents Genus: Phyllophaga Species: Phyllophaga hornii Similar to: Phyllophaga anxia, P. diffinis, P. drakii, P. foxii, P. hirsuta, P. latifrons, P. luctuosa, P. mican, P. subtonsa, P. sylvatica Dichotomous Key: Scarabaeoid Beetles of Nebraska, Key to Phyllophaga, Page 332

Scarab Beetles of South Carolina Key to Phyllophaga, Page 75 http://www- museum.unl.edu/research/entomology/Guide/S carabaeoidea/ Scarabaeidae/Melolonthinae/Melolonthinae-Generic- pages/Melolonthini/Phyllophaga/Phyllophaga-species/P- hornii/P-hornii.html

--Could not find species descriptions on P. barda & P. vilifrons--

Genus: Phyllophaga Species: Phyllophaga implicita (cited in literature) Similar to: Phyllophaga balia, P. ephilida, P. fraternal P. forsteri, P. fusca, P. futillis, P. hirticula, P. ilicis, P. inversa, P. marginalis, P. rugosa, Dichotomous Key: Scarabaeoid Beetles of Nebraska, Key to Phyllophaga, Page 332

Scarab Beetles of South Carolina Key to Phyllophaga, Page 75

http://bugguide.net/node/view/525168/bgimage

*All photographs are linked to their copyright holder. 39

Table of Contents Genus: Phyllophaga Species: Phyllophaga tristis Similar to: Phyllophaga glaberrima (cited in the literature) P. gracilis, P. prununculina (cited in literature), P. quercus Dichotomous Key: Scarab Beetles of South Carolina Key to Phyllophaga, Page 75

http://bugguide.net/node/view/537952/bgimage http://bugguide.net/node/view/279854/bgimage

Genus: Polyphylla

Species: Polyphylla variolosa

Dichotomous Key: Genus Description in The Scarabaeoid Beetles of Nebraska, Page 385

Genus Description in the Scarab Beetles Of South Carolina, Page 92

http://bugguide.net/node/view/195786/bgimage http://scarabaeoidea.lifedesks.org/pages/1123

*All photographs are linked to their copyright holder. 40

Table of Contents Tribe Sericini Genus: Maladera Species: Maladera castanea Dichotomous Key: Species Description in the Scarab Beetles Of South Carolina, Page 72

http://bugguide.net/node/view/302886/bgimage

Genus: Nipponoserica (Serica) Species: Nipponoserica peregrine

For species information visit the link below

http://www- museum.unl.edu/research/entomology/Guide/Scarabaeoide a/Scarabaeidae/Melolonthinae/Melolonthinae-Generic- pages/Sericini/Nipponoserica/Nipponoserica.html

Genus: Serica Species: Serica intermixta Similar to: Serica georgiana, S. mystaca, S. parallela, S. vespertina Dichotomous Key: Scarab Beetles of South Carolina Key to Serica, Page 66

The Scarabaeoid Beetles of Nebraska Key to Serica, Page 390

http://bugguide.net/node/view/612175/bgimage

* All photographs are linked to their copyright holder. 41

Table of Contents Genus: Serica Species: Serica iricolor Dichotomous Key: Scarab Beetles of South Carolina Key to Serica, Page 66

http://bugguide.net/node/view/302886/bgimage

Genus: Serica Species: Serica sericea Similar to: Serica blatchleyi, S. loxia (cited in literature), S. opposita (cited in literature) Dichotomous Key: Scarab Beetles of South Carolina Key to Serica, Page 66

http://museum.unl.edu/research/entomology/Guide/Scaraba eoidea/Scarabaeidae/Melolonthinae/Melolonthinae-Generic- pages/Sericini/Serica/Serica.html

--Could not find species descriptions on S. atracapilla (atricapilla), S. floridana, & S. imitans--

Subfamily Rutelinae Tribe Anomalini Genus: Anomala Species: Anomala binotata Dichotomous Key: The Scarbaeoid Beetles of Nebraska Key to Anomala, Page 414

http://bugguide.net/node/view/393759/bgimage

* All photographs are linked to their copyright holder. 42

Table of Contents Genus: Anomala Species: Anomala flavipennis Similar to: Anomala marginata, A. semilivida Dichotomous Key: Scarab Beetles of South Carolina Key to Anomala, Page 98

A. flavipennis-The Scarabaeoid Beetles of Nebraska, Key to Anomala, Page 414

http://bugguide.net/node/view/416361/bgimage

Genus: Anomala Species: Anomala innuba Similar to: Anomala oblivia Dichotomous Key: Scarab Beetles of South Carolina Key to Anomala, Page 98

A. innuba-The Scarabaeoid Beetles of Nebraska, Key to Anomala, Page 414

http://bugguide.net/node/view/448799/bgimage

Genus: Anomala Species: Anomala lucicola Similar to: Anomala umbra Dichotomous Key: Scarab Beetles of South Carolina Key to Anomala, Page 98

http://bugguide.net/node/view/14141/bgimage

*All photographs are linked to their copyright holder. 43

Table of Contents Genus: Anomala Species: Anomala orientalis Dichotomous Key: Scarab Beetles of South Carolina Key to Anomala, Page 98

http://bugguide.net/node/view/290796/bgimage

Genus: Anomala Species: Anomala undulata

Dichotomous Key: Scarab Beetles of South Carolina Key to Anomala, Page 98

The Scarabaeoid Beetles of Nebraska Key to Anomala, Page 414

http://bugguide.net/node/view/183542/bgimage

Genus: Popillia Species: Popillia japonica

Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 427

Scarab Beetles of South Carolina, Page 103

http://bugguide.net/node/view/565927/bgimage

*All photographs are linked to their copyright holder. 44

Table of Contents Genus: Strigoderma Species: Strigoderma pygmaea Similar to: Strigoderma arbicola Dichotomous Key: Scarab Beetles of South Carolina Key to Strigoderma, Page 102

S. arbicola-Scarabaeoid Beetles of Nebraska, Page 429

http://bugguide.net/node/view/307068/bgimage

Tribe Rutelini Genus: Cotalpa Species: Cotalpa lanigera

Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Cotalpa, Page 405

Scarab Beetles of South Carolina Key to Tribe Rutelini, Page 103

http://bugguide.net/node/view/504037/bgimage

Genus: Parastasia Species: Parastasia brevipes Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Rutelini, Page 103

http://www.marylandbiodiversity.com/viewSpecies.php?spec ies=11318

*All photographs are linked to their copyright holder. 45

Table of Contents Genus: Pelidnota Species: Pelidnota punctata Dichotomous Key: Scarab Beetles of South Carolina Key to Tribe Rutelini, Page 103

The Scarabaeoid Beetles of Nebraska Species Description, Page 409

http://bugguide.net/node/view/8644/bgimage

Subfamily Scarabaeinae Tribe Ateuchini Genus: Ateuchus Species: Ateuchus histeroides Similar to: A. lecontei (cited in literature) Dichotomous Key: Scarab Beetles of South Carolina Key to Ateuchus, Page 16

A. histeroides-The Scarabaeoid Beetles of Nebraska, Page 268

http://bugguide.net/node/view/638383/bgimage

Tribe Canthonini Genus: Canthon

Species: Canthon vigilans Similar to: C. chalcites, C. pilularius, C. (Boreocanthon) probus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Canthon, Page 283

http://bugguide.net/node/view/183542/bgimage

*All photographs are linked to their copyright holder. 46

Table of Contents Genus: Canthon Species: Canthon viridis Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Canthon, Page 283

http://bugguide.net/node/view/626455/bgimage

Genus: Melanocanthon Species: Melanocanthon bispinatus Dichotomous Key: Species Description in Scarab Beetles of South Carolina, Page 15

http://bugguide.net/node/view/189719/bgimage

Tribe Coprini Genus: Copris Species: Copris minutis Similar to: C. fricator

Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Canthon, Page 283

http://bugguide.net/node/view/595299/bgimage http://bugguide.net/node/view/93153/bgimage

*All photographs are linked to their copyright holder. 47

Table of Contents Genus: Dichotomius Species: Dichotomius carolinus Dichotomous Key: Species Description in The Scarabaeoid Beetles of Nebraska, Page 274

Species Description in Scarab Beetles of South Carolina, Page 17

http://bugguide.net/node/view/281178

Tribe Onthophagini Genus: Onthophagus Species: Onthophagus concinnus

Dichotomous Key: Beetle Genus Onthophagus—Howden, Cartwright Key to Onthophagus, Page 10

http://bugguide.net/node/view/522918/bgimage

Genus: Onthophagus Species: Onthophagus hecate Similar to: O. pennsylvanicus, O. striatulus, O. taurus O. tuberculifrons

Dichotomous Key: Beetle Genus Onthophagus—Howden,

Cartwright Key to Onthophagus, Page 10

The Scarabaeoid Beetles of Nebraska Key to Onthophagus, Page 253 http://bugguide.net/node/view/353332/bgimage http://bugguide.net/node/view/353335/bgimage

*All photographs are linked to their copyright holder. 48

Table of Contents Genus: Onthophagus Species: Onthophagus nuchicornis Dichotomous Key: Beetle Genus Onthophagus—Howden, Cartwright Key to Onthophagus, Page 10

http://bugguide.net/node/view/564461/bgimage

Genus: Onthophagus Species: Onthophagus orpheus Dichotomous Key: Beetle Genus Onthophagus—Howden, Cartwright Key to Onthophagus, Page 10

The Scarabaeoid Beetles of Nebraska Key to Onthophagus, Page 253

http://bugguide.net/node/view/613227/bgimage

Tribe Phanaeini Genus: Phanaeus (Moved from Tribe Coprini) Species: Phanaeus vindex

Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Phanaeus, Page 276

http://scarabaeoidea.lifedesks.org/pages/1228

*All photographs are linked to their copyright holder. 49

Table of Contents TROGIDAE

Subfamily Troginae Genus: Omorgus (Trox) Species: Omorgus (Trox) monachus Similar to: Omorgus (Trox) asper, O. (Trox) tytus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Omorgus, Page 92

Scarab Beetles of South Carolina Key to Species of Troginae (Trogidae), Page 61

http://bugguide.net/node/view/169337/bgimage

Genus: Omorgus (Trox) Species: Omorgus (Trox) scabrosus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Omorgus, Page 92

http://bugguide.net/node/view/207686/bgimage

Genus: Omorgus (Trox) Species: Omorgus (Trox) suberosus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Omorgus, Page 92

http://bugguide.net/node/view/420122/bgimage

*All photographs are linked to their copyright holder. 50

Table of Contents Genus: Trox Species: Trox scaber Similar to: Trox aequalis, T. affinis, T. capillaris, T. hamatus, T. laticollis, T. sordidus, T. terrestris, T. tuberculatus Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Trox, Page 69

T. affinis & T. terrestris- Scarab Beetles Of South Carolina, Key to Troginae (Trogidae), Page 61

http://bugguide.net/node/view/168449/bgimage

Genus: Trox Species: Trox spinulosus Similar to: Trox foveicollis

Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Trox, Page 69

http://bugguide.net/node/view/351715/bgimage

*All photographs are linked to their copyright holder. 51

Table of Contents Genus: Trox Species: Trox striatus Similar to: Trox unistriatus

Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Trox, Page 69

http://bugguide.net/node/view/642925/bgimage

Genus: Trox Species: Trox variolatus

Dichotomous Key: The Scarabaeoid Beetles of Nebraska Key to Trox, Page 69

http://bugguide.net/node/view/176768/bgimage

*All photographs are linked to their copyright holder. 52

Table of Contents

References BugGuide. Iowa State University Entomology. http://bugguide.net/

Borowiec, L. “Psammodius sulcicollis (Illiger, 1801)”. Iconographia Coleopterorum Poloniae. http://coleoptera.ksib.pl/kfp/search.php?img=6954

Cartwright, O. L. and H. F. Howden. 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.

Dabry, Julien. “Acrossus rufipes”. Portraits d’éboueurs. Last Visited July 25, 2012. http://typhaeus.chez-alice.fr/images/Acrossus_rufipes.jpg

Gordon, R. D. and P. E. Skelley. 2007. A monograph of the Aphodiini inhabiting the United States and Canada (Coleoptera: Scarabaeidae: Aphodiinae). Memoirs of the American Entomological Institute, 79: 580 p.

Guzman, D. “Lucanus elaphus”. Flickr from Yahoo. Last Visited July 25, 2012. http://www.flickr.com/photos/entomopixel/5098352100/

Harpootlian, P. J. 2001. Scarab Beetles (Coleoptera: Scarabaeidae) of South Carolina. Biota of South Carolina. Volume 2. Clemson University Public Service Publishing.

Jameson, M.L. and B.C. Ratcliffe. “Generic Guide to New World Scarab Beetles”. University of Nebraska State Museum Division of Entomology. http://museum.unl.edu/research/entomology/Guide/Guide-introduction/Guideintro.html

Majka, Christopher. “Gnorimella maculosa (Knoch)”. Empty Mirrors Press. Last Visited July 25, 2012. http://www.chebucto.ns.ca/environment/NHR/Gnorimella_maculosa.html

Manci, Cosmin-Ovidiu. “Psammodius laevipennis (A. Costa, 1844—Aphodiinae)”. Forum Entomologi Italiani.

Murray, Tom. “Aphonus castaneus”. PBase. Last Visited July 25, 2012. http://www.pbase.com/tmurray74/image/125304612

*All photographs are linked to their copyright holder. 53

Table of Contents

Orozco, J. 2012. Monographic Revision of the American Genus Euphoria Burmeister, 1842 (Coleoptera: Scarabaeidae: Centoniinae). The Coleopterists Bulletin, 11: 1—182.

Picture of Pleurophorus atlanticus taken from http://unibio.unam.mx/irekani/bitstream/123456789/25622/1/9310.jpg Last Visited July 25, 2012

Picture of Teuchestes fossor taken from http://www.galerie- insecte.org/galerie/view.php?adr=image%2Fdos6%2Ftemp%2Fteuchestes_fosso.jpg Last Visited July 25, 2012.

Price, Dana L. “Phanaeus vindex (MacLeay)”. eScarab LifeDesks. Last Visited July 25, 2012. http://scarabaeoidea.lifedesks.org/pages/1228

Ratcliffe, B.C. and M.J. Paulsen. 2008. The Scarabaeoid Beetles of Nebraska (Coleoptera: Scarabaeoidea). University of Nebraska State Museum Bulletin 22: 1—570.

Shaughney, Jennifer. “Phyllophaga variolosa Hentz”. eScarab LifeDesks. Last Visited May 27, 2014. http://scarabaeoidea.lifedesks.org/pages/1123

Staines, C. L., “An Annotated Checklist of the Scarabaeoidea (Coleoptera) of Maryland”. 1984. Maryland Entomologist 2(4): 79—89.

*All photographs are linked to their copyright holder. 54